New records of Pennsylvanian trigonotarbid arachnids from West Bohemia, Czech Republic

2013·5.9 MB·English

by  Ivana Hradská

2013. The Journal ofArachnology 41:335-341
New records of Pennsylvanian trigonotarbid arachnids from West Bohemia, Czech Republic
Ivana Hradska' and Jason A. Dunlop-: 'West Bohemian Museum Pilsen, Department ofZoology, Kopeckeho sady 2, 301
16 Pilsen, Czech Republic; -Museum fiir Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at
the Humboldt University Berlin, Invalidenstrasse 43, D-10115 Berlin, Germany
Abstract. New recordsoftheextinctarachnidorderTrigonotarbidaaredescribed from UpperPennsylvanian(Moscovian:
Bolsovian [=Westphalian C])spoil heaps associated with theTynecminenearthevillageofTynecin West Bohemia, Czech
Republic.Threespecimensarerecorded,twoofwhichareincompleteopisthosomasassignedtoTrigonotarbidaincertaeseclis.
Athirdfossilismorecompleteandisdescribedhereas Tynecotarhusticluivekigen.etsp.nov. Itsfamilialpositionisuncertain,
but the presence ofa weakly lobed carapace and finely tuberculate body ornament suggests affinities with the ‘eophrynid
assemblage’ semu Dunlop & Brauckmann (2006) and particularly the family Lissomartidae from Mazon Creek, USA. In
order to be comprehensive in our study, we include a complete list ofCzech trigonotarbids.
Keywords: Fossil, Tynec, stratigraphy, new genus and species
Trigonotarbids are an extinct order of arachnids that to features such as ratios ofbody proportions, and we expect
ranged from the Upper Silurian (Pfidoli) (Jeram at al. 1990) that some of the named species will eventually prove to be
to the Lower Permian (Sakmarian) (Dunlop & Rossler 2013). synonyms. Historical descriptions ofBohemian trigonotarbids
Sixty“five valid species are currently recorded in the literature can be found in Stur (1877), Kusta (1883, 1884), Eric (1901,
(Dunlop et al. 2013), and they occur most frequently in 1904), Petrunkevitch (1953) and Pfibyl (1958), with more
Pennsylvanian sediments in Europe and North America. Here recent summaries in Oplustil (1985, 1986:fig. 1) and a revision
they seem to represent one ofthemorecommon and abundant ofthree genera by Dunlop (1995a).
arachnid groups in Coal Measures ecosystems, their fossils Here, we describe three Pennsylvanian trigonotarbids from
regularly turning up at appropriate localities. Trigonotarbids a new Bohemian locality; namely spoil heaps near the village
are placed in the arachnid taxon Pantetrapulmonata Shultz ofTynec. Two ofthese records are incomplete and are treated
2007 [see also Shear et al. (1987) for details of synapomor- as Trigonotarbida incertae sedis. A third fossil is much better
phies] as the sister-group of the orders Araneae (spiders), preserved and appears to represent a new genus and species,
Amblypygi (whip spiders), Thelyphonida (whip scorpions) and which we describe in detail below.
Schizomida (schizomids). They also share characters with the
METHODS
rare order Ricinulei (ricinluleids), which has led to speculation
that ricinuleids may also be part ofthis wider pantetrapulmo- The three specimens were obtained from the private
nate assemblage (see Dunlopet al. 2009). Trigonotarbid fossils collection of Mr. Frantisek Tichavek and have subsequently
are characterized by a segmented opisthosoma with eight or been deposited in the West Bohemian Museum, Pilsen. All are
nine dorsally visible tergites, most of which are divided preserved as compression fossils in a gray-brown mudstone.
longitudinally into median and lateral plates. These animals We whitened the fossils with ammonium chloride and studied
evidently had mouthparts modified for biting (e.g., Shearet al. them under incident light using a binocular microscope.
1987) and are generally regarded as probably having been Photographs were made using an Olympus E410 camera.
cursorial predators in Paleozoic terrestrial ecosystems (see Immersion in 70% alcohol also improved the detection of
overview in Garwood & Dunlop 2010). morphological details, particularly cuticle. We studied the
Most ofthe Pennsylvanian trigonotarbids have been found holotype of the new species under scanning electron micros-
at classic Westphalian localities associated with coal mining copy (SEM) using a JEOL JSM-6380LV at the Institute of
districts, such as the Saar and Ruhr areas of Germany Geology and Palaeontology, Charles University, Prague. All
(Guthorl 1934; Jux 1982), Silesia in Poland (Karsch 1882), measurements are in millimeters. —
former collieries or clay pits such as Coseley in the English Locality and geological setting. All three specimens de-
West Midlands associated with the British Middle Coal scribed here were found among spoil deposits of the Tynec
Measures (Pocock 1911; Petrunkevitch 1949) and Mazon (formerly Masaryk or Austria 2) mine near the village of
Creek in the USA (Petrunkevitch 1913). Trigonotarbids are Tynec in West Bohemia, Czech Republic. In this mine,
also well represented in the Coal Measures of central and bituminous coal of Pennsylvanian (Moscovian) age was
western Bohemia in the Czech Republic. Fifteen currently extracted between 1899 and 1965 from coal seams of the
valid species have been described so far from this area Radnice (Bolsovian substage) and Nyfany (Asturian substage)
(Table 1), although these are largely based on compression groups of the Kladno Formation; the former group being
fossils that are prone to post-mortem alteration; e.g., shearing, more important (Havlena 1964; Pesek 1996). This spoil heap
stretching, truncation of body parts, etc. A number of these was recently under threat ofdisplacement due to intensive fire
taxa are currently defined on rather trivial characters, relating clay extraction and subsequent reclamation. During this
process (between 2008 and 2010), a rich Pennsylvanian fiora
^Corresponding author. E-mail: [email protected] was collected in dark gray mudstones and examined in detail
335
— ——
) )
336 THE JOURNAL OF ARACHNOLOGY
Table 1. - Trigonotarbids from the Czech Republic; arranged stratigraphically from youngest (above) to oldest (below). Data based on
Oplustil (1985, 1986), Dunlop et al. (2013), Dunlop & Rbssler (2013) and the primary literature, updated to rellect recent changes in
nomenclature (cf. Dunlop 1995a; Harvey & Sclden 1995; Garwood & Dunlop 2011). Only valid species listed; see Dunlop et al. (2013) for
synonyms. Abbreviations: ISB Intra Sudetic Basin, PB - Pilsen Basin, KRB Kladno Rakovnik Basin, USB - Upper Silesian Basin.
Species Locality Age Ma
Aiilhraconuirttis nuhaniceiisis (Oplustil 1985) Radvanice, ISB Gzhelian 299-304
Aiuhnicoiiuirtiis hohcniicus (Eric 1901) Nyfany, PB Moscovian 311-315
[Asturian]
Anthrdcoiiuiriiis carcinoides (Eric 1901) Nyfany, PB Moscovian 311-315
[Asturian]
Antliraconuirtiis edegans Eric 1901 Nyfany, PB Moscovian 311-315
[Asturian]
Aiitliracoiuartiis uyrancnsis (Petrunkevitch 1953) Nyfany, PB Moscovian 311-315
[Asturian]
Nyranylarhiis liofiiianni (Eric 1901 Nyfany, PB Moscovian 311-315
[Asturian]
Nyrcinytarhiis longipes (Eric 1901 Nyfany, PB Moscovian 311-315
[Asturian]
Tynecotarhus tichaveki gen. et sp. nov. Tynec, PB Moscovian 309-310
[Bolsovian]
Aplumtonuirlus arcolatiis Pocock 1911 Libusin near Kladno, KRB Moscovian 309-310
(in Oplustil 1985) [Bolsovian]
Aphantoiuuniispiistiikitiis Scudder 1884 Libusin near Kladno, KRB Moscovian 309-310
(in Rbssler 1998) [Bolsovian]
Trigonoimirtiis spp. (in Oplustil 1985) Libusin near Kladno, KRB Moscovian 309-310
[Bolsovian]
Aphantomartidae gen et sp. Indet. Mine Pokrok near Radnice, KRB Moscovian 309-310
(in Oplustil 1985) [Bolsovian]
Anthracosironidae gen. et sp. Indet. Mine Gottwald-lll, Kladno, KRB Moscovian 309-310
(in Oplustil 1985) [Bolsovian]
AntliracoDKirlus jcincie (Oplustil 1986) Vinafice near Kladno, KRB Moscovian 309-310
[Bolsovian]
Aiithracoinarlus kustac Petrunkevitch 1953 Rakovnik, KRB Moscovian 309-310
[Bolsovian]
Aiilhracoiiiartiis minor Kusta 1884 Rakovnik, KRB Moscovian 309-310
[Bolsovian]
Planonuirtus kre/cii (Kusta 1883) Rakovnik, KRB Moscovian 309-310
[Bolsovian]
Pelroviciaproditoria Eric 1904 Petrovice near Rakovnik, KRB Moscovian 309-310
[Bolsovian]
Stcnotrogidus sainni (Stur 1877) Ostrava-Karvina, USB Serpukhovian 323-331
in order lo gather data about the diversity ofplant species in emended to Anthracomartida and Trigonotarbida. The fea-
the Kladno Formation. During the course ofthese paleolloral tures Petrunkevitch used to separate these taxa werechallenged
investigations, a total of 32 species of fossil plant were found by Dunlop (1996), and both groups were reunited under the
together with rare Pennsylvanian faitnal elements (Tichavek—& then more widespread and clearly defined name Trigonotar-
Bures 2010), including the lophophorate Microcoiuiiiis bida. Foracompletesynonymy listoftrigonotarbid highertaxa
originally thought to be the worm Spirorhis, but see Taylor & and further discussion, see Garwood & Dunlop (201 1).
Vinn (2006) for a reinterpretation of pre-Mesozoic records
and the trigonotarbid arachnids. Although no stratigraphically Family uncertain
indicative plant remains are associated with the arachnid
specimens, the character of mudstone resembles that from the Genus Tynecotavhus gen. nov.
—
roofofthe Upper RadniceCoal. Thiswould renderit Bolsovian Type species. Tynecotarhus (ic/uiveki gen. et sp. nov.
in age (ca. 309-310 Ma), etiuivalent to the Westphalian C of Etymology. From the type locality of Tynec in West
Western European stage terminologies. The followingAsturian Bohemia and the typical trigonotarbid suffix tarbus; derived
substage is thus ec]uivalent to the Westphalian D. from the Greek tarhos, meaning terror/alarm.
Diagnosis. Trigonotarbids characterized by a kidney-
SYSTEMATIC PAEEONTOLOGY shaped to subtriangular carapace bearing a raised, triangular
median region that hosts both the median eyes anteriorly and
Order Trigonotarbida Petrunkevitch 1949 a row of three tubercles more centrally; carapace also with
Remarks. Petrunkevitch (1949) divided Karsch’s (1882) faint lateral lobation. Opisthosoma oval, and entire dorsal
original order Anthracomarti into two orders; subsecjiiently body surface ornamented with fine, granular tuberculation.
HRADSKA & DUNLOP TRIGONOTARBIDS FROM BOHEMIA
Figure 3. Tyiiccolarhus liiiuivcki gen. el sp. nov. Holotype,
scanning electron micrograpli (SEM) of the central carapace region.
Note again the granular cuticle ornament and the three larger,
rounded tubercles towards the midline (arrowed). Seale bar = mm.
1
—
Remarks. Nine families of trigonotarbids arc currently
recognized, but there is no robust phylogenetic framework for
their interrelationships. Our new Bohemian fossil does not fit
comfortably into any of the accepted groupings. It was
provisionally listed (and figured) as a member of the family
Anthracosironidae by Tichavek & Bures (2010). Based on its
granular cuticle, it was also initially suspected to be an
anthracomartid; see Garwood & Dunlop (201 for a revision
1 )
of Anthracomartidae. However, the absence of lateral eye
tubercles on the carapace unequivoeally excludes the families
Anthracomartidae, Palaeocharinidae, Archaeomartidae (see
especially Poschmann & Dunlop 2()10:fig. 9), and probably the
usually long-bodied Anthracosironidae, too. The remaining
Figures I -2. Tynccolurbus ticliavckigen. et sp. nov.. a new genus five trigonotarbid families are potentially a monophyletic
and speeies belonging to the extinet araehnid order Trigonotarbida group, united by this character oflateral eyes absent. Ofthese,
from the Pennsylvanian (Moseovian, Bolsovian substage) ofTynee in
the Pilsen Basin ofWest Bohemia, Czech Republic. 1. Overviewofthe Trigonotarbidae have a similarly raised median carapace
holotype. West Bohemian Museum Pilsen. No. M()()758; 2. Thesame, region (cf. Pocock 1911; Petrunkevitch 1949), but their
detail of the carapace region showing the fine granular ornament of carapace is more obviously triangular and they lack both the
the cuticle arrowed are the putative median eyes, three larger granular body ornament seen in the new fossil and the faint
tubercleson themidlineand thefaint lateral divisionsofthecarapace. lobes dividing the lateral carapace margins.
Scale bars = 5 mm (1) and 2 mm (2). Three ofthe remaining four families also probably represent
a monophylum, namely Aphantomartidae, Kreischeriidae and
Eophrynidae. They were provisionally termed the 'eophrynid
assemblage’ by Dunlop & Brauckmann (2006) and can be
characterized by the potential synapomorphies of an often
338 THE JOURNAL OF ARACHNOLOGY
clypeus
—
Figure 4. Interpretativedrawingoftheholotype ofTynecotarbus tichavekigen. et sp. nov.; opisthosomal segmentsnumbered. Scalebar = 5
mm.
heavily ornamented dorsal body surface and deeply lobed Our new fossil preserves a unique combination ofcharacters
carapacemargins, typicallywith furtherlobation ofthemedian for trigonotarbids (see Diagnosis), which we believejustifies a
carapace region as well; see, e.g., illustrations in Petrunkevitch new genus. We suspect that with its granular ornament and
(1953, Figs—. 81, 82) and Dunlop (1995a). In this context, our weakly lobed carapace, it falls phylogenetically somewhere into
new fossil with only weak and barely discernible carapace this transitional zone between Trigonotarbidae and the eo-
lobation—cannot be accommodated into any of these ‘eo- phrynid assemblage. Two other genera appear to belong here, I'
phrynid’-like families sensu stricto. In a wider context, there too. Namaurotarbus Poschmann & Dunlop 2010 (family
seemtohavebeen transitional formsbetween therathersmooth uncertain) was raised for a trigonotarbid from Hagen Vorhalle
and simply-constructed Trigonotarbidae and the larger and in Germany, originally described by Dunlop & Brauckmann j
more robust and ornamented ‘eophrynid’-Hke trigonotarbids. (2006: Figs. 1, 2). This rather squat German fossil has a more
HRADSKA & DUNLOP—TRIGONOTARBIDS FROM BOHEMIA 339
—
Figure 5. Sketch reconstruction of the probable appearance of
Tynecotarbus tichavekigen. et sp. nov. in life; distal endsofpedipalps
and legs equivocal in the original fossil and omitted here.
triangular carapace and quite strongly defined lateral carapace
lobes. Unlike our new material, the median region of the
carapace is also divided down the middle, and there is no
granular cuticle ornament. We exclude our new fossil on these
characters from Namaurotarbus.
A much better candidate is Lissomartus Petrunkevitch 1949
from Mazon Creek, USA. This genus was originally placed in
Trigonotarbidae, but was raised to a new family, Lissomarti-
dae, by Dunlop (1995b), who also redescribed its two
constituent species. As in our new fossil, the carapace in i.im.'
Lissomartus is medially raised, has some larger tubercles wMlm
behind theeyes toward themiddleofthecarapace, and there is
weakly-defined lateral lobation of the carapace (Dunlop
1995b: Figs. 1^, 7). However, Lissomartus differs from the
—
new Bohemian arachnid in having a more triangular carapace Figures 6-7. Additional specimens assigned to Trigonotarbida
profile, in lackinggranularcuticle ornament, and in havingthe incertae sedis held in the West Bohemian Museum Pilsen. 6.
terminal (ninth) tergite divided into median and lateral plates. No M()0759; 7. No M00760. Scale bars = 2 mm.
Inthenewfossil, tergite9iscomposedofasingleplateonly. On
these characters, the new fossil does not fit clearly into
Lissomartidae as it is currently defined, although we suspect noted above, with its weakly lobed carapace and light body
this is where its affinities lie. Our concern would be that ornament it seemsto be partofa trigonotarbid lineagethat was
formally including it as a lissomartid cannot presently be approaching the condition ofthe eophrynid assemblage.
justified by explicit apomorphies and may render the family
paraphyletic with respect to more derived trigonotarbid taxa. Tynecotarbus tichaveki gen. et sp. nov.
Pendingadetailedphylogeneticanalysis,weprefernottocreate (Figs. 1-5)
another monotypic family group at this stage, and we suggest Trigonotarbida, Anthracosironidae: Tichavek & Bures
treatingthenewgenusasbeingofuncertainfamilyaffinities. As 2010:135-136, Figs. 6-7.
THE JOURNAL OF ARACHNOLOGY
340
—
Material. Holotypeandonlyknownspecimen,No. M00758, circular pygidium (diameter 1.5) visible and cuticle locally
Department ofPalaeontology, West Bohemian Museum Pilsen, preserved. Sparse granulation, unlike No. M00758 (the
Czech Republic (ex private collection ofF. Tichavek). holotype of Tynecotarhus tichaveki gen. et sp. nov.) and
—
Type locality and horizon. Tynec, West Bohemia, Czech No. M00759—, with tubercles oflarger diameter (ca. 0.5).
Republic. Pilsen Basin, most probably the roof of the Upper Remarks. The shape of the sternites and presence of a
Radnice Coal, Kladno Formation. Pennsylvanian, Moscov- pygidium on the underside of the opisthosoma allow both of
ian, Bolsovian substage (= Westphalian C). these fossils to be assigned with some confidence to
—
Diagnosis, As for the genus. Trigonotarbida. They are, however, too incomplete to place
—
Etymology. In honor of Mr. Frantisek Tichavek, who in any particular family or genus.
discovered the—holotype and kindly made it available for study. ACKNOWLEDGMENTS
carDaepsaccrei,ptioopn.isthSopseocmiamenanidn daorpsaarltivailewle(gF.igCsu.ti1c,l2e) roefveeanltiinrge We thank Frantisek Tichavek for making these specimens
specimen with fine granular ornament (Fig. 3); average available for our study, Stanislav Oplustil (Charles University,
tubercle size ca. 0.1. Total body length 20.8. Carapace Prague) for advice on Czech stratigraphy and the reviewers and
somewhat kidney-shaped to subtriangular, length 7.6, maxi- editorsforhelpfulcommentsonanearlierdraftofthetypescript.
mum width 8.7; slightly rounded anteriorly where it becomes LITERATURE CITED
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