Jurassic crabs: new characters of carapace diagnostic to known tanidromitid species

Jurassic crabs: new characters of carapace diagnostic to known tanidromitid species

Natalia Starzyk

With 8 figures

Abstract: Representatives of Tanidromitidae are known from the Middle Jurassic (Bajocian) to the Late Jurassic (Kimmeridgian). The genus Tanidromites comprises now eight species. Four of them are present in large numbers in Polish localities. The differences between all species are based upon new features of the anterior part of the carapace, distinguishing them in a transparent way. Their development is described. Distinguishing T. etalloni from T. insignis is based on new material and new characters.

Key words: Tanidromites, Homolodromioidea, Brachyura, Decapoda, Jurassic, Oxfordian.

1. Introduction

The family Tanidromitidae is one of the oldest brachy-uran families. It comprises two genera: Tanidromites Schweitzer & Feldmann, 2008a and Gabriella col-linS et al., 2006. Representatives of the Tanidromi-tidae are known from the Middle Jurassic (Bajocian; FörSter 1985) to the Late Jurassic (Kimmeridgian; Schweitzer & Feldmann 2010a). The family was originally erected for the only genus Tanidromites, comprising now eight species: T. richardsoni wood-ward, 1907 – latest Bajocian (Schweitzer & Feld-mann 2010a), T. montreuilensis crônier & BourSicot, 2009 – Early Callovian (crônier & BourSicot 2009), T. lithuanicus Schweigert & KoppKa, 2011 – Middle Callovian (Schweigert & KoppKa 2011), T. insignis von meyer, 1857 – the stratigraphic range of which is Middle Oxfordian to latest Kimmeridgian (hyžný et al. 2011), T. etalloni collinS in collinS & wierz-BowSKi, 1985 – Late Oxfordian (collinS & wierz-BowSKi 1985), T. sculpta QuenStedt, 1857 – Late Jurassic (Schweitzer & Feldmann 2008a), T. schef-

fnerae Schweigert & KoppKa 2011 – Late Kimmer-idgian (Schweigert & KoppKa 2011) and T. pustulosa von meyer, 1840, from the Tithonian (Schweitzer & Feldmann 2010b).

The genus Gabriella described by collinS et al. (2006) was transferred to Tanidromitidae by Sch-weitzer & Feldmann (2008a) and was restricted to five species: G. plana van Straelen, 1936, G. anfrac-ta Schweitzer & Feldmann, 2009a, G. patula Sch-weitzer & Feldmann, 2009a, G. biburgensis wehner, 1988, and G. lugobaensis FörSter, 1985. All these species come from European localities (Schweitzer & Feldmann 2009a) except G. lugobaensis which is from Tanzania.

Diagnostic features of the genus Tanidromites are: ventral extension of the branchiocardiac groove initial-ly strong and then diminishing ventrally, and a large portion of the lateral flank ventral to the branchiocar-diac groove (Schweitzer & Feldmann 2009a). Species specific characters in Tanidromites include: shape and proportions of the carapace and its regions (collinS & wierzBowSKi 1985; von meyer 1857), shape and

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DOI: 10.1127/0077-7749/2013/0343 0077-7749/2013/0343 $ 4.00

N. Jb. Geol. Paläont. Abh. 269/2 (2013), 173–187 ArticleStuttgart, August 2013

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174 Natalia Starzyk

length of the augenrest; presence and size of the outer orbital spine (crônier & BourSicot 2009; Schweigert & KoppKa 2011) and the distance from the end of the augenrest to the cervical groove (Schweitzer & Feld-mann 2008a). Species also differ in the length of the rostrum and its tip which might be sharp or blunt.

Other characters, such as cervical pits, were men-tioned in the descriptions of a few species, but were not treated as diagnostic. This comes probably from the small number of specimens allotted to the tanidromit-id species. Besides T. insignis, the remaining four spe-cies treated herein were described based on single in-dividuals (T. lithuanicus, T. etalloni, T. richardsoni, T. scheffnerae).

I had access to a new large collection of brachy-urans housed in the Institute of Systematics and Evo-lution of Animals, Polish Academy of Sciences in Cracow which is stepwise being studied (Brachyura:

StarzyK et al. 2011; StarzyK et al. 2012; Anomura: Fraaije et al. 2012a-c). Among them the tanidromitid species were found in great numbers. This enabled me to find and verify new characters diagnostic for the species. This also allowed to confirm the position of T. etalloni as a separate species, and not a juvenile of T. insignis.

2. Material and methodsSpecimens treated herein come from the collection of the Museum of the Geological Institute, University of Warsza-wa (MWGUW), Staatliches Museum für Naturkunde in Stuttgart (SMNS), University of Greifswald (GG), and the Institute of Systematics and Evolution of Animals, Polish Academy of Sciences in Cracow (ISEZ PAN, I-F/MP).

Specimens housed in the ISEZ PAN were collected by a family of amateur collectors: iwona, roBert and Karolina BoreK. In total, there are about 680 specimens determined

Fig. 1. Studied area, the southern Polish Uplands, northwest of Cracow (Poland). Abbreviations of the localities: O – Ogrodzieniec, B – Bzów, G – Grabowa, W – Wysoka, N – Niegowonice.

Jurassic crabs: new characters of carapace diagnostic to known tanidromitid species 175

so far. For this study I have chosen 49 of the best preserved specimens.

In some specimens, the carapaces were coated with am-monium chloride prior to being photographed, to make the details better visible (according to Feldmann 1989).

3. Ages of localities

Specimens from the collection of the Geological In-stitute, University of Warszawa come from several lo-calities in the Polish Jura and Wieluń Upland. Speci-mens from the collection of the Staatliches Museum für Naturkunde in Stuttgart come from several Kim-meridgian localities in Germany, described previously (Schweigert & KoppKa 2011; Schweitzer & Feldma-nn 2008a).

The material described below originates from five localities in the southern Polish Uplands, northwest of Cracow (Poland), namely Ogrodzieniec, Bzów, Grabowa, Wysoka and Niegowonice (Fig. 1).Bzów – This pit, the northerly portion of which is in the village of Bzów, represents a prolongation of the large quarry at Ogrodzieniec. Ammonites indicate the Middle Oxfordian Transversarium Zone. A sin-gle specimen in this lot might represent Cardioceras tenuiserratum, which would indicate the upper Plica-tilis Zone (matyja & wierzBowSKi 1994).

Grabowa – The quarry is east of the quarry of Ne-gowonice and very close to it. The age of these sedi-ments was until now unknown from the literature. One accompanying ammonite was recently determined by E. głowniaK and A. wierzBowSKi (2007) as Taram-elliceras externnodosum, which correlates with the Late Oxfordian (uppermost Bifurcatus to lowermost Bimammatum zones, Hypselum Subzone).Niegowonice – The sequence exposed at this quarry ranges from the upper Elisabethae Subzone to the up-per Wartae Subzone – Middle and Late Oxfordian age (głowniaK 2006, StarzyK et al. 2011).Ogrodzieniec – On ammonite evidence, the section exposed at Ogrodzieniec Quarry can be dated as Early and Middle Oxfordian. With the exception of the dis-continuous Mariae Zone, all zones and subzones from the Cordatum to the Transversarium have been docu-mented here (głowniaK 2006; StarzyK et al. 2011).

Wysoka – Ammonites indicate the age of these sediments ranging from Early Oxfordian Cordatum Zone (Cordatum Subzone) to Middle Oxfordian Trans-versarium Zone (Buckmani Subzone) (przyBylSKi et al. 2010).

Localities from other regions of Poland were treat-ed previously. The Opoczno locality is in the Polish Jura, and it is dated as Middle Oxfordian. Raciszyn

Fig. 2. Groundplan of the morphological structures in the species of Tanidromites on the example of T. sculpta.

176 Natalia Starzyk

and Draby are located in the Wieluń Upland. These are sediments of the Late Oxfordian Bifurcatus Zone (oSiKa et al. 1972).

4. New diagnostic features

The following four species-dignostic features were recognized among the five species of Tanidromites. Additionally, these characters were checked on pic-tures of two other species: T. montreuilense (crônier & BourSicot 2009: 1282, pl. 1, figs. 1-8, text-fig. 3A-D) and T. richardsoni (Schweigert & KoppKa 2011: 5, fig. 5A-B; Schweitzer & Feldmann 2008a, pl. 6A).

1) Presence of a pair of elongated depressions (shallow grooves) in the intersection of the anterior margin of the augenrest and the carapace margin, named here the anterior grooves (ag; Fig. 2). They are conspicuous in T. insignis and T. montreuilensis, weakly developed in T. lithuanicia, T. sculpta and T. richardsoni, and ab-sent in T. etalloni and T. scheffnerae (Fig. 3).2) Presence and size of a pair of pits positioned sym-metrically in the middle of the hepatic regions. They were mentioned before (Schweigert & KoppKa 2011) but noted only in three species (T. insignis, T. lithu-anica, T. sculpta). They are named here the hepatic pits (hp; Fig. 2). They are largest in T. insignis, T.

lithuanicus and a little smaller in T. richardsoni, T. montreuilensis, T. sculpta. Tanidromites etalloni and T. scheffnerae do not have them at all (Fig. 3).3) Presence and appearance of two elongated tubercles in the hepatic grooves, one on each side of the meso-gastric region, named here the hepatic tubercles (ht; Fig. 2). These tubercles are very distinct and large in T. scheffnerae, T. insignis and T. richardsoni, and more shallow and hence poorly visible in T. sculpta and T. montreuilense. In T. etalloni and T. lithuanicus they are absent (Fig. 3).4) Shape of a pair of cervical pits positioned in the middle of the cervical groove (cp; Fig. 2). They were observed before (Feldmann et al. 2006; Schweitzer & Feldmann 2009a) but their shape was not treated as a diagnostic feature. The cervical pits can be oval (T. etalloni), rounded (T. lithuanica), or crescent (T. in-signis, T. sculpta, T. scheffnerae) (Fig. 3). Just as the hepatic tubercles, cervical tubercles are clearly visible with the cuticle present and absent (Figs. 4B-C, 6A, C, F, 7C, E, 8E).5) Additionally, the measurements indicated in Fig. 3 were applied and found to be helpful for diagnostic purposes. These are: RtC – length from the rostrum to the cervical groove and AtC – length from the end of the outer augenrest angle (outer orbital spine if pres-ent) to the cervical groove.

Fig. 3. Reconstruction of the anterior part of the carapace and the augenrest of described species of Tanidromites. The area of augenrest is shaded.


5. Systematic palaeontology

Order Decapoda latreille, 1802Infraorder Brachyura linnaeuS, 1758Section Dromiacea de haan, 1833

Superfamily Homolodromioidea alcocK, 1900Family Tanidromitidae Schweitzer & Feldmann,

2008a

Genus Tanidromites Schweitzer & Feldmann, 2008a

Type species: Prosopon insigne v. meyer, 1857, by original designation.

Other species included: T. etalloni (collinS in collinS & wierzBowSKi, 1985), as Coelopus; T. lithuanica Schweigert & KoppKa, 2011; T. montreuilensis crônier & BourSicot, 2009; T. pustulosa (v. meyer, 1840), as Pithonoton; T. rich-ardsoni (woodward, 1907), as Prosopon; T. scheff-nerae Schweigert & KoppKa, 2011; T. sculpta (QuenStedt, 1857), as Prosopon.

The stratigraphic range of the genus is latest Bajocian to latest Kimmeridgian.

Fig. 4. Tanidromites insignis (v. meyer, 1857). A – Specimen, 6245, Wysoka; B – specimen, 6250, Wysoka; C – specimen, 4561, Niegowonice; D – rostrum, 6243, Wysoka; E – augenrest, anterior view, 6244, Wysoka; F – specimen, 6256, Wysoka; G – augenrest, anterior view, 6273, Wysoka; H - lateral view, IGPUW C/1/14, Opoczno. Scale bars equal 1 mm.

178 Natalia Starzyk

Tanidromites insignis v. meyer, 1857Figs. 3-4

1857 Prosopon insigne von meyer, p. 556.1858 Prosopon insigne (von meyer). – v. meyer, p. 61.1860 Prosopon insigne (von meyer). – v. meyer, p. 193,

pl. 23, fig 4.1925 Avihomola insignis (von meyer). – van Straelen,

p. 344.1925 Prosopon insigne (von meyer). – Beurlen, p. 468.1928 Pithonoton insigne (von meyer). – Beurlen, p.149.1929 Pithonoton insigne (von meyer). – glaeSSner, p.

320.1933 Pithonoton insigne (von meyer). – glaeSSner, p.

181.1985 Pithonoton insigne (von meyer). – collinS in col-

linS & wierzBowSKi, p. 83, pl. 3, figs.1-2.1988 Pithonoton insigne (von meyer). – wehner, p. 71,

pl. 5, figs. 1-3.2000 Pithonoton insigne (von meyer). – müller et al.,

fig.17K, L.2002 Pithonoton insigne (von meyer). – garaSSino &

KroBicKi, figs. 3a-b; 4.2008a Tanidromites insignis (von meyer). – Schweitzer &

Feldmann, p. 138, pl. 6, fig. A-F.2008 Pithonoton insigne (von meyer). – KroBicKi &

zatoń, p. 38.2010 Tanidromites insignis (von meyer). – Schweitzer et

al., p. 61.2011 Tanidromites insignis (von meyer). – Schweigert &

KoppKa, p. 7, pl. 6.2011 Tanidromites insignis (von meyer). – hyžný et al.,

p. 4, fig. 2A-E.

Emended diagnosis: Large-sized species (maximal cara-pace length 22.5 mm, width 15.6 mm). Carapace is convex transversally and longitudinally. The distance from outer or-bital spine to cervical groove (AtC) is long (0.26-0.38 x RtC length). Rostrum is short, bluntly terminated. Augenrest is flat and short (length about 0.8 x width), well visible, termi-nation with a short outer orbital spine. The anterior groove is present, conspicuous. Hepatic pits are well-defined; the surface around it is not depressed. Hepatic tubercles are large and distinct. Cervical pits are of crescent shape. Car-diac region is well defined by the branchiocardiac groove.

Type material examined: Holotype: SMNS 61641; Braunenberg near Aalen-Wasseralfingen (south-west Ger-many), Early Kimmeridgian.

Comparative material: Germany: SMNS 61662, Albstadt-Ebingen, Lower Kimmeridgian, Pseudomutabilis Zone; SMNS 61640, Sauserbrunnen near Laufen/Eyach, Lower Kimmeridgian; SMNS 61666, Nusplingen, Lower Kimmer-idgian. Poland: MWG UW C/1/013, MWG UW C/1/014, both Opoczno (Polish Jura).

Additional material: Specimens from Bzów: I-F/MP/2846/1530/08. From Grabowa: I-F/MP/5611/1543/09. From Niegowonice: I-F/MP/199/1495/08; I-F/MP/4561/

1534/08; I-F/MP/4569/1534/08; I-F/MP/5072/1543/09; I-F/MP/5325/1543/09; I-F/MP/6246/1588/11; I-F/MP/6248/1588/11; I-F/MP/6253/1588/11. From Ogrodzieniec: I-F/MP/5/1489/08; I-F/MP/118/1489/08; I-F/MP/6269/1599/12. Specimens from Wysoka: I-F/MP/3393/1532/08; I-F/MP/4400/1532/08; I-F/MP/4646/1534/08; I-F/MP/6243/1588/11; I-F/MP/6244/1588/11; I-F7MP/6245/1588/11; I-F/MP/6250/1588/11; I-F/MP/6256/1588/11; I-F/MP/6273/1599/12 (further on only the numbers in bold will be referred to).

Dimensions: Carapace width ranges from 5.62 to 15.6 mm; length of the largest specimen is 22.5 mm.

Additional description: The carapace is about 1.5 x longer than wide, convex (Fig 4A, H), widest across the epibran-chial region. Lateral borders are parallel, smooth, except the outer orbital spine. The distance from outer orbital spine to cervical groove (AtC) is long (0.26-0.38 x RtC length) (Figs. 3, 4A-C, F).

Rostrum is downturned, not long, narrowing anteriorly, with blunt termination. The axial groove is deep and reach-es almost to the anterior border of rostrum. Rostrum is best preserved on specimens 6243, 6244 and 6245 (Fig. 4D).

Augenrest is flat and short (length about 0.8 x width), with clearly visible border, and ending with short outer or-bital spine (in contrast to T. etalloni, in which this spine is absent). The anterior groove is along the intersection of the anterior margin of the augenrest and the carapace margin (Fig. 4A, D, F). Details of the augenrest are best visible in specimens 4646, 4836, 6243, 6244, 6246 and 6273 (Fig. 4E, G).

Epigastric regions are elongated. Mesogastric region is pyriform, distinctly delimited by grooves in its anterior and posterior part, and very weakly in the middle. In the middle of its posterior border there is a small incision, on both sides of which there are strongly developed muscle scars in the shape of a pair of symmetrical, horizontal elevations ex-tending across the entire posterior border of the mesogastric region. They are elongated and correspond to attachments of the gastric muscles. In front of them there is a pair of similar, smaller scars (Fig. 4F).

Hepatic pits are well defined; the surface around is not depressed (Figs. 3, 4B-C). Hepatic tubercles are large and distinct (Fig. 4A-C). Cervical pits are crescent (Figs. 3, 4A-C, F).

The postcervical groove ends with depressions on both sides. Together with the branchiocardiac groove they form narrow areas facing the cardiac region. The tubercle on the urogastric region is very small, positioned in the middle of the postcervical groove. Cardiac region is well delimited by the branchiocardiac groove, and is provided with two tu-bercles in the anterior part and one in the posterior part.

Epibranchial region is more convex and a little shorter than the branchial region.

The cervical groove is deep. The middle part of the branchiocardiac groove is weaker than the lateral parts. It reaches the posterior end of the carapace, separating it into halves. Posterior border of the carapace is usually badly pre-served, incised in the middle.


Tanidromites etalloni (collinS in collinS & wierz-BowSKi, 1985)

Figs. 3, 5

1985 Coelopus etalloni collinS in collinS & wierz-BowSKi, p. 81, pl. 2, fig. 1a, 1b.

1988 Pithonoton insigne (collinS in collinS & wierz-BowSKi). – wehner, p. 71.

2008a Tanidromites etalloni (collinS in collinS & wierz-BowSKi). – Schweitzer & Feldmann, p. 140, pl. 6, fig. G-H.

Emended diagnosis: Small-sized species (maximal cara-pace width 8.6 mm). Carapace is strongly convex transver-sally and longitudinally. Lateral borders are parallel, lack-ing outer orbital spine. The distance from the posterior end of the augenrest to cervical groove (AtC) is long (0.28-0.32

x RtC length). Rostrum is short as in T. insignis but with sharp termination. Augenrest is flat and sharply ended; its length is about 0.95 x width. Mesogastric region is triangu-lar shaped. A pair of oval shaped cervical pits is positioned in the cervical groove on both sides of the incision on the mesogastric region. Hepatic pits and hepatic tubercles are absent.

Type material: Holotype: MWG UW C/1/010, Raciszyn (Wieluń Uppland, Poland), Early Kimmeridgian (Planula Zone).

Additional material: Specimens from Bzów: I-F/MP/1740/1517/08; I-F/MP/1905/1517/08; I-F/MP/2033/1517/08; I-F/MP/3585/1533/08. From Niegowonice: I-F/MP/6254/1588/11 (further on only the numbers in bold will be re-ferred to).

Fig. 5. Tanidromites etalloni (collinS in collinS & wierzBowSKi, 1985). A – Holotype, MWG UW C/1/10, Raciszyn); B – specimen, 2033, Bzów; C – specimen, 3585, Bzów; D – augenrest, anterior view, IGPUW C/1/10; E – augenrest, anterior view, 3585; F – lateral view, IGPUW C/1/10. Scale bars equal 1 mm.

180 Natalia Starzyk

Dimensions: Carapace width ranges from 3.74 to 8.6 mm. The length of the holotype, which is the biggest specimen, is more than 11 mm.

Additional description: The carapace is longer than wide, strongly convex transversally and longitudinally (Fig. 5A, C), widest across the epibranchial region. Lateral borders are parallel, smooth, without the outer orbital spine. The distance from the posterior end of the augenrest to cervical groove (AtC) is very long (0.28-0.32 x RtC length) (Figs. 3, 5A, C).

Rostrum is short as in T. insignis but with sharp end, downturned. The axial groove in the middle is moderately deep and reaches the anterior border of rostrum.

Augenrest is flat and sharply ended (length is about 0.95x width), with clearly visible border. In contrast to T. insignis, there is no outer orbital spine right behind the dis-tal angle. The anterior groove is not visible. Details of the augenrest are best visible in specimens 1740 and 2033 (Fig. 5C-D).

Epigastric regions are elongated. Mesogastric region is triangular shaped. It is distinctly bordered by grooves in its anterior and posterior part, very weakly in the middle part. In the middle of its posterior border there is a small incision. Gastric muscle scars on both sides of this incision are very weakly developed. A pair of oval pits is lying in the cervical groove, on both sides of the incision (Figs. 3, 5A-B). Hepatic pits and hepatic tubercles are absent.

The tubercle on the urogastric region is very weakly vis-ible.

The postcervical groove is faint. The areas facing the cardiac region, formed together with the branchiocardiac groove and corresponding to attachments of the dorsoven-tral muscles, are almost not visible.

Borders of the cardiac region are weakly distinguished. There are two tubercles lying in the anterior part of this re-gion and one very small in the posterior part.

Epibranchial region is convex and the branchial region is more flat. Branchial region is not much longer than the epibranchial region. The posterior border of the carapace is not fully preserved in any specimen.

The cervical groove is deeper than the branchiocardiac groove. The middle part of the branchiocardiac groove is weaker than the lateral parts and does not reach the poste-rior end of the carapace.

Remarks: Since its description (collinS & wierzBowSKi 1985), Tanidromites etalloni was only known from the ho-lotype, and for this reason it had been treated as a possible juvenile form of T. insignis (wehner 1988; Schweigert & KoppKa 2011).

Material from the collection of the ISEZ PAN provide the evidence that both species are distinct. The following features distinguish T. etalloni from T. insignis: absence of outer orbital spine (which is distinct in T. insignis). Meso-gastric region is different in shape: triangular in T. etalloni and without clearly differentiated narrow and wide parts (Figs. 3, 5A-B), which are well visible in T. insignis (Figs. 3, 4A-C, F). Rostrum of T. etalloni is sharply ended, while ros-trum of T. insignis has a blunt end. Hepatic pits and hepatic tubercles are absent in T. etalloni, but present in T. insignis.

The possibility that the hepatic pits and tubercles are

invisible in the holotype of T. etalloni because of its small size, must be rejected, as they are well visible even in small specimens of T. insignis (of the size comparable to T. etal-loni).

Tanidromites scheffnerae Schweigert & KoppKa, 2011

Figs. 3, 6

2011 Tanidromites scheffnerae Schweigert & KoppKa, p. 10, fig. 8.

Emended diagnosis: Large-sized species (maximal cara-pace length 27.34 mm). Carapace is more convex transver-sally and longitudinally than in T. insignis and T. etalloni. The distance from outer orbital spine to cervical groove (AtC) is very long (0.32-0.35 x RtC length). Rostrum is quite long with sharp termination. Augenrest is flat and very short (length about 0.6 x width), clearly visible, rounded. The outer orbital spine is absent. The anterior groove is ab-sent. Hepatic pits are absent. Hepatic tubercles are large and distinct. Cervical pits are of crescent shape.

Type material examined: Holotype: SMNS 67145, Gun-delsheim (Germany), Late Kimmeridgian (Pseudomutabilis Zone).

Additional material: Specimens from Bzów: I-F/MP/5150/1543/09. From Niegowonice: I-F/MP/4676/1534/08; I-F/MP/6236/1588/11; I-F/MP/6272/1599/12. From Wysoka: I-F/MP/6261/1588/11.

Dimensions: The length of the biggest specimen is 27.34 mm.

Additional description: The carapace is longer than wide, strongly convex (Fig. 6C-D), widest across the epibranchial region. Lateral borders are parallel, smooth, lacking outer orbital spine. The distance from outer orbital spine to cer-vical groove (AtC) is relatively very long (0.32-35 x RtC length).

Rostrum is short and downturned, with a sharp termina-tion. The axial groove in the middle is shallow and does not reach the end of the rostrum (Figs. 3, 6A-C).

Augenrest is flat, very short and rounded (length is about 0.6x width), with clearly visible border. Outer orbital spine and the anterior groove are absent. Details of the augenrest are best visible in specimens 6236 and 6261 (Figs. 3, 6G-H).

Epigastric regions are elongated. Mesogastric region is pyriform, distinctly bordered by grooves in its anterior and posterior part, very weakly in the middle part. In the middle of its posterior border there is a small incision. Scars cor-responding to attachments of the gastric muscles are elon-gated, strongly developed, positioned on both sides of this incision and extend across the entire posterior border of the mesogastric region (Fig. 6E-F). Proximally they are accom-panied by a parallel pair of smaller scars.

Hepatic pits are absent. Hepatic tubercles are large and distinct (Figs. 3, 6A-C, E-F, I). Cervical pits are of crescent


shape (Figs. 3, 6A, C, F).Postcervical groove with deeper depressions laterally.

Together with the branchiocardiac groove they form narrow areas facing the cardiac region.

Urogastric region with a very small tubercle in the mid-

dle of postcervical groove. Borders of the cardiac region are weakly distinguished. There are two tubercles lying in the anterior part of this region and one very small in the pos-terior part. Epibranchial region is more convex and a little shorter than the branchial region.

Fig. 6. Tanidromites scheffnerae Schweigert & KoppKa, 2011. A – specimen, 6236, Niegowonice; B – holotype, SMNS 67145, Gundelsheim, Germany; C – specimen, 6261, Wysoka; D – lateral view, 1222, Niegowonice); E, F – mesogastric region (E: 6272, Niegowonice, F: 6236); G, H – augenrest, anterior view (G: 6272, H: 6236); I – mesogastric region, 6236. Scale bars equal 1 mm.

182 Natalia Starzyk

The cervical groove is deep. The middle part of the branchiocardiac groove is weaker than the lateral parts and almost reaches the posterior end of the carapace. Posterior border of the carapace is mildly incised medially.

Remarks: T. scheffnerae was described from an Upper Kimmeridgian locality (Schweigert & KoppKa 2011). Spec-imens from the collection of the ISEZ PAN are older than the holotype; they come from Middle and Late Oxfordian localities.

Tanidromites sculpta (QuenStedt, 1857)Figs. 3, 7

1857 Prosopon sculptum QuenStedt, p. 778, pl. 95, fig. 48.

1858 Prosopon lingulatum von meyer, p. 61.1860 Prosopon lingulatum (von meyer). – v. meyer, p.

205, pl. 23, fig. 17.1925 Prosopon lingulatum (von meyer). – Beurlen, p.

476.1929 Pithonoton lingulatum (von meyer). – glaeSSner,

p. 322.1933 Prosopon lingulatum (von meyer). – glaeSSner, p.

180.1939 Prosopon würgauensis n. sp. – Kuhn, p. 492, pl. 15,

fig. 41.1988 Pithonoton lingulatum (von meyer). – wehner, p.

77, pl. 5, figs. 6-7.

Fig. 7. Tanidromites sculpta (QuenStedt, 1857). A – specimen, (248, Niegowonice); B – specimen, 735, Ogrodzieniec; C – cervical pits, 3414, Niegowonice; D – development of the hepatic pits and hepatic tubercles, 5532, Niegowonice; E – speci-men, MWG UW C/1/044, Draby; F – augenrest, anterior view, 5532; G – lateral view, 735, Ogrodzieniec. Scale bars equal 1 mm.


2008a Tanidromites lingulata (von meyer). – Schweitzer & Feldmann, p. 140, pl. 6, fig. I-J.

2010 Tanidromites sculpta (QuenStedt). – Schweitzer et al., p. 61.

2011 Tanidromites sculpta (QuenStedt). – Schweigert & KoppKa, p. 9, fig. 7.

Emended diagnosis: Small-sized species (maximal cara-pace length 12.85 mm, width – 10.97 mm). Lateral borders of the carapace are parallel or narrowing posteriorly. The carapace is flattened, widest in the hepatic region. The dis-tance from outer orbital spine to cervical groove (AtC) is very short (0.18-0.22 x RtC length). Rostrum is long with blunt end, axial groove deep. Augenrest is flat and relatively long (length about 0.95 x width), ended with well visible, blunt outer orbital spine. The anterior groove is weakly de-veloped. Hepatic pits are poorly visible, inside flat depres-sions. Hepatic tubercles are also faint. Cervical pits are of crescent shape.

Comparative material: MWG UW C/1/044, from Draby (Wieluń Upland).

Additional material: Specimens from Bzów: I-F/MP/1806/1517/08. From Grabowa: I-F/MP/5352/1543/09. From Niegowonice: I-F/MP/248/1495/08; I-F/MP/1084/1508/08; I-F/MP/1086/1508/08; I-F/MP/3051/1532/08; I-F/MP/3414/1532/08; I-F/MP/4533/1534/08; I-F/MP/4904/1534/08; I-F/MP/5012/1543/09; I-F/MP/5013/1543/09; I-F/MP/5066/1543/09; I-F/MP/5103/1543/09; I-/MP/5114/1543/09; I-F/MP/5532/1543/09. From Ogrodzieniec: I-F/MP/628/1502/08; I-F/MP/735/1502/08 (further on only the numbers in bold will be referred to).

Dimensions: The length of the biggest specimen is 12.85 mm. Carapace width ranges from 1.66 to 10.97 mm.

Additional description: The carapace is 1.49 x longer than wide, flattened, with lateral borders parallel or narrowing posteriorly (Fig. 7A-B, G), widest across the hepatic region. Lateral borders are smooth, except for the blunt outer or-bital spine. The distance from outer orbital spine to cervical groove (AtC) is relatively very short (0.18-22 x RtC length) (Figs. 3, 7A-B, E).

Rostrum is long and downturned, with a blunt end. The axial groove is deep and reaches end of the rostrum (Figs. 3, 7A).

Augenrest is flat and relatively long (length is about 0.95 of the width), with a blunt outer orbital spine. Suborbital margin projects beyond the upper orbital margin in apical view. Details of the augenrest are best visible on specimens 248, 4533, 5066 and 5532 (Fig. 7F).

The anterior groove is weakly developed. Epigastric regions are elongate. Mesogastric region is

pyriform, distinctly bordered by grooves in its anterior and posterior part, very weakly in the middle. In the middle of its posterior border there is a small incision, on both sides of which there are strongly developed muscle scars in shape of a pair of symmetrical horizontal elevations extending across the entire posterior border of the mesogastric region. Proxi-mally they are accompanied by a parallel pair of poorly de-

veloped smaller scars.Hepatic pits are faint, the surface around them is slightly

depressed (Figs. 3, 7A-B, D-E). Hepatic tubercles are also faint (Figs. 3; 7A-B, D-E). Cervical pits are of crescent shape (Figs. 3, 7B-C, E).

Urogastric region with a very small tubercle in the middle of the postcervical groove. There are depressions on lateral sides of the postcervical groove. Together with the branchiocardiac groove they form narrow areas facing the cardiac region.

In the anterior part of the cardiac region there are two tubercles and one very small in the posterior part. Epibran-chial region is slightly convex but more than the branchial region. Branchial region is much longer than the epibran-chial region.

The cervical groove is deep. The middle part of the branchiocardiac groove is weaker than the lateral parts and almost reaches the posterior end of carapace. Posterior bor-der of the carapace is mildly incised medially.

Tanidromites lithuanica Schweigert & KoppKa, 2011

Figs. 3, 8

2011 Tanidromites lithuanicus [sic] Schweigert & Kopp-Ka, p. 4, fig. 4.

Emended diagnosis: The carapace is 1.6 x longer than wide, with convex regions and smooth lateral borders nar-rowing posteriorly. The distance from outer orbital spine to cervical groove (AtC) is rather short (0.28 x RtC length). Augenrest is more concave than in other species, quite short (length about 0.6x width), with a blunt outer orbital spine. The anterior groove is distinct. Mesogastric region is more distinctly bordered by grooves than in other species. Gastric muscle scars along posterior border of this region are not visible. Hepatic pits are distinctly visible. Hepatic tubercles are absent. Cervical pits are round. Cardiac region is better defined by the branchiocardiac groove than in other species of Tanidromites.

Type material: Holotype: GG-350-2; type locality: Papartinė near Papilė (Popilani), Lithuania; age: Middle Callovian (Coronatum Zone, Grossouvrei Subzone). The holotype was not examined by myself, but the diagnosis and description below are based on the excellent photographs provided by jenS KoppKa.

Dimensions: The length of the holotype is 14.71 mm, width 9.38 mm.

Additional description of holotype: The carapace is 1.6 x longer than wide, with convex regions and smooth lateral borders narrowing posteriorly (Fig. 8A, C), widest across the epibranchial region. Lateral borders are smooth, with a small outer orbital spine. The distance from outer orbital spine to cervical groove (AtC) is short (0.28 x RtC length) (Figs. 3, 8A).

184 Natalia Starzyk

Rostrum is quite long and downturned, with damaged tip. The axial groove along rostrum is deep (Fig. 8B).

Augenrest is more concave than in T. insignis, T. scheff-nerae, T. etalloni and T. sculpta described above, quite short (length is about 0.6x width), with a blunt outer orbital spine (Fig. 8D). Sub- and upper orbital margin project weakly be-yond the surface of the augenrest. The anterior groove is distinct (Figs. 3, 8B-C).

Epigastric regions are rounded. Mesogastric region is pyriform, more distinctly bordered by grooves than in other species. In the middle of its posterior border there is a small incision. Gastric muscle scars on both sides of this incision are not visible.

Hepatic tubercles are absent. Hepatic pits are distinctly visible (Fig. 8A-C). Cervical pits are round (Fig. 8A, E).

Urogastric region without tubercle. The deep lateral de-pressions of the postcervical groove are not fused with the branchiocardiac groove.

The anterior part of cardiac region is very convex, not divided into two tubercles. One small tubercle is in the pos-terior part.

Epibranchial region is more convex and shorter than the branchial region.

Grooves are very deep. The middle part of the branchio-cardiac groove is weaker than the lateral parts but also well visible; it reaches the posterior end of the carapace. Poste-rior border of the carapace is incised medially.

6. Discussion

6.1. Importance of new diagnostic characters

Characters described above proved to be reliable with-in the species of the genus Tanidromites. Their pres-ence and appearance is stable; the paired structures are distributed symmetrically which helps to discern them from casual artifacts of the matrix. This was confirmed on numerous specimens from four museum collections (ISEZ PAN, MWGUW, GG and SMNS) which host the conspecific specimens from various lo-calities. Therefore, it may be stated with confidence that these features really represent the characteristics of the species, and not of the local population.

An important fact is that within a given species they do not change with age of the crab and are found on every specimen regardless of its size. The best ex-ample is T. insignis, represented by dozens of speci-mens in a wide size range. Characters like hepatic pits are well visible even in the smallest specimens, while they are always absent in T. scheffnerae, in specimens of size comparable to the largest T. insignis. Hepatic tubercles characteristic of T. insignis and of T. scheff-

Fig. 8. Tanidromites lithuanica Schweigert & KoppKa, 2011. Holotype, GG-350-2, Papartinė near Papilė, Lithuania; A – specimen; B – rostrum; C – lateral view; D – augenrest, lateral view; E – cervical pits. Scale bars equal 1 mm.


nerae are visible in big and small specimens of both species. Specimens of T. etalloni differ from the smallest specimens of T. insignis by the absence of he-patic pits and tubercles, and it can be stated with confi-dence that both species are separate. Anterior grooves, which are rather subtle structures, are identified fairly easily in T. sculpta despite small size of this species, while they can never be found in large specimens of T.scheffnerae.

The combination of pits, tubercles and grooves characteristic to the species permits identification of even damaged and incomplete specimens, especially with the aid of the characteristic shape of the augenrest (the latter feature proved to be very helpful also within the genus Goniodromites; StarzyK et al. 2012).

6.2. The plausible role of the pits and grooves in the carapace

Deep relief of the carapace, as well as additional struc-tures such as grooves, sutures, pits and tubercles were characteristic to a majority of the first crabs. Good examples might be species of the genera Laeviproso-pon, Planoprosopon, Gabriella or Bucculentum (ex. Schweitzer & Feldmann 2008b, fig. 1; Schweitzer & Feldmann 2009a, figs. 1, 3; Schweitzer & Feld-mann 2009b, fig. 4; Schweitzer & Feldmann 2009c, fig. 3; StarzyK et al. 2011, figs. 3-4). By analogy with recent crabs the elements of carapace ornamentation are related to muscles attachments (Schweitzer & Feldmann 2009a; Schweigert & KoppKa 2011; ng & guinot 2008; Kato et al. 2010; crônier & BourSicot 2009). Thus, the scars on the posterior border of the mesogastric region and a pair of cervical pits just be-low in the cervical groove correspond to attachments of the gastric muscles, while hepatic pits correspond to attachments of the eye muscles. Therefore it is not surprising that they are characteristic for the species and are present in all growth stages. Probably their presence and appearance is just a mere trace of deep differences between the species in structures that are not preserved, such as antennae, chelae and eyes.

Acknowledgements

I am very grateful to jenS KoppKa for taking the photographs of the Tanidromites lithuanica holotype. Critical reviews by carrie Schweitzer and guenter Schweigert helped me to improve this paper. This project was supported by the grant No. N N303 811940 of the Polish Ministry of Science and education to n. StarzyK and e. KrzemińSKa.

References

alcocK, a. (1900): Materials for a carcinological fauna of India. No. 5. The Brachyura Primigenia, or Dromiacea. – Journal of the Asiatic Society of Bengal, 68 (2): 123-169.

Beurlen, K. (1925): Über Brachyuren- und Anomurenreste des Schwäbischen Jura. – Neues Jahrbuch für Mineralo-gie, Abteilung B, Beilage-Bände, 52: 464-532.

Beurlen, K. (1928): Die fossilen Dromiaceen und ihre Stammesgeschichte. – Paläontologische Zeitschrift, 10: 144-183.

collinS, j.S.h. & wierzBowSKi, a. (1985): Crabs from the Oxfordian sponge megafacies of Poland. – Acta Geo-logica Polonica, 35 (1-2): 73-88.

collinS, j.S.h., roSS, a.j., genzano, g. & manzian, h. (2006): Earleria gen. nov. and Gabriella gen. nov., re-placement names for Foersteria arai and BrinKmann-voSS, 1980 (Cnidaria, Hydrozoa, Mitrocomidae) and Foersteria wehner, 1988 (Crustacea, Decapoda, Pro-sopidae), junior homonyms of Foersteria Szepligeti, 1896 (Insecta, Hymenoptera, Braconidae). – Bulletins of the Mizunami Fossil Museum, 33: 125-126.

crônier, c. & BourSicot, p. y. (2009): A new decapod crustacean faunule from the Middle Jurassic of north-west France. – Palaeontology, 52 (6): 1275-1289.

Feldmann, r.m. (1989): Whitening fossils for photographic purposes. – The Paleontological Society, Special Publi-cations, 4: 342-346.

Feldmann, r.m., lazăr, i. & Schweitzer, c.e. (2006): New crabs (Decapoda: Brachyura: Prosopidae) from Jurassic (Oxfordian) sponge bioherms of Dobrogea, Ro-mania. – Bulletins of the Mizunami Fossil Museum, 33: 1-20.

FörSter, r. (1985): Frühe Anomuren and Brachyuren (De-capoda, Crustacea) aus dem mittleren Dogger. – Mittei-lungen der Bayerischen Staatssammlung für Paläontolo-gie und historische Geologie, 25: 45-60.

Fraaije, R.H.B., KrzemińSKi, W., van BaKel, B.W.M., KrzemińSKa, E. & jagt, J.W.M. (2012a): The earliest record of pylochelid hermit crabs from the Late Jurassic of southern Poland, with notes on paguroid carapace terminology. – Acta Palaeontologica Polonica, 57: 647-654.

Fraaije, R.H.B., KrzemińSKi, W., van BaKel, B.W.M., KrzemińSKa, E. & jagt, J.W.M. (2012b): The sixth ab-dominal tergites of paguroid anomurans – a newly rec-ognized crustacean macrofossil type. – Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 266: 115-122.

Fraaije, R.H.B., KrzemińSKi, W., van BaKel, B.W.M., KrzemińSKa, E. & jagt, J.W.M. (2012c): New Late Jurassic symmetrical hermit crabs from the south-ern Polish Uplands and early paguroid diversifica-tion. – Acta Palaeontologica Polonica, http://dx.doi.org/10.4202/ app. 2012.022.

garaSSino, a. & KroBicKi, m. (2002): Galicia marianae n. gen., n. sp. (Crustacea, Decapoda, Astacidea) from the Oxfordian (Upper Jurassic) of the southern Polish uplands. – Bulletins of the Mizunami Fossil Museum, 29: 51-59.

http://dx.doi.org/10.4202/

http://dx.doi.org/10.4202/

186 Natalia Starzyk

glaeSSner, m.F. (1929): Crustacea Decapoda. – In: pom-pecKj, J.F. (Ed.): Fossilium Catalogus: Animalia, pars 41: 1-464; Berlin (Junk).

glaeSSner, m. F. (1933): Die Krabben der Juraformation. – Centralblatt für Mineralogie, Geologie, und Paläonto-logie, Abteilung B, 3: 178-191.

głowniaK, e. (2006): Correlation of the zonal schemes at the Middle – Upper Oxfordian (Jurassic) boundary in the Submediterranean Province: Poland and Switzer-land. – Acta Geologica Polonica, 56: 33-50.

głowniaK, e. & wierzBowSKi, a. (2007): Taxonomical revision of the perisphinctid ammonites of the Upper Jurassic (Plicatilis to Planula Zones) described by józeF SiemiradzKi (1891) from the Kraków Upland. – Volu-mina Jurassica, 5: 27-137.

haan, w. de (1833-1850): Crustacea. In: de SieBold, p.F. (Ed.): Fauna Japonica sive Descriptio Animalium, quae in Itinere per Japoniam, Jussu et Auspiciis Superiorum, qui Summum in India Batava Imperium Tenent, Suscep-to, Annis 1823 – 1830 Collegit, Notis, Observationibus et Adumbrationibus Illustravit.– i-xvii, i-xxxi, ix-xvi, 243 pp.; Lugduni-Batavorum (J. Müller & Co.).

hyžný, m., Schlögl, j. & KroBicKi, m. (2011): Tanidromites insignis (von meyer, 1857) (Crustacea: Decapoda: Brachyura) from Late Jurassic non-biohermal facies of the Western Tethys (Pieniny Klippen Belt, Western Car-pathians, Slovakia). – Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 262 (2): 213-226.

KaraSawa, h., Schweitzer, c.e. & Feldmann r.m. (2011): Phylogenetic analysis and revised classification of po-dotrematous Brachyura (Decapoda) including extinct and extant families. – Journal of Crustacean Biology, 31 (3): 523-565.

Kato, h., taKahaShi, t. & taira, m. (2010): Late Jurassic Decapod Crustaceans from Northeast Japan. – Paleon-tology, 53 (4): 761-770.

KroBicKi, m. & zatoń, m. (2008): Middle and Late Jurassic roots of brachyuran crabs: Palaeoenvironmental distri-bution during their early evolution. – Palaeogeography, Palaeoclimatology, Palaeoecology, 263: 30-43.

Kuhn, o. (1939): Beiträge zur Fauna des Oxford und Kim-meridge in Nordbayern. – Neues Jahrbuch für Minera-logie, Geologie und Paläontologie, Beilage-Bände, (B), 80: 464-497.

latreille, p.a. (1802-1803): Histoire naturelle, générale et particuliére, des crustacés et des insects, 3: 1-468, Paris (Dufart).

linnaeuS, c. v. (1758): Systema naturae per regna tria natu-rae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis (10th edi-tion). Volume 1. – 824 pp., Stockholm (Laurentii Slavii).

matyja, B.a. (1977): The Oxfordian in the south-western margin of the Holly Cross Mts. – Acta Geologica Po-lonica, 27 (1): 41-64.

matyja, B.A. & wierzBowSKi, a. (1994): On correlation of Submeditcrranean and Boreal ammonite zonations of the Middle and Upper Oxfordian: new data from Cen-tral Poland. – Geobios, 17: 351-358.

meyer, h. v. (1840): Neue Gattungen fossiler Krebse aus Gebilden vom bunten Sandsteine bis in die Kreide. – 23 pp.; Stuttgart (Schweizerbart).

meyer, h. v. (1857): Briefliche Mittheilungen. – Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Pe-trefaktenkunde, 1857: 556.

meyer, h. v. (1858): Briefliche Mitteilungen. – Jahrbuch für Mineralogie, Geologie, und Paläontologie, 1858: 61.

meyer, h. v. (1860): Die Prosoponiden oder die Familie der Maskenkrebse. – Palaeontographica, 7: 183-222.

müller, p., KroBicKi, m. & wehner, g. (2000): Jurassic and Cretaceous primitive crabs of the family Prosopidae (Decapoda: Brachyura) – their taxonomy, ecology and biogeography. – Annales Societalis Poloniae, 70: 49-79.

ng, p.K.l., guinot, d. & davie, p.j.F. (2008): Systema Brachyurorum: Part I. An annotated checklist of extant brachyuran crabs of the world. – The Raffles Bulletin of Zoology, 17: 1-286.

oSiKa, r., pożarySKi, w., rühle, e. & znoSKo, j. (1972): Geological map of Poland without Cenozoic formations; Warszawa (Wydawnictwa Geologiczne).

przyBylSKi, p.a., głowniaK, e., ogg, j.g., ziółKowSKi, p., SidorczuK, m., gutowSKi, j. & lewandowSKi, m. (2010): Oxfordian magnetostratigraphy of Poland and its correlation to Sub-Mediterranean ammonite zones and marine magnetic anomalies. – Earth and Planetary Science Letters, 289: 417-432.

QuenStedt, F.a. (1856-1858): Der Jura. – 842 pp.; Tübingen (Laupp).

Schweigert, g. & KoppKa, j. (2011): Decapods (Crustacea: Brachyura) from the Jurassic of Germany and Lithua-nia, with descriptions of new species of Planoprosopon and Tanidromites. – Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 260 (2): 203-210.

Schweitzer, c.e. & Feldmann, r.m. (2008a): A new Clas-sification for some Jurassic Brachyura (Crustacea: De-capoda: Brachyura: Homolodromioidea): Families Go-niodromitidae Beurlen, 1932 and Taniodromitidae new family. – Senckenbergiana lethaea, 87 (2): 119-156.

Schweitzer, c.e. & Feldmann, r.m. (2008b): Revision of the genus Laeviprosopon glaeSSner, 1933 (Decapoda: Brachyura: Homolodromioidea: Prosopidae) including two new species. – Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 253 (3): 273-285.

Schweitzer, c.e. & Feldmann, r.m. (2009a): Revision of Gabriella collinS et al., 2006 (Decapoda: Brachyura: Tanidromitidae) with new Jurassic species. – Neues Jahrbuch für Geologie und Paläontologie, Abhandlun-gen, 252 (1): 1-16.

Schweitzer, c.e. & Feldmann, r.m. (2009b): New species of Longodromitidae Schweitzer and Feldmann, 2009, from the Ernstbrunn Formation, Late Jurassic (Tithoni-an), Austria. – Annalen des Naturhistorischen Museums in Wien, 111A: 207-224.

Schweitzer, c.e. & Feldmann, r.m. (2009c): Revision of some Prosopinae sensu glaeSSner, 1969 (Crustacea: Decapoda: Brachyura) including four new families, four new genera, and five new species. – Annalen des Naturhistorischen Museums in Wien, 110 A: 55-121.

Schweitzer, c.e. & Feldmann, r.m. (2010a): The oldest Brachyura (Decapoda: Homolodromioidea: Glaess-neropsoidea) known to date (Jurassic). – Journal of Crustacean Biology, 30 (2): 251-256.

Schweitzer, c.e. & Feldmann, r.m. (2010b): The genus


Coelopus Étallon, 1861 (Brachyura: Glaessneropsoi-dea: Longodromitidae) with new species. – Neues Jahr-buch für Geologie und Paläontologie, Abhandlungen, 258 (1): 51–60.

Schweitzer, c.e., garaSSino, a., KaraSawa, h. & Schwei-gert, g. (2010): Systematic list of fossil decapod crus-tacean species. – Crustaceana Monographs, 10: 222 pp.; Leiden (Brill).

StarzyK, n., KrzemińSKa, e. & KrzemińSKi, w. (2011): In-traspecific variation in the Jurassic crab Bucculentum bucculentum (Decapoda: Homolodromioidea: Buccu-lentidae). – Neues Jahrbuch für Geologie und Paläon-tologie, Abhandlungen, 260 (2): 203-210.

StarzyK, n., KrzemińSKa, e. & KrzemińSKi, w. (2012): A new crab species from the Oxfordian of Poland (Deca-poda: Brachyura: Goniodromitidae). – Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 263 (2): 143-153.

van Straelen, v. (1925): Contribution à l’étude des crustecés décapodes de la période jurassique. – Mémoi-res d’Academie Royale de Belgique, Classe des Scienc-es, series 2, 7: 1-462.

van Straelen, v. (1936): Crustecés Décapodes nouveaux ou peu connus de l’époque Crétacique. – Musée Royal d’Historie Naturelle de Belgique, 12 (45): 1-50.

wehner, g. (1988): Über die Prosoponiden (Crustacea, Decapoda) des Jura. – Dissertation zur Erlangung des Doktorgrades der Fakultät für Geowissenschaften der Ludwig-Maximilians-Universität zu München. – 154 pp.; München (published by the author).

wierzBowSKi, a., głowniaK, e. & pietraS, K. (2010): Am-monites and ammonite stratigraphy of the Bimmama-tum Zone and lowermost Planula Zone (Submediterra-nean Upper Oxfordian) at Bobrowniki and Raciszyn in the Wieluń Upland, central Poland. – Volumina Jurassi-ca, 8: 49-102.

wierzBowSKi, a., matyja, B.a. & ŚluSarczyK-radwan, d. (1983): New data on Upper Jurassic strata in the Wieluń Upland and vicinities of Burzenin and their economic value. – Kwartalnik Geologiczny, 27 (3): 517-533.

woodward, h (1907): On a new brachyurous Crustacean from the ‘Clypeus-Grit’ (Inf. Oolite) of the Cotteswold Hills. – Geological Magazine, new series, 4: 79-81.

Manuscript received: May 8th, 2013.Revised version accepted by the Stuttgart editor: May 25th,

2013.

Address of the author:

natalia StarzyK, Institute of Systematics and Evolution of Animals, Polish Academy of Sciences; ul. Sławkowska 17; 31-016 Kraków, Poland;e-mail: [email protected]

Jurassic crabs: new characters of carapace diagnostic to known tanidromitid species

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