GEOLOGICA BALCANICA, 34. 1-2, Sofia, Jun. 2004, p. 47-50

IU S UNES 0 IGCP Project 469

Atlas of animals from the Late Westphalian of Writhlington,

Ed. A. Jarzembowski

Maidstone Museum & Bent/if Art Gallery, St Faith 's St, Maidstone, Kent M£14 1LH and S.H.E.S., U11iversity of Reading, UK (E-mail: edjarzembowski@ maidstone.gov. uk)

3o. A. f!:JK:eM6oBcKu - AmJiac n030HeBecmrjjaJibCKUX Abstract. Representative animals from the Writhlington as­ :>ICUBOmflblX PaitmJiuflzmoHa (06?JeOuHeHHOe KopoJieBc­ semblages are collectively illustrated for the first time with mBo). IIpHse,neHhi rrepBhie o6o6l.lleHHhie HnniOcTpa~HH notes on palaeoecology, association and distribution. They rrpe,ncTaBHTenhHhiX )KJiBOTHhiX PaihnHHITOHCKHX acco~H­ are from a comparatively well sampled, terrestrial-fresh wa­ a~Hil:.lfnniOcTpa~~om COIIpOBO)K,LiaiOTCSI 3aMeTKaMH 06 HX ter fauna of nematodes, bivalves, arthropods and vertebrates rraneoe3KOnorHH H pa3Mel.lleHHH. ATnac H3fOTosneH Ha from the Upper Westphalian D of the UK. The Writhlington OCHOBaHHH cpaBHHTenhHO XOpOlliO OT06paHHhiX o6pa3- fauna provides a baseline for comparative faunal work in the ~0B CyX03eMHOH H rrpeCHOBO,LIHOH ayHa rrpe.nocTaBnSieT OCHOBHYIO HH¢lOpMa~HlO ,nng cpaBHHTenhHhlX

Jarzembowski, E.A., 2004. Atlas of animals from the Late Westphalian ofWrithlington, United King­ dom. -Geologica Bale., 34, 1-2;47-50. Key words: , Carboniferous, Faunas.

Introduction zone). The 'shales' are mostly from above the No. 10 Coal Seam although a small amount asso­ Writhlington Geological Nature Reserve ciated with the No. 1 seam is also found. The No. (W.G.N.R.) lies in the Radstock Syncline of the 10 seam was worked jointly with the former Foreland Basin at Lower Writhlington in the Kilmersdon Colliery nearby (Jarzembowski, Coalfield, UK (National Grid Refer­ 1989; Thomas and Cleal, 1994). ence ST 703 553; Latitude 51 o 17.8' North, Lon­ gitude 2° 55.6' West). W.G.N.R. is the former colliery tip of a closed deep mine and now a Site Body fossils (terrestrial fauna) of Special Scientific Interest (S.S.S .I.) and Re­ gionally Important Geological Site (R.I.G.S.) The terrestrial fauna is dominated by arthropods, Systematic collecting since 1984 has yielded a in particular insects (Pl. I, figs 4-6, 9, 11), arach­ well-preserved compression flora and fauna in­ nids (Fig. 1; Pl. 1, figs 1-2) and arthropleurids cluding over 1,200 insects and other animals (Fig. 2). These occur mainly in the No. 4 assem­ (Austen, 2001). The unburnt tip is composed blage of Proctor (1994), which is characterised by mainly of Roof Shales (grey silt- and mudstones, lycophyte debris. Arthropods are found less of­ occasionally sideritic) from the Farrington For­ ten in assemblages 5 and 6, which are dominated mation dating from the upper part of the West­ by sphenopsid and fern/pteridosperm remains re­ phalian D Stage (Dicksonites plueckenetii Sub- spectively. Nos 4 and 6 are the most common

47 PLATE I Fig. 1. Phalangiotarbus sp. (extinct arachnid) body alongside Cyperites bicarinatus leaf, BMB 014846 [W 79a] coil. B. Jarzembowski. Scale line in mm. Fig. 2. Pleophrynus verrucosa (armoured spider), opisthosoma showing upper and lower surfaces (after Jarzembowski, 1989). Scale line in mm. Species also known from Cantabrian/Westphalian D of Mazon Creek, South Wales Coalfield and possibly the Central Bohemian Region (Dunlop, 1994b). Fig. 3. Phagophytichnus sp. (trace attributable to a mandibulate insect) on left side of Macroneuropteris scheuchzeri leaf; conchoidal fracture on right side (after Jarzembowski, 1992). Scale line in mm. Fig. 4. Cockroach larva (nymph) distal portion of exuvium missing. BMB 014876 [W 105] coli. P. Austen. Scale line in mm. Fig. 5. Bechlya ericrobinsoni (damselfly-like dragonfly), piece of C. bicarinatus below (after Jarzembowski and Nel, 2002). Scaleinmm. Fig. 6. Blattodean insect (mylacrid cockroach). Upper view- head shield (pronotum) with folded forewings: note fern-like wing venation (after Jarzembowski & Ross, 1993). Length 17 mm. Fig. 7. Euproops danae (horseshoe crab). BMB 014850 coli. J. Latham. 45 x 25 mm. Species also known from Mazon Creek and the South Wales Coalfield (Anderson, 1994). Fig. 8. Adelophthalmus imhofi (water scorpion) body after Proctor ( 1999). Note cuticular sculpture. Scale line in mm. Spe­ cies widespread including Mazon Creek. Fig. 9. Protorthopteran (gerarid) insect, apical portion offorewing: note cross veins present. NHM In. 64605. Preserved wing length 32 mm, C. bicarinatus leaf below. Fig. 10. Palaeoxyris cf. carbonaria (shark egg case). BMB 014848 [W 467 b], counterpart ofJarzembowski (1989, fig. 17). Maximum width 6 mm. Fig. 11 . Palaeodictyopteran insect, hindwing base: note archedictyon present (after Jarzembowski, 1988). Scale line in mm. Fig. 12. Anthraconauta tenuis (fresh water bivalve) after Jarzembowski (1989). Scale line in mm. Species also present in the Sydney Coalfield and associated with Anthraconaia at Writhlington (Eagar, 1994).

PLATE II Fig. 1. Diplichnites cf. cuithensis (giant millipede trackway). W 901, Manchester Museum, coli. E. Jarzembowski. Width of trackway (external) 88 mm. Fig. 2. Kouphichnium aff. variabilis (walking trackways of horseshoe crabs). BMB 014878 [W 75a], counterpart of Jarzem­ bowski (1989, fig. 6). Scale line in mm. Fig. 3. cf. Pseudobradypus sp. (pelycosaur (mammal-like) reptile trace) associated with unnamed ostracod ichnofossils (PL. II, fig. 4). Footprint 63 x 29 mm (after Milner, 1994). Fig. 4. Ostracod (seed shrimp) trails and resting traces after Jarzembowski (1989). Illustrated area is a 17 x 30 mm area of the bedding plane. Fig. 5. Arthropod coprolites. BMB 014872 [W 186c] coli. P. Hardy. Scale in mm. Fig. 6. Cochlichnus sp. (nematode (roundworm) burrow after J arzembowski, 1989). Scale in mm. Rare, larger forms are cf. Lunichnium (Pollard & Hardy, 1991).

plant assemblages. No. 4 represents minimally Trace fossils transported material from low-lying forest (domi­ nated by Lepidodendron aculeatum) and growing In addition .to body fossils in the above plant-rich in a back-basin mire. In contrast, No. 6 is more crevasse splay depositional environments, there travelled material from drier, higher diversity levee are animal trace fossils from the margins of a forest. Animals are absent from the coal seams. floodplain lake (Pl. II). Diverse ethological as well as morphological data are thus available (see below). Body fossils (aquatic fauna) Remains of aquatic animals include horseshoe Interpretation crabs (Pl. I, fig. 7), water scorpions (Pl. I, fig. 8), conchostracans (Fig. 3), shark egg capsules (Pl. I, A numerical breakdown of the fauna is shown fig. 10) and non-marine bivalves (Pl. I, fig. 12). graphically (Fig. 4) to give a visual impression of Animals are rare in river channel sandstones. relative abundances. More animals have been The shelly lacustrine mudstone associated with found since the last published census but the the No. 1 seam is a distinctive facies. overall asymptotic 'curve' is unchanged for the

48 PLATE I

12:- • • lj. ,..= Ed. A. Jarzembowski, Atlas of animals from the Late Westphalian ... Geologica Balcanica, 34, l-2, 2004 PLATE II ·"'· /:'.'-.~: . ·.' ~ ,

Fig. 3. Anomalonema reumau.xi (conchostracan or spinicau­ date) carapace after B. Jarzembowski (1985). Length 5 rnm. Also known from northern France

lower 'shale' (i .e. excluding shell counts for the upper 'shale'). The main change is an increase in accessory taxa suggesting that W.G.N.R. has been well sampled. Thus we now know that the 'in­ det.' (indeterminate) category includes unique examples of a protozygopteran odonate, an am­ blypygid, and a eurypterid (Fig. 1; Pl. I, fig. 5; 8). Phagophytichnus seems confirmed and an un­ named seed-piercing ichnofossil (possibly the work of palaeodictyopteroids) has been added Fig. 1. Protophrynus carbonarius? (whipspider) restored by (Fig. 5; Pl. I, 3). There have been some inevita­ Proctor and Jarzembowski (1999) on lycophyte bark, based Eophrynus on Dunlop (1994a) ble taxonomic changes and is now known as Pleophrynus (Pl. I, fig. 2). The general palaeoenvironment at W.G.N.R. is a forested 'upper delta plain' with fresh water wetland and waterways as summarised above. The fauna is divided here into land and aquatic elements. Representative taxa are illustrated in ascending systematic order. More information and reconstructions are available in primary sourct?s, and work is continuing on a number of groups, e.g. the Araneida (true spiders) are being investigated separately at the University of Man­ chester. Some of the photographed specimens have been enhanced with ammonium chloride (e.g. Pl. I, fig. 5) or polarised light (e.g. Pl. I, fig. 9). The specimens illustrated are from the lower 'shales' at Writhlington except PL. I, fig. 12 from the upper 'shales' and PL. I, fig. 11 from Kilm­ ersdon colliery tip. Specimen registration num­ bers for newly figured material have their deposi­ tory prefixed as follows: BMB, Booth Museum Brighton; BR, Bristol City Museum; LL, Man­ chester University Musem; NHM, Natural Histo­ ry Museum, London. It is hoped that this atlas will aid Carboniferous geologists and palaeobotanists undertaking com­ parative studies and, moreover, encourage the recognition and recovery of faunas which must exist on the numerous colliery tips across Eura­ merica. Chance finds may be the first indicator of untapped potential. Acknowledgements. This is a contribution to IGCP 469's r~. 2. Arthropleu.ra armata (giant millipede) body seg­ Sofia meeting gratefully supported by the Royal Society. wents. BRSMG Cd 4057. Drawing by C. Proctor, scale line Thanks to Dr J. Pollard (Manchester) for comment on Text­ em. Species widespread in Westphalian D fig. 13.

- Geologica Balcanica, 1-2/2004 49 700

600

500

400 ~ Joo :5"'

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Fig. 4. Numbers (y-axis) of animals (x-axis) at W.G.N.R. to show relative abundances (from Jarzembowski, 1989)

Fig. 5. Seed hole (attributable to a palaeodictyopteroid insect) in Trigonocar- ---1 pus sp. Left, after Jarzembowski (2001). Scale line in mm

References Jarzembowski ,_~ .A. 2001. Review of early insects and pal­ aeocommumties. -In: Deuve, T. (ed.) Origin of the Hexa­ Anderson, L.I. 1994. Xiphosurans fom the Westphalian D of poda. -Ann. Soc. Entom. Fr., N. S., 37; 11-19. the Radstock Basin, , the South Wales Jar~em?owski, E.A., N~l •. A. 2002. The earliest damselfly­ Coalfield and Mazon Creek, Illinois.- Proc. Geol. Assoc. like msect and the ongm of modern dragonflies (Insecta: lOS; 265-275. ' Odonatoptera: Protozygoptera).- Proc. Geol. Assoc. 113· 165-169. , ' Austin, P.A. 2001. The Writhlington experience.- In: Bas­ sett, M.G., King, A.H., Larwood, J.G., Parkinson, N.A. Jarzembowski, E. A., Ross, A. 1993. Time flies: the geolog­ and De1sler, V.K. (eds) A future for fossils. Nat. Mus. .ical record of insects.-Geology Today, 9; 218-223. Wales, Geol. Se1:, 19; 67-70. Milner, A.C. 1994. A Carboniferous reptile footprint from the Somerset Coalfield. - Proc. Geol. Assoc. 105· Dunlop, J.A. 1994a. An Upper Carboniferous amblypygid 313-315. , , from the Writhlington Geological Nature Reserve. - Proc. Geol. Assoc., 105; 245-250. Pollard, J.E., Hardy, P.O. 1991. Trace fossils from the West­ Dunlop, J.A. 1_994b. The palaeobiology of the Writhlington phalian D of the Writhlington Geological Nature Reserve, tngonotarb1d arachnid. - Proc. Geol. Assoc. 105· nr Radstock, Avon. - Proc. Geol. Assoc., 102; 169-178. 287-296. , , Proctor, C.J. 1994. Carboniferous fossil plant assemblages Eagar, R.M.C. 1994. Non-marine bivalves from Writhling­ and palaeoecology at the Writhlington Geological Nature ton Geological Nature Reserve, Avon. - Proc. Geol. As­ Reserve. - Proc. Geol. Assoc., 105; 277-286. · soc., 105; 251-264. Proctor, C:.J. _1999. An Upper Carboniferous eurypterid from Jarzembowski, B.J. 1985. Fossil arthropods from Writhling­ the Wnthhngton Geological Nature Reserve.-Proc. Geol. ton tip. Unpublished report, London. Assoc., 110; 263-265. Jarzembowski, E.A. 1988. Prospecting for early insects. ­ Proctor, C.J., Jarzembowski, E.A. 1999. Habitat reconstruc­ tions in the late Westphalian of southern . Proc Open Univ. Geol. Soc. Jl, 9; 34-40, 2 pis. Jarzembowski, E.A. 1989. Writhlington Geological Nature Int. Palaeoemom. Conf (Moscow); 125-129. Thomas, B.A., Cleal, C.J. 1994. Plant fossils from the Writh­ Reserve. - Proc. Geol. Assoc., 100; 219-234. Jarzembowski, E.A. 1992. Writhlington Geological Nature lington Geological Nature Reserve. - Proc. Geol. Assoc., Reserve. - Palaeontology Newsletter, 13; 17-18. 105; 15-32.

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