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Substrate Specificity in the Devonian Tabulate Coral Pleurodictyurn

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Substrate Specificity in the Devonian Tabulate Coral Pleurodictyurn Substrate specificity in the Devonian tabulate coral Pleurodictyurn CARLTONE. BRETT AND JOHN F. COTTRELL Brett, Carlton E. & Cottrell, John F. 19820815: Substrate specificity in the Devonian tabulate coral LETHAIA pleurodicryum. Leihain, "01.15. pp 247-262. o110 ~ssN0024-1164. The tabulate coral Pleurodiciyum nmericonum Roemer has been cited as an example of a host-specific organism occurring exclusively on the shells of gastropods, particularly Paloeozvsopkuro homihonioe (Hall). Examination of over 16W specimens of P. omericonum, from the Middle Devonian Hamilton Group of wcrtcrn New York, reveals additional cornplcriticr which require reinterpretation. While substrate selectivity for Palaeoiygopleuro shells is evident in all 42 subsamples, a variety of other substrates were also utilized by Pleurodictyum including corals, brachiopods, other molluscs and pebbles. Recent rcleractinian corals inhabiting soft bottoms show similar substrate preference, selecting for the tubes of live serpulids, or gastropod shells (invariably with a secondary sipunculid host), but also occasionally settling on unoccupied shells or pebbles. Shell surfaces of P. harniltoniae, preserved as external molds on the Pieurodiciyum epitheca, exhibit cnciustation by worm tubes and bryozoans as well as boring5 and mechanical shell damage, suggesting that these were not ihc shells of live gastropods. However, the invariant aperture-downward orientation and thc high degrcc of selectivity of P. arneri- canurn strongly suggest that the shells were occupied by secondary hosts. O Subsrrarespecifi~iry,cornmen- mlirm, tobulore coral, garfropod, sipunculid, Devonian, Hamilton Group, New York. Corlron E. Brert ond John F. Cornell, Depcrimsnr of Geologicoi Sciences, the Univrrsiry of Rochester, Rochesrer, New York 14627; 15th March, 1981. Larvae of many marine invertebrates require substrate specificity in Pleurodictyum; however, hard, stable substrates for successful settlement it also reveals additional complexities which re- on the sea floor (Thorson 1950; Gray 1974). quire reinterpretation of the relationship. Many species exhibit nonrandom colonization of particular suhstrate types, including the skele- tons of other living host organisms (Scheltema Materials and methods 1974; Knight-Jones 1951; Crisp & Meadows 1967; and others). Substrate specificity can also Pleurodictyum is a small tabulate coral with a be demonstrated among fossil organisms in roughly-. discoidal corallum composed of relative- which the original attachment site is incorporated ly large, flaring, prismatic corallites and hearing into the adult skeleton or into the skeleton of a wncentricallv wrinkled basal eoitheca (Hill & another organism (Clarke 1908, 1921; Teichert Stumm 1957). The corallum exhibits a high de- 1945; Thayer 1974; Brett 1978; MacNamara gree of size and morphologic variability, ranging 1978; Conway Morris 1981). The Devonian tabu- from large hemispherical, flat-based forms, late coral Pleurodictyum has previously been which are generally the most abuhdant morpho- cited as an ancient organism showing suhstrate type, to conical and biconical forms, and irregu- specificity (Ager 1963). As early as 1908, John lar or aberrant morphotypes (Fig. 1). All forms M. Clarke noted the frequent association of this are intergradational and prohabiy represent eco- coral with particular -gastropods and inferred the phenotypic variants of a single species (Ross existence of a commensal relationship, in which 1953). Pleurodictyum americanum occurs ahnn- the coral larvae selected live snail hosts on which dantly in the Hamilton Group of New York to settle. While this suggestion has been much State, in a wide variety of lithologies. The corals quoted, the relationship has never been thor- are most abundant and largest in offshore medi- oughly documented and the hypothesis remains um to dark gray shales of western New York untested. Our detailed study of approximately (e.g. Wanakah and Windom members), but also 1600 specimens of Pleurodictyum americanum occur rarely in coarser clastic facies in the Hamil- Roemer from the Hamilton Group of western ton Group of eastern New York (e.g. King Fer- New York State supports Clarke's suggestion of ry). Pleurodictyum frequently occurs in sparsely 248 Carlton E. Bren and John F. Comell Pleurodictyurn occur in the Jaycox Member of the Ludlowville Formation and in the Kashong Shale Member of the Moscow Formation (Baird 1979). The highest stratigraphic occurrences of Pleurodictyum in western New York are in two richly fossiliferous zones of the Windom Shale in the upper portion of the Moscow Formation (Brett 1974; Baird & Brett, in press). A total of 42 large samples of Pleurodictyum were obtained from various horizons at 31 local- ities in western and west-central New York State (Fig. 3; Appendix). By far the largest samples were derived from the Wanakah Shale Pleurodic- tyum beds and the basal Windom Shale coral beds, both in Erie County. Specimens were pre- pared by boiling in Quaternary-0 for several hours and then treated with an ultrasonic clean- er. For most specimens in the soh shales of western New York this treatment served to re- move the matrix completely from basal surfaces of the corals and their attachment scars. After cleaning, latex casts were prepared of certain samples that showed attachment scars of Pleuro- dictyum. In three sites, where Pleurodictyum Fig. 1. Morphologic vanation in the corallum of Pleurodictyum specimens were particularly common, large cf. americonum Roemer. OA. Morphotype 1: planar base with blocks of matrix rock were collected and disag- nearly planar 'upper' orallite surface, BMS E 24955. OB. gregated in the lab. All contained fossils and Morphotype 2: planar base with convex upper surface, BMS E 24954. OC. Morphotype 3: convex base with planar upper their preservations were recorded to determine suriace, BMS E 24956. OD. Morphotype 4: convex base with the bulk composition of the Pleurodictyum-bear- convex upper surface, BMS E 24957. OE, F. Morphotype 5: ing fossil assemblages. All corals included in the specimens exhibiting irregular coralla, BMS E 24958. E 24959. present study are believed to represent a single B others x from Loc. 5A, all from Loc. 5B. AU views 1.2. species, Pleurodictyum americanum; however, further detailed taxonomic studies of the genus Pleurodictyum are needed. fossiliferous mudstones which lack rugose corals. It evidently was capable of inhabiting relatively soft mud bottoms which otherwise supported Substrates of Pleurodictyum only small brachiopods and mollusks. Indeed, Pleurodictyum has been cited as an example of The basal epitheca of Pleurodictyum commonly an organism adapted for soft substrates (Thayer incorporates the remains or impressions of ob- 1975). jects which provided initial attachment sites for Pleurodictyum is largely confined to several the corals (Fig. 4). Thus, in the majority of cases, thin but persistent horizons (Fig. 2; Cooper 1930, it is possible to identify the initial settlement 1957). The lowest horizon occurs at the base of substrate. A wide variety of hard substrates was the Ludlowville Formation in the transition be- utilized by Pleurodictyum including both skeletal tween the Centerfield Limestone and the Led- and inorganic materials such as phosphate peb- yard Shale (McCollum 1980). A very widespread bles and hiatus concretions (Fig. 4L, M). Over 60 unit termed the 'Pleurodictyum beds' by Grahau different skeletal substrates have been identified (1899) occurs at the base of the Wanakah Shale in our study (Tables 1 and 2). These include from Lake Erie to the central Finger Lakes re- mgose and auloporid corals (Fig. 4F), brachio- gion. Two or three other levels exist in the higher pods (Fig. 4C), and a variety of mollusks includ- units of the Wanakah and the laterally equivalent ing cephalopods, bivalves, and gastropods (Figs. King Feny Shale in the Seneca-Cayuga Lake 4A, B, E, G-I, K). Thus, statements which im- region (Baird 1981). Scattered specimens of ply that Pleurodictyum americanum occurs exclu- JO uo!$ualuoa aql ~ioddnsisal s!ql JO sqnsax .sa%e[q .sa~dmesy1nq pue alerpqns aql qaoq u! a8qq -masse a!io!q Ieg!u! aq, u!q!m spodo1lse8 asaql -massa l!ssoj aqllo siaqmam laqio lsu!r?8e av!uoJ JO aauepunqe Leu!ploelixa ue )aaUai d[dm!s -[!uvy vlnaldo8i(zoav~vd$0 aauepunqe aq) arud ~q%!rusa)eilsqns se v.inaldo8i(zoavlvd JO uoyod -moa 01 alqel Laua8u!luoa z x z e 8u!sn 'saldmes -old q%!q aql 'rahamoH .i(lyy!aads aleqsqns lsa2 aaiq) aq) $0 qaea 103 elep asaq uo pamiopad 4ns elep mo laadsai s!q) UI .alerasqns [elalays sem aauapuadapu! jo pa] ,X v .a%e~qmasseqlo!q JO sauads a[Zu!s uommoa Isom aql s! vrnaldo8 paleraosse aql jo % E ueq, ssa1 sasudmoa a! pay -i(zoav1vd 'saldmes lsom UI .Lalle~eu8!epueue~ -mexa suamuads un4xpodnald aq, jo %og ueql aql $0 aIeqs qeyemfi aql u! % 18 01 bun03 aux aiom u! alensqns Iuamqaene ue se srnaao vlnald $0 aleqs mopu!M a41 u! % LZ moq sa%uelvlnald -o2i(zoav~vdseaiaqm IE~Ismoqs 'huno3 aug -02zoav~vduo aauarrnaao 30 a8emaarad aq) 'uogeas yooig Bu~rd~ae apqS qeyeuefi lam01 'uozuoq a!qde@lens al8u!s e moq suam~aads aql 30 paq uni(rxpo.n~aldaqlloj elep xulem aq) unti03!polnald arolu lo 0s jo saldmes UI (,u!e%e pue aleilsqns aqljo uosuedmos e 'aldmexa rod aauo aep!)emauoxoT aqi q$!m sapads s!ql Bu!pn[a '(E alqe~)urnAj3!polnald $0 Sa)ellsqns mamqael -u! zuelrem Lem qalqm qauoaoloid aqljo s[!e)ap -,e 40 4!iolem aq) sasudmoa I! aiaqm sa!l!Ieao[ a~asaidel[eloa urnAj2!polnald lo aseq aql uo awes aql le sa8elqmasse pssoj pa~asaidIEIOI aq, splom
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