Mor Phol Ogy and Palaeo Ec Ol Ogy of New, Non-Ma Rine Microconchid Tubeworm from Lower Car Bonif Er Ous (Up Per Mis Sis Sip Pian) of West Vir Ginia, Usa

Annales Societatis Geologorum Poloniae (2013), vol. 83: 37–50.

MOR PHOL OGY AND PALAEO EC OL OGY OF NEW, NON-MA RINE MICROCONCHID TUBEWORM FROM LOWER CAR BONIF ER OUS (UP PER MIS SIS SIP PIAN) OF WEST VIR GINIA, USA

Micha³ ZATOÑ1 & Rob ert L. PECK2

1 Univer sity of Silesia, Faculty of Earth Sciences, Bêdziñska 60, PL-41-200 Sosnowiec, Poland; e-mail: [email protected] 2 Concord Univer sity, Divi sion of Natu ral Sciences, Athens, West Virginia 24712, USA; e-mail: [email protected]

Zatoñ, M. & Peck, R. L., 2013. Mor phol ogy and palaeoec ol ogy of new, non-ma rine microconchid tubeworm from Lower Carboniferous (Upper Mis sissip pian) of West Vir ginia, USA. Annales Societatis Geologorum Poloniae, 83: 37–50.

Ab stract: A new species of a non-ma rine microconchid (Tentaculita) tubeworm, Microconchus hintonensis, from the Lower Car bonif er ous (Upper Mis sissip pian, Chesterian) of West Vir ginia, USA, is de scribed. Non-ma rine microconchids oc cur abun dantly in the de posits of the Bluefield, lower Hinton, Prince ton and Bluestone Forma - tions of the Mauch Chunk Group, where they are ei ther as soci ated with land plant remains and bi valve shells, or are pre served loose in the host sed i ment. The spec i mens at tached to plant re mains and bi valve shells, are poorly pre served, but those occur ring loose in the de posits are well-pre served in three di mensions. The in ter pre ta tion presented here, is that the loose speci mens of Microconchus hintonensis sp. nov. also orig i nally en crusted plants (land plants, al gae) and bi valve shells, but be came de tached af ter sub strate deg ra da tion and dis solu tion. The as soci a tion of land plant re mains, charophyte gyrogonites, bi valves, ostracodes, conchostracans, and fish teeth and scales, and the con comi tant lack of strictly marine fos sils in di cate that the microconchid-bear ing de posits of the lower Hinton, Princeton and Bluestone Forma tions were de posited in fresh-water en vi ron ments. Microconchus hintonensis sp. nov. is re garded as a highly fe cund, op por tu nistic spe cies that in large num bers col o nized ev ery available substrate in its habi tat. Its abundance in the depos its inves ti gated indi cates that the spe cies was well- adapted to the en vi ron ments it oc cu pied, even dur ing ep i sodes of higher sed i men ta tion rates and/or com pe ti tion with other soft-bod ied encrusters. During such ep i sodes, microconchids were able to grow ver ti cally by uncoil ing and el e vat ing their tubes, in or der to es cape po ten tial burial and/or overgrowth by other encrusters.

Key words: Microconchids, Mis sis sip pian, Car bon if er ous, encrusters, palaeo ec ol ogy, fresh-wa ters.

Manu script re ceived 29 April 2013, ac cepted 18 June 2013

IN TRO DUC TION Microconchids (Or der Microconchida Weedon, 1991) indi cate their lophophorate affin ity (Vinn and Mutvei, are ex tinct, seden tary, tentaculitoid tubeworms. Having cal- 2009; Taylor et al., 2010; Wil son et al., 2011; Zatoñ et al., citic skele tons, they are char acter ised by an excel lent fos sil 2012b), being even related to such suspen sion-feed ers as record, ranging from the Late Ordo vi cian to the Middle Ju- phoronids (Taylor et al., 2010). However, their true bio log i - rassic (Taylor and Vinn, 2006; Vinn and Mutvei, 2009; cal af fin ity is still un cer tain and spec i mens with ex cep tion- Zatoñ and Vinn, 2011). Origi nat ing in marine envi ron ments ally preserved soft tissues would be highly desir able for de- at least since the Early De vo nian, microconchids started to ci pher ing their po si tion on the an i mal phylo gen etic tree. invade fresh-water habi tats as well (Taylor and Vinn, 2006; Car bon if er ous microconchids, like oth ers from dif fer ent Caruso and Tomescu, 2012). Their occur rence in a wide suite systems, used to be treated and described under the polycha- of en vi ron men tal set tings, from nor mal ma rine through ete ge neric names Spirorbis or Serpula in the older (e.g., Mc- brack ish- to fresh-wa ter hab i tats (Zatoñ et al., 2012a), and Coy, 1844; Etheridge, 1880; Whitfield, 1882; Branson, 1937; their clear domi na tion on hard substrates and microbialites Elias, 1957; Howell, 1964; Leeder, 1973; Sando, 1984; Kie- during times follow ing mass extinc tions (Fraiser, 2011; tzke, 1990) and some recent liter a ture (e.g., Lescinsky, 1997; Zatoñ and Krawczyñski, 2011a; He et al., 2012; Zatoñ et al., Aitkenhead et al., 2002; Cassle et al., 2003; Falcon-Lang, 2013), make them a group of oppor tu nis tic organ isms. Al- 2005; Williams et al., 2005), even though many of them have though in cluded in the Class Tentaculita by Weedon (1991), been observed in strictly non-marine settings. On the basis the mor pho log i cal and microstructural features of the tube of their ce mented mode of life and lamellar tube mi cro- 38 M. ZATOÑ & R. L. PECK

Fig. 1. Lo ca tion of study area. A. Sketch map of West Vir ginia State, showing Missis sip pian de posits (shaded) and study area. B. Study area between Hinton and Bluefield. Circles indi cate loca tions of sam pled sections (see Table 1 for details) struc ture, some Lower Car bon if er ous ‘spirorbids’ were detailed study of the Lower Carbon ifer ous (Upper Missis - re-inter preted as vermetid gastro pods by Burchette and Rid- sippian, Chesterian) microconchids from the Mauch Chunk ing (1977), who later were supported by Wright and Wright Group of southern West Virginia, USA. On the basis of a (1981), Be³ka and Skompski (1982) and Paszkowski and rich col lec tion of spec i mens, thor ough mor pho log i cal and Szyd³ak (1986). It is notewor thy that Be³ka and Skompski mi crostruc tural ob ser va tions en abled rec og ni tion of a new (1982) were the first to in vesti gate the tube exter nal mor- microconchid species. Apart from taxon omy, its palaeoeco- phology, with the aid of the scanning electron micro scope logy is also discussed on the basis of taphonomic obser va - (SEM), an indis pens able tool in modern studies of this tions of many speci mens preserved in the host sedi ments. group of fossils. Except for the studies of Weedon (1990, This is thus the first detailed, taxo nomic and palaeoeco logi cal 1991), who clas si fied the Car bon if er ous ‘ver mi form’ gas- study of fresh- and brackish-wa ter microconchids in gen eral. tropods and ‘spirorbids’ of earlier authors in the Order Mi- croconchida, there is a lack of modern studies deci pher ing the tax on omy of Car bon if er ous microconchid tubeworms, GEO LOG I CAL SET TING which def i nitely would as sist rec og ni tion of the di ver sity of these enigmatic fossils during that period. Although the ma- The geo log i cal set ting of the Up per Mis sis sip pian of jority of modern studies were focused on marine forms southeastern West Virginia (Fig. 1A) has been fairly well (e.g., Vinn, 2006; Vinn and Taylor, 2007; Zatoñ and Kraw- studied due to the in ter est in the Missis sippian – Pennsyl va- czyñski, 2011b), there is a signif i cant gap in taxo nomic nian boundary that is ex posed in the study area. The rocks stud ies, con cern ing fresh- and brack ish-wa ter spe cies. Rec- of the Upper Missis sip pian Mauch Chunk Group show a ogni tion of the diver sity of such species would provide the transition from shal low, open marine limestones of the answers to many questions, such as: 1) are there any mor- Greenbrier Se ries, into the mixed, ter res trial de pos its with pho log i cal and microstructural sim i lar i ties/dif fer ences be - thin, marine incur sions of the Mauch Chunk Group (Fig. 2), tween fresh-and brackish-wa ter and marine forms?, 2) and then into the coal-bearing, Pennsyl va nian siliciclastic could the same species have lived in both fresh- and brack- depos its (Ettensohn, 2009). This influx of clastics re flects a ish-wa ter or even marine envi ron ments?, or 3) in what envi - major change in sed imen tation that was at least partly con - ron ments and at what times did they at tain the great est di - ditioned by the movement of the Appa la chian Basin north- versity? To an swer these inter est ing questions, some weight wards into a more humid, trop ical cli mate belt (Cecil et al., should now be put on fresh- and brackish-wa ter settings. 2004). The change in lithol ogy from limestones to siliciclas- In the present paper we turn the empha sis toward the tics was primar ily a re sult of tec tonic plate move ments. This under stand ing of some of the problems of the fresh- and could well reflect the change from the tec toni cally passive brack ish-wa ter microconchids. As an ex am ple, we pres ent a con di tions of the Greenbrier to the more active tectonism of NON-MA RINE MICROCONCHID TUBEWORM, LOWER CARBONIFEROUS 39

the Mauch Chunk and Pennsyl vanian along the Alleghanian orogen. This could be a product of the early Alleghanian Orog eny and in creas ing, tec tonic ac tiv ity (Chesnut and Greb, 2009) or a relaxational response to previ ous conver - gence at the SE margin of Laurussia in the fi nal phases of the Acadian Orogeny (Ettensohn, 2009). The Lower Car bon if er ous (Mis sis sip pian) Sys tem of the Ap pa la chian Ba sin com prises a third-or der se quence, defined by an uncon formity or abrupt transi tion at the base of the Sunbury Shale and by an early Pennsyl vanian uncon - formity at the top (Ettensohn, 2009). The Upper Missis sip - pian Mauch Chunk Group in southeastern West Virginia is subdi vided into the Bluefield, Hinton, Princeton, and Blue- stone For ma tions, with a max i mum ag gre gate thick ness of approx i mately 1051 m, of which approx i mately 650 m are ex posed in the map area (Matchen et al., 2011). The basal Up per Mis sis sip pian Bluefield For ma tion (Fig. 2), compris ing gray, calcar e ous shales and blocky, red- dish mudstones, begins the upward trend to ward siliciclastic depos its with the Glenray and Reynolds Limestones, the products of two spasms of marine incur sion near the bottom of the For ma tion. Sub se quently, siliciclastic de pos its were the rule. The Coney Shale of Reger (1926) near the top of the for ma tion con tains ma rine fos sils, in clud ing brachi o - pods. The thin Coney Limestone, which only has microconchids, fish teeth and scales, and ostracodes (Stencil, 2012), gives way to the Hinton Forma tion and the Stony Gap Sand - stone Member (Maynard et al., 2006). The Stony Gap Sandstone, the basal mem ber of the Hinton Forma tion (Fig. 2), mainly comprises a white, fine- to medium-grained quartz arenite, although it changes locally to a light gray, fine-grained, lithic arenite. The unit is typ ically cross-bedded, with both trough and planar cross- beds present. It has a fluvial-estuarine ori gin, asso ci ated with palaeovalley inci sion and a basinward shift of fluvial Fig. 2. Stra tig ra phy of Up per Mis sis sip pian Mauch Chunk envi ron ments during a time of lowered, rela tive sea level Group with general section, showing mainly siliciclastic depos its (Miller and Eriksson, 2000). Alter na tively, Englund (1979) (simpli fied after Stencil, 2012) in stratigraphic frame work as sec- ond-or der se quence (af ter Maynard et al., 2006). Shaded, hor i zon - suggested that the Stony Gap Sandstone was a series of ma- tal bars in up per Hinton For mation re fer to ma rine members (af ter rine bars. Above the Stony Gap Sandstone and a fossilife- Beuthin & Blake, 2004). Shaded ver ti cal bars re fer to gen eral rous shale (mostly non-marine bivalves) overly ing it, the strati graphic range of sam pled sec tions at par tic u lar lo cal i ties (for rocks be come terrigenous depos its, consist ing mainly of red numbers, see Fig. 1B and Ta ble 1). SGSM – Stony Gap Sand stone mudstones and thin, interbedded, lentic u lar or channel-fill Member, LSGM – Little Stone Gap Member, FMM – Five Mile sandstones. Occa sional inter vals of black mudstones and Mem ber, EMM – Eads Mill Member, TST – transgressive sys tem tan to yel lowish or gray-green, limey mudstones (Fig. 3A) tract, MFS – max i mum flooding surface, HST – highstand systems contain microfossils (microconchids, ostracodes, and fish tract teeth and scales). Charophyte oogonia and conchostracan Hemicycloleaia speci mens have been found in sepa rate ho- dom i nately micritic and ar gil la ceous and con tains a typ i cal ri zons, re flecting a rather fresh-water palaeoenvironments Chesterian fauna of brachio pods, bryozoans, corals, bival- (R. L. Peck, field obser va tions). However, most of the mud- ves, gastro pods, ostracodes, trilobites, pelmatozoans, and stones are red and often ex hibit char acter istics of palaeo- cepha lopods (Reger, 1926; Gordon and Henry, 1981; Beu- sols. They proba bly origi nated as overbank flood depos its. thin and Blake, 2004; Matchen et al., 2011). The Little Stone The mudstones and sand stones reflect de po sition in terres - Gap Member provides a brief change from the red mudstones trial and coastal envi ron ments (Miller and Eriksson, 2000). of the lower Hinton Forma tion that lack the region ally identi - Hori zons of plant fossils are found from the Stony Gap fiable, marine units (Beuthin and Blake, 2004). A vari able Sandstone through the entire Forma tion into the Princeton succes sion of mudstone, sandstone, and limestone, with lim- Sand stone, sup port ing the in ter pre ta tion that subaerial ex- ited oc cur rences of coal, occurs above the marine Lit tle Stone posure occurred fairly frequently. Gap Mem ber. In this part of the forma tion (upper Hinton For- Near the middle of the Hinton Forma tion, a marine mation), thin, fairly wide spread, marine zones are found in limestone, the Lit tle Stone Gap Member (for merly the Avis the Fivemile and Eads Mill members (see Fig. 2; Beuthin and Limestone, see Beuthin and Blake, 2004) occurs. It is pre- Blake, 2004; Vance, 2007; Matchen et al., 2011), contain ing 40 M. ZATOÑ & R. L. PECK

Fig. 3. Ex am ples of sam pled sec tions. A. Sec tion with de posits of lower Hinton For mation at State Route 20, Bluestone Lake, Sum- mers County (lo cal ity no. 8). Beds, indi cated by as ter isks, re fer to fossiliferous mudstones with myalinid bi valves, Hemicycloleaia bran - chi opods, ostracodes, fish teeth and scales, as well as abundant microconchids. B. Sec tion with de posits of Bluestone For mation at County Road 11, Crane Creek Road, Montcalm, Mer cer County (lo cal ity no. 13). Sam pled gray to black, shaly mudstone is indi cated by as ter isk a typi cal Chesterian fauna of brachio pods, bryozoans, corals, spec imens of microconchids were retrieved from the Blue- bivalves, gastro pods, ostracodes, trilobites, pelmatozoans field Forma tion. In general, however, microconchids were and cephalo pods (Reger, 1926; Cooper, 1948, 1961; Henry no ticed in both fresh-, brack ish and marine depos its of the and Gordon, 1979, 1992). Mauch Chunk Group in the area studied, but not all de- The Hinton Forma tion is overlain by the Princeton For- served special atten tion, owing to their state of preser va tion. mation (Fig. 2), com prising medium- to coarse-grained, For exam ple, the marine deposit of the Eads Mill Mem ber quartzose sandstone to quartz arenite, and contain ing quartz of the upper Hinton Forma tion contained Composita brachi- pebbles and conglom erate beds. Thin beds of mudstone, o pods with some encrust ing microconchids, but their state sandstone, coal, and palaeosols are also locally ob served of pres er va tion was in suf fi cient for de tailed study. The above the sandstone (Matchen et al., 2011). The sandstone same might be said of the microconchids encrust ing bivalve is another incised valley fill (Miller and Eriksson, 2000). shells and plant frag ments, pre served in the siliciclastics of Above the Princeton Forma tion, the Bluestone Forma - the Bluefield, lower Hinton, Princeton and Bluestone For- tion (Fig. 2) occurs, in the form of mudstones, shales, silt- mations. Although they may be numer ous where attached to stones, and sandstones (Fig. 3B), with discon tin u ous beds these substrates, their poor state of preser va tion obscured of coalesced, authigenic limestone and sid erite nodules the de tails of their tubes. Ad di tion ally, many spec i mens (Matchen et al., 2011). A few thin, discon tin u ous, im pure were pre served as traces of their tubes on the shelly and coal beds are pres ent, as well (Matchen et al., 2011). Blue- plant substrates. Such speci mens, of course, are not suit able stone marine zones (Bramwell Mem ber) con tain a typ ical for mor pho log i cal and microstructural in ves ti ga tion. How- Up per Mis sis sip pian (Chesterian) fauna of brachi o pods, ever, they do provide inter est ing mate rial for taphonomic bryozoans, corals, bivalves, gastro pods, ostracodes, trilo- and palaeo eco logi cal ob ser va tions. Many sam ples with bites, pelmatozoans, and cephalo pods (Reger, 1926; Henry such microconchid-encrusted bivalve shells and plant frag- and Gordon, 1979, 1992; Hoare, 1993). ments (shoots and leaves) were collected for these purposes. Corre la tion of biostratigraphic data with Euro pean suc- A number of siliciclastic rock samples from the lower ces sions in di cates that the Bluefield For ma tion cor re sponds Hinton Forma tion were rich in microconchid tubes, scat- to the Upper Viséan, and the Hinton to Bluestone Forma - tered in the host sedi ment along with ostracode cara paces. tions corre spond to the lower part of the Namurian (see On the bedding planes, the tubes appeared to be well-pre- Beuthin and Blake, 2004; Maynard et al., 2006; Ettensohn, served. There fore, it was de cided to re trieve the fos sils by 2009). boiling the rock samples with Quater nary-O. To do so, the rock sam ples (0.5 to ca 3 kg) were put in a pot, covered with water, and about a table spoon of Quater nary-O was added. MA TE RIAL AND METH ODS This prepared mix was then slowly boiled for several hours. Next, the samples were gently washed with hot wa ter to All the mate rial in ves tigated here, co mes from the flush out the floating parti cles and the Quater nary-O. After Lower Car bon if er ous (Up per Mis sis sip pian) Mauch Chunk drying the sam ples, they were sieved through a mesh of 1.4 Group of West Virginia, USA (Figs 1, 2). The de posits of mm (No. 14), 0.5 (No. 35) and 0.212 mm (No. 70) and the all forma tions of the Mauch Chunk Group were inspected re sult ing res i dues were ex am ined for fos sils. Microcon- with respect to microconchids. The bulk of speci mens were chids, along with other microfossils, occurred to be in the found in the Hinton and Bluestone forma tions, while a few resi dues of the 1.4 and 0.5 mm sieves. These resi dues were NON-MA RINE MICROCONCHID TUBEWORM, LOWER CARBONIFEROUS 41

Ta ble 1 Details on prove nance of samples used for extrac tion of microconchids from Upper Missis sip pian Mauch Chunk Group of West Virginia, USA

Locality Sample Stratigraphy Lithology River Road, County Road 26, Raleigh Yellow tan to olive gray, mottled, 1 182 Coney Limestone, Bluefield Formation County poorly lithified, calcareous mudstone Tug Creek Mountain Road, County 2 130 Gray member, Bluestone Formation Black, coaly, poorly lithified mudstone Road 44/6, Summers County Elk Knob Road, County Road 9, Bellepoint Limestone?, lower Hinton Yellow tan to gray black, calcareous, 3 107 Hinton, Summers County Formation shaly mudstone Leatherwood Road, County Road 44/7, Gray to black, whitish when weathered, 4 153-154 Lower Hinton Formation Hinton, Summers County thinly laminated, shaly mudstone Black silty mudstone, thin to thick State Route 20, below Bluestone Dam, Lower Bellepoint Shale?, lower Hinton 5 070, 157 laminations, with ostracodes and Hinton, Summers County Formation Carbonicola bivalves Beds of tan and gray to black shaly, State Route 20, below Bluestone Dam, Lower Bellepoint Shale?, lower Hinton 6 111-113, 115 silty mudstones sandwiched between Hinton, Summers County Formation sandstones and siltstones State Route 20, S of Leatherwood Blackish mudstone to claystone with 7 Road, County Road 44/7, Hinton, 195, 197 Lower Hinton Formation ostracodes Summers County 007, 020, 045-046, Shaly, laminated mudstones, gray at the State Route 20, Bluestone Lake, 088, 094, 121, 147, bottom with myalinid bivalves and 8 Lower Hinton Formation Summers County 171-172, 175, 198, Hemicycloleaia branchiopods to black, 199-201 coaly, poorly indurated mudstone at top County Route 20/2, Bluestone State Lower Bellepoint Shale?, lower Hinton Black, highly carbonaceous to almost 9 097, 099-100 Park, Summers County Formation coaly, shaly mudstone Yellow tan mudstone to claystone with County Route 20/2, Bluestone State 086, 090-081, 122, Lower Bellepoint Shale?, lower Hinton 10 Carbonicola bivalves, charophyte Park, Summers County 166, 178, 184 Formation gyrogonites, and ostracodes State Route 20, S of Bluestone Bridge, Lower Bellepoint Shale?, lower Hinton 11 093 Gray to black mudstone True, Summers County Formation County Road 44/11, Road to Bull Falls, Bellepoint Limestone?, lower Hinton Orange tan to gray, calcareous 12 170 Bluestone Lake, Summers County Formation mudstone County Road 11, Crane Creek Road, 13 162 Gray Member, Bluestone Formation Gray to black shaly mudstone Montcalm, Mercer County US Route 460, Princeton, Mercer Black, thinly laminated, shaly mudstone 14 048, 168 Gray Member, Bluestone Formation County with bivalves and ostracodes Local ity num bers corre spond to those in Figure 1B. Each sam ple num ber is preceded by acro nym, GIUS 5

rinsed 3 to 6 times in hot wa ter, dried and then picked for un der the binoc u lar micro scope, those from two sam ples of microfossils. We used this method for 42 samples from dif- the lower Hinton Forma tion and two samples from the Blue- ferent hori zons of the Mauch Chunk Group (Fig. 2) in Ra- stone Forma tion were discarded, owing to their poor state of leigh, Sum mers and Mer cer Coun ties, near Hinton and preser va tion. From the remain ing 38 samples, the best-pre- Prince ton (Fig. 1B; for de tails, see Ta ble 1): the Bluefield served spec imens were se lected for further, detailed ob ser - Forma tion (one sam ple), the lower Hinton Forma tion (37 vations. In total, 175 microconchids were mounted on steel samples) and the Bluestone Forma tion (4 samples). Surpris - tables using carbon tape, and exam ined in an uncoat ed state ingly, this method provided a number of microconchid spe- using a Philips XL 30 low-vacuum envi ron men tal scan ning cimens, the major ity of which come from the lower Hinton electron micro scope (ESEM), housed at the Faculty of Earth For ma tion. Sci ences in Sosnowiec, Po land. Im ages were gen er ated The preser va tion of spec imens varies, from tubes being using back scat tered elec trons (BSE de tec tor). Ad di tion ally, completely obliter ated, owing to ferrous oxides, to those 8 speci mens were embed ded in epoxy resin and polished for with cal car e ous tubes. The lat ter com prise spec i mens with microstructural obser va tion of the tubes. Measure ments of both flattened tubes due to compac tion and those with the microconchids were performed directly on the ESEM three-dimensionally well-preserved tubes. The num ber of photomicrographs. spec imens per treated sample dif fers widely, from as few as The spec imens are housed at the Fac ulty of Earth Sci - 4 speci mens to as many as dozens to hundreds of speci mens ences, Univer sity of Silesia in Sosnowiec, Poland, desig - for a given sample. Af ter initial in spection of all spec imens nated GIUS 5-3620. 42 M. ZATOÑ & R. L. PECK

SYS TEM ATIC PALAE ON TOL OGY mens, the jux ta po si tion of per pen dic u lar ridges and lon gi tu di nal striae re sults in the for ma tion of thick ened el e va tions, tu ber cles or Class TENTACULITA Bouèek, 1964 even node-like struc tures (Fig. 4A, C, K). The in ten sity of the lat- Order MICROCONCHIDA Weedon, 1991 ter structures varies widely in dif ferent speci mens. However, inter - me di ate stages of their de vel op ment are ev i dent. In spec i mens, in Ge nus Microconchus Murchison, 1839 which the out er most tube layer is worn away, per pen dic ular ridges Type spe cies Microconchus carbonarius Murchison, 1839 are the only or namen tal features. Di ag no sis: Tube planispirally coiled, with a ten dency for he li cal Tube or i gin bul bous, el lip ti cal in outline, ca 243 µm in width un coil ing in later on tog eny. Ex te rior sur faces or na mented with and 260 µm in length, sep a rated from the rest of the tube by dis- var i ously de vel oped growth lines, per pen dic u lar ridges, lon gi tu di - tinct constric tion. Its ex te rior is or na mented by straight, per pen - nal striae and tuber cles or nodes. Minute punctae pene trate lamel- dic u lar ridges, ca 63 µm apart (Fig. 6). lar tube microstructure. Tube consists of lamellar layer with microlaminae, pene trated Re marks: The ge nus Microconchus dif fers from Palaeoconchus by tiny punctae (Fig. 7A–C). The punctae are cir cu lar in out line (see Vinn, 2006; Zatoñ and Krawczyñski, 2011a) in pos sessing and ca 2.2 µm in diam e ter (Fig. 7B, C). tiny pores (punctae) pen e trat ing the tube. From Punctaconchus Discussion – variabil ity : The species Microconchus hintonensis (see Vinn and Taylor, 2007) it dif fers in hav ing much smaller sp. nov. shows a clear, intraspecific variabil ity with respect to coil- punctae and a tube with a ten dency to un coil. From Annuliconchus ing pat tern and tube or na menta tion (Fig. 4). The coil ing pat tern (Vinn, 2006) it dif fers in lack ing in ter nal annulation. The ge nus may vary widely among dif fer ent in di vid u als. Some tubes show Helicoconchus (see Wilson et al., 2011) dif fers in lack ing punctate ev i dence of reg u lar coil ing dur ing the an i mal growth, leav ing the tube microstructure and in bud ding new tubes from ex ist ing ones. umbi li cus open through out (e.g., Fig. 4B, F, H), while oth ers are tightly coiled in the last whorl, re sult ing in a nar row or even closed Microconchus hintonensis new species umbi li cus (Fig. 4C, E). Many spec i mens were at tached through out their growth, which may be re flected in the flat tube base along its Figs 4, 6–8 growth di rec tion. How ever, oth ers have the ter minal part of their Di ag no sis: Tube planispirally coiled at the first stages of tube tube ori ented ver ti cally up wards. This may be a ten dency for tube growth, fol lowed by he li cal un coil ing of it. Tube ex te rior or na - uncoil ing and ver ti cal growth at some point in their devel opment, mented with var i ously spaced, thicker, trans verse rib-like ridges as many speci mens in differ ent samples have even the greater part crossed by thin ner, lon gi tu di nal striae. The crossing ridges and of their tubes uncoiled, and with a ver ti cal ori en ta tion (Fig. 4F). striae form dis tinct, but var i ously de vel oped tu ber cles and nodes. This prob lem is discussed in de tail in the palaeoeco logi cal sec tion. Tube or i gin (protoconch) bul bous, or na mented with widely- The or na men ta tion of the tube ex te rior shows con sid er able spaced, sharp, per pen dic u lar ridges. vari a tion. This should not be sur pris ing, since in other mi cro- Et y mol ogy: From the name of the Hinton For mation. conchids such vari abil ity of or na menta tion also was noted (e.g., Zatoñ and Krawczyñski, 2011a). The vari abil ity not only af fects Types: Holotype: GIUS 5-3620/121/01 (Fig. 4J), Hemicycloleaia the trans verse ridges, which may be straight or sin u ous, and thin or bed, lower Hinton For mation, Mauch Chunk Group, Upper Mis - thicker, but also the lon gi tu di nal, thin striae, crossing the ridges sissip pian, State Route 20, Bluestone Lake, Summers County, per pen dic ularly. The striae may be fine along the tube. However, West Virginia, USA. Paratypes: GIUS 5-3620/020/02, GIUS more commonly they have an ir reg u lar thick ness, re sult ing in a 5-3620/046/03, GIUS 5-3620/086/01, GIUS 5-3620/086/06, GIUS wrinkle-like or wavy appear ance in places, where they are thicker 5-3620/097/02, GIUS 5-3620/097/05, GIUS 5-3620/097/07, GIUS (see Fig. 4). More over, in some in di vid uals they de velop into 5-3620/097/12, GIUS 5-3620/097/15, GIUS 5-3620/099/02, GIUS forms of tuber cles or nodes, where they cross the ridges. Their ap - 5-3620/121/01, lower Hinton Forma tion, Mauch Chunk Group, pear ance is also irreg ular on the tube ex te rior. Such tuberculation Upper Mis sissip pian, West Vir ginia, USA; GIUS 5-3620/130/05, may ap pear very early in microconchid on tog eny, oc cur ring just Bluestone For mation, Mauch Chunk Group, Upper Missis sip pian, af ter the protoconch. In many spec i mens, such tuberculation is ab - West Virginia, USA. sent. In such spec i mens, the lon gi tu di nal striae are also lack ing Ma te rial: Hundreds of var i ously pre served spec i mens, of which and only transverse ridges are present. However, in many cases 183 spec i mens (includ ing 8 sec tioned ones) have been stud ied in such speci mens are charac ter ised by a worn tube exte rior. That the de tail us ing ESEM. wrin kle and tu ber cu late or na men ta tion may de pend on the state of De scrip tion: Tube small, planispirally (dextrally) coiled (e.g., pres er va tion is well ex empli fied by par tially crushed tubes: in de- Fig. 4A, C–E, G–I) up to ca 2170 µm in diam e ter, be ing later heli - pres sions, where such or na menta tion is well-preserved, and on the cally un coiled (Fig. 4B, F). Umbi li cus open, dif fer ing in width in top, where it is lacking (Fig. 8A). Therefore, it is believed that on dif fer ent spec i mens (129–568 µm, mean = 322 µm, n = 79), with the one hand, the pres ence of tu ber cles/nodes may de pend on the rounded mar gin and gently in clined slope. Ap er ture rounded to state of pres er va tion of the spec i mens, be ing de vel oped on tubes semi-rounded, up to 978 µm in di ame ter. The in crease in tube hav ing a well-pre served outer sur face. On the other hand, the pres - width proceeds as the tube diam e ter increases. The um bili cal ence of tuberculation also may de pend on well-de vel oped striae, width, on the other hand, is not so well cor re lated with tube di ame - crossing the transverse ridges. The pres ence of tubes with more or ter growth (Fig. 5), suggest ing a greater de vel op mental plastic ity less well-de vel oped tuberculation in the spec i mens studied in di - in the latter feature. cates the presence of a morpho log i cally vari able species that lived Tube ex te rior or na mented by fine growth lines and more or during sed i menta tion of both the lower Hinton and Bluestone For- less thicker, trans verse rib-like ridges, run ning straight or sin uous ma tions. The char ac ter is ti cally or na mented protoconches, in the to tube base or around the tube in uncoiled speci mens (Fig. 4). The form of transverse, widely spaced ridges, oc cur ring in both tu ber- ridges and growth lines are ir reg ularly crossed by lon gi tu di nally culated tubes and in those, where the ex te ri ors are de void of well- run ning (in the tube growth di rec tion) thin ner striae. The striae de vel oped lon gi tu di nal striae, also may in di cate the pres ence of a may be de vel oped on a whole tube surface, or may be con fined sin gle, al beit mor pho log i cally vari able spe cies. This may also be only to its lat eral side. They also may run con tin uously along the supported by the presence of simi lar sizes of punctae, a feature that tube growth di rec tion, or, more commonly, be in ter rupted, giv ing may vary be tween differ ent species of the same genus (e.g., Punc- a wrin kle-like ap pear ance (e.g., Fig. 4B, D, G, L). In many spec i - taconchus, see Vinn and Tay lor, 2007). NON-MA RINE MICROCONCHID TUBEWORM, LOWER CARBONIFEROUS 43

Fig. 4. Microconchids Microconchus hintonensis sp. nov. from Upper Missis sip pian Mauch Chunk Group of West Vir ginia, USA. A. GIUS 5-3620/020/02, B. GIUS 5-3620/046/03, C. GIUS 5-3620/086/01, D. GIUS 5-3620/086/06, E. GIUS 5-3620/097/15, F. GIUS 5-3620/097/05, G. GIUS 5-3620/097/02, H. GIUS 5-3620/097/12, I. GIUS 5-3620/097/07, J. GIUS 5-3620/121/01 (holotype), K. GIUS 5-3620/099/02, L. GIUS 5-3620/130/05. Arrows indi cate healed in ju ries. A–K – lower Hinton Forma tion, L – Bluestone For mation. BSE ESEM im ages of un coat ed spec i mens 44 M. ZATOÑ & R. L. PECK

Fig. 5. Plots of umbil i cal width and whorl width against tube di ame ter for Microconchus hintonensis sp. nov.

Fig. 6. Mor phol ogy of tube or i gin (ar rowed) in Microconchus hintonensis sp. nov. from lower Hinton (A, B) and Bluefield (C) For mations. A. GIUS 5-3620/097/01, B. GIUS 5-3620/168/03, C. GIUS 5-3620/182/03. BSE ESEM images of un coat ed spec i mens

Fig. 7. Tube microstructure of Microconchus hintonensis sp. nov., show ing wavy microlaminae (A, B), punc tured by tiny punctae (A–C, ar rowed). A. GIUS 5-3620/099/06, B. GIUS 5-3620/097/04, C. GIUS 5-3620/020/06. BSE ESEM images of un coat ed spec i mens

A hand ful of spec i mens, de rived from the Bluefield For ma- that em ployed a bin oc u lar mi cro scope. There fore, any de tailed tion (Co ney Lime stone), seem to be de void of in di vid u als with the doc u men ta tion, com pris ing SEM pho to mi cro graphs of the tube wrin kle-like and tu ber cu late or na men ta tion, char ac ter is tic for the surface and tube microstructure, are sim ply lack ing. More over, de - spec i mens from the lower Hinton and Bluestone For mations. scrip tions of each microconchid spe cies in the 19th Cen tury lit er a - However, the major ity of speci mens are small (presum ably ju ve- ture are sup ple mented only with hand-draw ings, showing a rather niles) and those larger (pre sumably adults) have the tube ex te rior gen eral tube ap pear ance. The only work, in clud ing de tailed SEM too poorly pre served for the orig i nal or na menta tion to be deci - pho to mi cro graphs of Car bon if er ous microconchids, is that of phered. But, as in the tubes from the lower Hinton and Bluestone Be³ka and Skompski (1982). The spec i mens came from the Polish Forma tions (Fig. 6A, B), the Co ney Lime stone spec i mens have Lower Car bonif er ous (Viséan), and then were con sidered to be simi larly or na mented protoconchs (Fig. 6C). Thus, it is plau sible archaeogastropods by the au thors. They are charac ter ised by heli - that they might repre sent the same species. cally un coiled tubes, with strong, thick and more or less reg u larly Dis cus sion – com par i sons: The com par i son of the spe cies de - spaced, trans verse ridges, dis sim i lar to the or na men ta tion pat terns scribed here, is rather lim ited as the great ma jority of previ ously of Microconchus hintonensis sp. nov. As Be³ka and Skompski de scribed and il lus trated Car bon if er ous microconchids (usu ally (1982) stated, the tubes are close to those de scribed un der the un der the name Spirorbis) were in ves ti gated us ing clas sic meth ods name Spirorbis caperatus by Mc Coy (1844). Microconchus pusi- NON-MA RINE MICROCONCHID TUBEWORM, LOWER CARBONIFEROUS 45

Fig. 8. Dif fer ent states of pres er va tion of Microconchus hintonensis sp. nov. tubes. A. Wrin kle or na men ta tion, pre served in de pressed (con cave) por tion of tube (arrowed), while top of three-dimensionally (con vex) pre served part is nearly de void of it. B. Mould of tube, show ing only rem nants of trans verse ridges, while per pen dic u lar wrin kle or na men ta tion is lack ing. C. Cross-section of tube, showing in - fill ing of it by car bon ate and py rite min eral phases. BSE ESEM images of un coat ed spec i mens llus (Martin) from Westphalian of Yorkshire is also dissim i lar, TAPHONOMY AND PALAEO EC OL OGY hav ing closely-spaced fine trans verse ridges and a completely smooth tube or i gin. How ever, it has simi lar, tiny tube punctation Microconchids from the Missis sip pian Mauch Chunk (see Tay lor and Vinn, 2006, fig. 1K, L). Sev eral spe cies of Car - Group of the area studied are differ ently preserved in partic - bon if er ous ‘Spirorbis‘ were described and illus trated from Great u lar forma tions. The spec imens, re trieved from bulk sam - Brit ain by Etheridge (1880). Of them, only one spe cies, ‘Spi- ples from depos its of the Bluefield, lower Hinton and Blue- rorbis‘ spinosa (de Koninck) may be somewhat simi lar in hav ing stone For mations, dif fer from the to tally or partially flat - “small, sharp prickles, or abor tive spines, ar ranged in quincunx” tened (Fig. 8A) to three-dimensionally pre served tubes. (Etheridge, 1880, p. 262) on the tube sur face. However, Mi cro- conchus hintonensis sp. nov. clearly dif fers from this species, as it Microconchids, attached to plant shoots and leaves (Blue- has no spines, possesses addi tional transverse ridges and lon gi tu di - field and Princeton Forma tions), as well as to bivalve shells nal striae, and its tube base is not crenulated. (lower Hinton Forma tion), are usually flattened, and the ex- Another form that is notewor thy is a spe cies de scribed as ter nal surfaces of their tubes are badly pre served (Fig. ‘Spirorbis‘ nodulosus ((Hall) (see Whitfield, 1882, pl. 9, fig. 31), 9A–C). Commonly, the microconchids, preserved on plants from the Lower Car bon if er ous of Indi ana, USA. Its tube, however, or shells, are also preserved in the form of traces of their possesses large nodes, reg ularly ar ranged in three rows; sharp, tube bases (Fig. 9C); the speci mens themselves most proba - short ridges near the umbil i cal mar gin and a dor sal crest, running bly had fallen out. Microconchid tubes, attached to plants or half of the way to the last whorl. So, al though they have nodes, shells in fresh-wa ter depos its, usually are poorly pre served, these are completely dif ferent from irreg u larly scat tered, small tuber cles in Microconchus hintonensis sp. nov. ‘Spirorbis‘ moreyi, a with tubes diagenetically al tered by do lo mite (Zatoñ and ma rine Mis sis sip pian spe cies from Wy o ming, de scribed by Bran- Mazurek, 2011; Caruso and Tomescu, 2012). Such a state son (1937, pl. 89, figs. 1, 2), pos sesses slightly sin u ous, trans verse of pres er va tion ham pers the di rect com par i son of these mi- ridges and prob a bly also fine, longi tu di nal striae, as may be de - croconchids to those well-preserved speci mens, retrieved duced from the descrip tion. However, it lacks the charac teris tics from the host sed i ment. How ever, it is be lieved that well- for Microconchus hintonensis sp. nov., in clud ing the ir reg ular, preserved microconchids, occur ring as both loose speci - wrinkled pat tern, formed by the crossing, trans verse ridges and mens, scattered in the host sedi ment (Fig. 9D) and attached longi tu di nal striae, as well as tuberculation. to organic substrate in the Bluefield and lower Hinton For- The Mis sissip pian spe cies, known as ‘Spirorbis‘ kentuckien- mations, belong to the same spe cies, Microconchus hinto- sis, de scribed by Howell (1964), pos sesses only thick, trans verse nensis sp. nov. The flatten ing of the tubes presum ably re- ridges. In ter est ingly, it is rep re sented by tubes of both dextral (clock wise) and sinistral (anticlockwise) coil ing and thus may be sulted from later sedi ment load on the speci mens. However, al lied to Mid dle De vo nian spe cies, such as ‘Spirorbis‘ arkonensis those un com pacted and three-dimensionally pre served (Nichol son, 1874). tubes were af fected by early diagenetic pre cip itates, es pe - The most simi lar species, with a tube orna mented with sim i- cially carbon ates and pyrite, fill ing the empty spaces within lar, transverse ridges and lon gi tu di nal striae, giv ing a wrin kle-like them (Fig. 8C). Although many speci mens from the host pat tern, is a microconchid de scribed as ‘Spirorbis‘ sp. A, from the sedi ments have the tubes worn, part of them still retains the ma rine Penn syl va nian de pos its of New Mex ico, USA (Kietzke, cal car e ous tube min er al ogy with a well-pre served lamellar 1990). Whether it is a sepa rate new species, or maybe a form microstructure (Fig. 7). The dif fer ences in the occur rences conspecific with Microconchus hintonensis sp. nov., is un cer tain. of such tube preser va tion may depend on changing micro- First, a de tailed study of a large num ber of the New Mex ico spec i - mens using SEM, should be con ducted and, sec ond, the pref er - en vi ron ments within the sedi ment. It is plau si ble that ep i - ences of Microconchus hintonensis sp. nov. for ma rine en vi ron- sodi cally the tubes were buried in sedi ment, rich in plant de- ments must be confirmed, on the ba sis of a large num ber of tri tus, the degra da tion of which may have low ered the al ka- well-pre served spec i mens. lin ity of the pore wa ters, thus result ing in tube etching and Oc cur rence: Upper Missis sip pian (Mauch Chunk Group, cer - dis so lu tion (Fig. 8B). tainly in the lower Hinton and Bluestone For mations, pos sibly in The presence of land plant remains, bivalves (myali- the Bluefield For mations, as well) from the en vi rons of Hinton and nids, Modiolus, Carbonicola, Anthraconaia), ostracodes, Bluefield, West Virginia, USA. branchi o pods, fish teeth and scales, as well as charophyte 46 M. ZATOÑ & R. L. PECK

Fig. 9. Microconchids and their sub strates. A. microconchid (ar rowed), en crust ing the land plant’s shoot, Bluefield For mation. B. Microconchids, en crust ing leaf let of land plant, Prince ton For mation. C. Sev eral microconchids en crust ing pos te rior part (ar rowed) of shell of Modiolus bi valve, lower Hinton For mation. D. Microconchids (ar rowed), de tached from their substrate and pre served loose in sed i ment to gether with abun dant ostracode car a paces, lower Hinton For mation. Im ages of spec i mens, coated with ammo nium chlo ride gyrogonites in some hori zons, and the lack of a strictly ma- Group. The presence of Microconchus hintonensis sp. nov. rine fauna in di cate that the siliciclastic de pos its of the lower in other forma tions, such as the Princeton Forma tion or Hinton, Princeton and Bluestone Forma tions were depos - within the marine units of the up per Hinton Forma tion ited in fresh-wa ter envi ron ments. It is very likely that the (Eads Mill Member), cannot be confirmed, owing to the palaeoenvironment took the form of terres trial flood plains, poor state of preser va tion of the speci mens. where overbank flood depos its left swamps and ephem eral Obser va tions of plant remains (shoots and leaves) and lakes. The tan Carbonicola bed (local ity 10), includ ing bi valves within the for ma tions in ves ti gated in di cate that ostracodes and charophyte gyrogonites, may repre sent such microconchids preferred both kinds of sub strates for at tach- a lac us trine en vi ron ment. Sim i larly the lam i nated mud- ment and later growth. It is not surpris ing, as the presence of stones with Hemicycloleaia bran chi o pods and bi valves (lo- Carbon if er ous microconchids on such types of substrate is cality 8) may repre sent overbank flood depos its. well known in the liter a ture (e.g., Trueman, 1942; Masta- The Co ney Limestone of the Bluefield Forma tion, con- lerz, 1996; Aitkenhead et al., 2002; Falcon-Lang, 2005; Za- tain ing microconchids, as well as fish teeth and scales, ostra- toñ and Mazurek, 2011; Florjan et al., 2012). A great num- codes and a sporadic marine fauna (Reger, 1926; Stencil, ber of spec imens also occur as detached tubes, free of sub - 2012), presum ably was depos ited in brackish waters. The strate (Fig. 9D). Those tubes, occur ring near bivalves, may presence of the abundant ostracode, Whipplella, in the Coney indi cate that the microconchids occu pied shells, but were Lime stone in di cates a tran si tional shore line-nearshore en vi- later detached, owing to, e.g., disso lu tion of the aragonitic ron ment with par tial car bon ate-form ing con di tions (Tibert shell. The pres ence of char ac ter is tic de pres sions from and Dewey, 2006), yet the ap pear ance of paraparchitaceans microconchid tubes on the bi valve molds (Fig. 9C) supports and kloedenellaceans suggests this may have been a shore- this sce nario. Microconchid tubes also may have detached, line tran si tional en vi ron ment (Sten cil, 2012). There fore, it is follow ing the degra da tion of the land plant remains, to ev ident that the spe cies Microconchus hintonensis sp. nov. which they were orig inally at tached during life. The preser - was able to thrive prolif i cally in fresh-water, and possi bly vation of plant frag ments on the tube bases of some of the also in brack ish-wa ter (Co ney Lime stone) con di tions dur ing spec imens may support this statement. However, the large sed i men ta tion of the Up per Mis sis sip pian Mauch Chunk num bers of microconchid tubes, dispersed in the depos its of NON-MA RINE MICROCONCHID TUBEWORM, LOWER CARBONIFEROUS 47 the Bluefield, lower Hinton and Bluestone Forma tion, away Ta ble 2 from any land plant re mains, is inter est ing. In this case, it is inter preted that the tubes either fell off dissolved bivalve Number of microconchids shells and/or, al ter na tively, be came de tached from algal en crust ing par tic u lar bi valve shells fronds, on which they may have origi nally been encrusted. It is gen er ally assumed that the pres ence of microconchid Bivalves Microconchid # tubes, detached from substrates, is an indi ca tion that they Modiolus sp. 1 origi nally encrusted hard, but aragonitic sub strates that dis- Modiolus sp. 2 solved, or organic substrates (soft-bodied taxa) that degra- Modiolus sp. 1 ded, leaving no trace (e.g., Vinn and Taylor, 2007; Zatoñ Modiolus sp. 8 and Krawczyñski, 2011b). The hypoth e sis of microcon- Carbonicola sp. 3 chids settling on algae is very possi ble, as the preser va tion Carbonicola sp. 1 of microconchid-encrusted al gal thalli is also known from Carbonicola sp. 4 the Upper Devo nian of Germany (Jux, 1964). The presence of gyrogonites in some hori zons may indi cate that at least Carbonicola sp. 2 some of the algae, serving as a substrate for microconchids, Carbonicola sp. 3 were repre sented by charophytes. Carbonicola sp. 3 The statement of Trueman (1942) about possi ble com- myalinid 2 mensalism be tween Carbon if er ous microconchids and some myalinid 1 non-marine bivalves is diffi cult to support and the authors do not subscribe to this hypoth e sis. Instead, it is believed that microconchids, as op por tu nis tic or gan isms, col o nized turbances, in the form of either higher sedi ment input or any suit able, firm and hard sub strate in a given envi ron - com pe ti tion with other en crust ing or gan isms. How ever, as ment. Currently, there are no data support ing the idea that the higher sed iment in put may be supported indi rectly by cer tain microconchid spe cies pre ferred a spe cial kind of the presence of land plant fragments, the com peti tion hy- substrate. Recently pub lished data from the marine Upper pothe sis is diffi cult to prove, because of the lack of evi - Devo nian (Zatoñ and Krawczyñski, 2011a; Zatoñ and dence. Borszcz, 2013) indi cated that microconchids col o nized On the basis of distinct signs of tube regen er a tion (Fig. brachio pod shells, since they served as an unlim ited source 4E, G), it may be concluded that some microconchids in a of hard substrate in the en viron ment. More over, no prefer - given popu la tion may have suffered from attempted preda - ences of microconchid set tle ment con cern ing par tic u lar tion by other ani mals, proba bly some kind of small arthro - sites on the brachio pod shells, have been detected. There - pods or fish. Proba bly while trying to catch the tenta cles, a fore, in the case of the Upper Missis sip pian microconchids po tential preda tor must have injured the tube along with the studied, it is suggested that Microconchus hintonensis sp. tube-se cret ing ep i the lium, caus ing a de vi a tion in its growth. nov. set tled on a va riety of suit able substrates, present in the Sim ilar devi a tions in tube growth, related to failed preda - soft-bot tom en vi ron ments, rang ing from al gae, trans ported tion, were noted in cornulitids (Vinn, 2009), which were land plants and bivalve shells. Thus, com peti tion for space close rel atives of the microconchids. There fore, pre da tion among these organ isms must have been mini mal or non-ex- on simi larly small-sized, tube-dwelling organ isms, such as istent. This may be supported by the fact that very few microconchids, must have occurred as well and, im por- microconchids are present, with respect to a given inspected tantly, was not lethal to all indi vid u als in a given popu la tion. plant fragment or bivalve shell (Table 2). This prob lem, how ever, awaits fur ther in ves ti ga tion. The microconchids in ves ti gated are rep re sented by three mor pho log i cal types, adapted to cer tain, eco log i cal and en vi ron men tal con di tions (Vinn, 2010): 1) planispiral CON CLU SIONS com pletely substrate-ce mented tubes; 2) planispiral tubes with ele vated aper tures, and 3) loosely coiled, soli tary The inves ti gated depos its of the Bluefield, lower Hin- tubes. Unlike morpho log i cal type 1, types 2 and 3 include ton and Bluestone Forma tions of the Upper Missis sip pian speci mens, growing on a hard substrate in the envi ron ment Mauch Chunk Group of the southern part of the West Vir- and ex pe ri enc ing at least pe ri odic dis tur bances. In the case ginia, USA re vealed the presence of abundant microcon- of type 2, the el eva tion of the tube ap erture was an escap ing chids, assigned to a new species Microconchus hintonensis. response to overgrow ing by neighbour ing encrusters. Type The new species shows con sider able vari abil ity in tube coil- 3 is char ac ter is tic of en vi ron ments, where faster sed i men ta- ing and or namen ta tion pattern, a feature known also in other tion rates forced the microconchids to grow upwards, result - described, microconchid species. Microconchus hintonen- ing in heli cal uncoil ing of their tubes (see also Burchette sis sp. nov., as an oppor tu nis tic, highly fecund spe cies, inha- and Rid ing, 1977). It must be noted, however, that morpho - bited in large numbers fresh-water (lower Hinton and Blue- logi cal type 3 may have also resulted from compe ti tion with stone For ma tions), and pos si bly also brack ish-wa ter (Co ney other soft-bodied encrusters (ani mals or algal cover) which Lime stone, Bluefield For ma tion) hab i tats, as ev i denced by were not fos silised. The pres ence of these three tube mor- fos sil as so ci a tions, com pris ing land plant re mains, charo- phologies in the microconchids studied indi cates that gener - phyte gyrogonites, freshwa ter bivalves, conchostracans, ally calm en vi ron men tal con di tions were in ter rupted by dis- ostracodes, as well as fish teeth and scales. There, micro- 48 M. ZATOÑ & R. L. PECK

conchids have been found, encrust ing bi valve shells and grams, 35: 600. land plant remains, although a large number of speci mens Cecil, C. B., Brezinski, D. & DuLong, F., 2004. The Pa leo zoic re - were also found loose in the host sedi ments. The latter spe- cord of changes in global climate and sea level: Cen tral Ap pa - cimens may have orig inally encrusted bivalve shells and la chian Ba sin. In: Southworth, S. & Bur ton, W. (eds), Geo- plants (includ ing al gal thalli, as well), but fell off the sub- logy of the National Cap i tal Re gion – Field Trip Guide book. U.S. Geo log i cal Sur vey Cir cu lar 1264: 77–135. strates, after disso lu tion of the aragonitic shells and deg ra- Chesnut, D. R., Jr. & Greb, S. F., 2009. In tro duc tion. In: Greb, S. dation of the plant remains. The oc currence of many speci - F. & Chesnut, D. R., Jr. (eds), Car bon if er ous of the Ap pa la - mens with heli cally uncoiled tubes may in di cate that they chian and Black War rior Basins . Ken tucky Geo log i cal Sur- responded with ver ti cal tube growth to epi sodic, high rates vey, Spe cial Pub li ca tion 10, Se ries XII: 3–9. of sed i men ta tion and/or more in tense com pe ti tion with Cooper, B. N., 1948. Sta tus of Missis sip pian stra tig ra phy in the other, soft-bodied encrust ing organ isms. The healed inju - cen tral and north ern Ap pa la chian re gion. The Journal of Ge - ries, marked in the tube by devi a tion in its growth, clearly ol ogy, 56: 255–263. indi cate that these microconchids also witnessed some pre- Cooper, B. N., 1961. Grand Ap pa la chian Ex cur sion. Virginia dation pressure from other ani mals in the envi ron ment. Poly tech nic In sti tute En gi neer ing Ex ten sion Se ries, Geo log i - Although microconchids are also present in other for- cal Guide book I, Blacksburg, 187 pp. mations, in cluding the marine inter vals of the up per Hinton Elias, M. K., 1957. Late Mis sissip pian fauna from the Redoak Forma tion (Eads Mill Member) and Bluefield Forma tion, Hol low Forma tion of south ern Oklahoma, Part I. Jour nal of their assign ment to the species Microconchus hintonensis Pa le on tol ogy, 31: 370–427. Englund, K. J., 1979. Mis sissip pian System and lower se ries of the sp. nov. is uncer tain, owing to the poor preser va tion of spe- Penn syl va nian Sys tem in the pro posed Penn syl va nian Sys tem cimens, which are attached to brachio pod shells. stratotype area. In: Englund, K. J., Arndt, H. H. & Henry, T. Fi nally, this study shows that bulk sam ple mac er a tion W. (eds), Pro posed Penn syl va nian Sys tem Stratotype, Vir - and sieving is a very prom ising method for microconchid ginia and West Vir ginia. Amer i can Geo log i cal In sti tute, Se- extrac tion from depos its, where any firm or hard substrate, lected Guide book Se ries No. 1, Guide book for Ninth Inter- in the form of plant re mains and ani mal shells respec tively, na tional Con gress of Car bon if er ous Stra tig ra phy and Ge ol - has been degraded and dissolved. ogy Field Trip No. 1: 69–72. Ettensohn, F. R., 2009. The Mis sissip pian of the Appa la chian Ba - sin. In: Greb, S. F. & Chesnut, D. R., Jr. (eds), Car bon if er ous Ac knowl edge ments of the Ap pa la chian and Black War rior Basins . Ken tucky Geolog i cal Sur vey, Spe cial Pub li ca tion 10, Se ries XII: 22–31. We would like to thank Mark Wil son (Ohio, USA) and Olev Etheridge, R., 1880. A con tri bution to the study of the British Car - Vinn (Tartu, Esto nia), the jour nal ref er ees, for their con struc tive bon if er ous tubicolar Annelida. Geo log i cal Mag a zine, 7: 258– re views and the jour nal ed i tors for their care ful check ing of the fi - 266. nal ver sion of the manu script. Falcon-Lang, H. J., 2005. Small cordaitalean trees in a marine-in - flu enced coastal hab i tat in the Penn syl va nian Joggins For mation, Nova Sco tia. Jour nal of the Geo log i cal So ci ety, Lon don, REF ER ENCES 162: 485–500. Florjan, S., Pacyna, G. & Borzêcki, R., 2012. First find of mi cro- Aitkenhead, N., Barclay, W. J., Brandon, A., Chadwick, R. A., conchids (Tentaculita) on Up per Car bon if er ous seed fern Ka- Chis holm, J. I., Coo per, A. H. & John son, E. W., 2002. The rinopteris daviesii from Nowa Ruda (Lower Silesia, Po land). Pennines and Ad ja cent Ar eas (4th edi tion). Brit ish Re gional Przegl¹d Geologiczny, 60: 273–275. [In Pol ish, Eng lish sum- Ge ol ogy Se ries. Brit ish Geo log i cal Sur vey, Keyworth, 206 mary]. pp. Fraiser, M. L., 2011. Paleoecology of sec ondary tierers from Be³ka, Z. & Skompski, S., 1982. A new open-coiled gas tro pod West ern Pangean trop i cal ma rine en vi ron ments dur ing the af - from the Viséan of Po land. Neues Jahrbuch für Geologie und ter math of the end-Perm ian mass ex tinc tion. Palaeo ge ogra - Paläontologie, Monatshefte, 7: 389–398. phy, Palaeoclimatology, Palaeo ec ol ogy, 308: 181–189. Beuthin, J. D. & Blake, B. M., 2004. Revised stra tig ra phy and no - Gordon, M. & Henry T. W., 1981. Late Mis sissip pian and Early mencla ture for the Up per Hinton For mation (Up per Mis sis- Penn syl va nian in ver te brate fau nas, east-cen tral Ap pa la chians sippian) based on rec ogni tion of re gional ma rine zones, – a pre limi nary re port. In: Rob erts, T. G. (ed.), GSA Cincin- southern West Vir ginia. South east ern Ge ol ogy, 42: 165–178. nati ‘81 Field Trip Guide books, Vol. 1: Stratig raphy, Sedi- Branson, C. C., 1937. Stra tig ra phy and fauna of the Sacajawea mentology. Geo log i cal So ci ety of Amer ica: 165–171. For ma tion, Mis sis sip pian, of Wy o ming. Jour nal of Pa le on - He, L., Wang, Y., Woods, A., Li, G., Yang, H. & Liao, W., 2012. tol ogy, 11: 650–660. Calcar e ous tubeworms as disas ter forms after the end-Perm- Burchette, T. P. & Rid ing, R., 1977. At tached ver miform gastro - ian mass ex tinc tion in south China. Palaios, 27: 878–886. pods in Car bonif er ous marginal ma rine stromatolites and bio- Henry, T. W. & Gordon, M., 1979. Late De vonian through Early stromes. Lethaia, 10: 17–28. Perm ian (?) in ver te brate fau nas in pro posed Penn syl va nian Caruso, J. A. & Tomescu A. M. F., 2012. Microconchid encrusters Sys tem stratotype area. In: Englund, K. J., Arndt, H. H. & col o niz ing land plants: the ear li est North Amer i can re cord Henry, T. W. (eds), Pro posed Penn syl va nian Sys tem Stra- from the Early De vonian of Wyo ming, USA. Lethaia, 45: totype, Virginia and West Virginia . 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