The epiplanktic anthozoan, Kolihaia eremita Prantl, 1946 (Cnidaria

Journal of the Czech Geological Society 46/3ñ4(2001) 239

The epiplanktic anthozoan, Kolihaia eremita Prantl, 1946 (Cnidaria),from the Silurian of the Prague Basin (Bohemia)

EpiplanktonnÌ kor·l, Kolihaia eremita Prantl, 1946 (Cnidaria),ze siluru praûskÈ p·nve (»echy)

(3 figs, 2 plates)

JIÿÕ KÿÕé 1 ñ JIÿÕ FR›DA 1 ñ ARNOäT GALLE 2

1 Czech Geological Survey, Kl·rov 3/131, 118 21 Praha 1, Czech Republic; e-mail: [email protected], [email protected] Geological Institute, Academy of Science of the Czech Republic, Rozvojov· 135, 165 00 Praha 6-Suchdol; e-mail: [email protected]

New material makes it possible to classify the epiplanktic species Kolihaia eremita Prantl, 1946, supposed to be a worm or cornulitid,from the Silurian of the Prague Basin as a member of the phylum Cnidaria and a possible member of Rugosa or Tabulata.

Key words: Silurian, Prague Basin, Bohemia, Cnidaria, Rugosa?, Tabulata?

Silurian strata of the Prague Basin, Bohemia. Prantl (1946)placed this ecologically interesting new genus and speciesin the phylum Annelida Lamarck, 1809 (Serpulidae Bur-meister, 1837). Fischer (1962, 1966) included the genusKolihaia Prantl, 1946 within the family Cornulitidae Fis-cher, 1962 with uncertain phylum, class and order rela-tionship together with the genera Cornulites Schlotheim,1820, Conchicolites Nicholson, 1872 and CornulitellaHowell, 1952. In 1990 Turek considered Kolihaia to beof uncertain affinity and supported the idea of its epiplank-tic mode of life in agreement with Prantlís original inter-pretation on the base of new observations in the field.

In 1975 one of us (J. K.) collected more than 1000specimens of Kolihaia eremita in the upper Wenlock(Monograptus belophorus Zone) at a new locality inKonÏprusy near Beroun (private property 200 m SW ofthe village cemetery) and later also at other localities. Thevery good preservation of the material makes it possibleto contribute to the systematic classification of the ge-nus Kolihaia.

Systematic paleontology

Genus Kolihaia Prantl, 1946

1946 Kolihaia gen. nov.; Prantl, p. 1ñ3.1962 Kolihaia Prantl, 1944; Fischer, p. W138.1966 Kolihaia Prantl, 1944; Fischer, p. W138.1992 Kolihaia Prantl, 1946; Gnoli, p. 383.

Ty p e s p e c i e s : Kolihaia eremita Prantl, 1946.

D i a g n o s i s : Solitary, loosely dendroid or gregarious,relatively small animal with almost regularly annulatedvery thin conical calcareous corallites, usually curved atits proximal part. In the proximal and also distal partscharacteristic, relatively long, thin, hollow and mostlybranched radiciform processes are developed. Internalfeatures most probably reduced to minimum due to theepiplanktic mode of life. Individual corallites increaselaterally by production of offsets.

Introduction

For more than two hundreds years zoologists have usedan artificial system for a classification of the animal king-dom. Amongst the many categories used for classifica-tion of species the phylum represents the most satisfac-tory taxa. There is a good general consensus about therank of phylum which should include all those animalssharing a particular body plan. Modern zoology has reco-gnised around 32ñ36 phyla in which all extant animalsmay be placed. However, our interpretation of the phy-lum-level position of extinct animals is much more dif-ficult because it is considerably limited by the loss ofmany characters during the fossilisation processes. Com-monly only hard body parts like skeleton or shell havebeen preserved. For this reason our interpretation of thehigher taxonomic position of any extinct animal is basedon a comparison of the preserved hard body parts withthose of extant animals or on their functional analysis.These limitations make the placement of fossils into thecategories used by zoologists much more difficult (some-times even impossible) than that of the living animals.In addition, many animal groups of the order or class lev-el are extinct and it is very probable that even somegroups of the phylum rank are extinct. The placement ofsome fossils in one of the extant phyla is not always theonly possible solution of their taxonomic position. Thus,a determination of the higher taxonomic position in manyextinct animals has to be considered as ìthe best solu-tionî based on the present level of our knowledge.

Change of mode of life is typically connected with aloss or reduction of some body structures and with thedevelopment of new body structures. In such cases a cor-rect interpretation of the ancestral body plan and subse-quent placement of the animal into any phylum is some-times difficult even in living animals. Determination ofcorrect taxonomic position in fossil animals which wereadapted to a quite different mode of life than their ances-tors is often a serious problem. The present paper is fo-cused on one of such fossils, Kolihaia eremita, from the

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240 Journal of the Czech Geological Society 46/3ñ4(2001)

R e m a r k s : Prantl (1946) related Kolihaia to the phy-lum Annelida Lamarck, 1809 (Serpulidae Burmeister,1837) and Fischer (1962, 1966) included the genus Ko-lihaia Prantl, 1946 within the family Cornulitidae Fis-cher, 1962 with uncertain phylum, class and order rela-tionship together with the genera Cornulites Schlotheim,1820, Conchicolites Nicholson, 1872 and CornulitellaHowell, 1952. According to Richards (1974) Cornuliti-dae represent a family of extinct worm-like organisms ofunknown biologic affinities showing a commonly epi-zoic mode of life on other fossils as solitary commen-sals, gregarious commensals, unattached cornulitids andparasitic cornulitids. When living as commensals they arecemented to the substrate by the proximal part of thetubes directly and absorb calcium carbonate partly fromthe host (Fischer 1962, 1966). Their vesicular wall (Bath-er 1923) is similar to calcareous hydroids, stromatopo-roids and some fusulines (Fischer 1962, 1966)

Kolihaia developed characteristic, relatively long, thin,hollow and mostly branched radiciform processes (Fig. 1,Pl. I/1, 4, 5, 6, Pl. II). On the other hand, no annelid canproduce similar radiciform processes in the proximal partof the tube. The shape of the tube and similar processesmay be expected in some cnidarians. Hydroidea are ex-cluded since polyps do not bud from an existing polypand new buds/offsets develop from the supporting sto-lons or stems of the polyps only (Sch‰fer 1972). Rugosaand Tabulata (Cnidaria, Anthozoa) represent the onlygroups of cnidarians characterized by increase of origi-

nal protocorallites by offsets produced laterally (Hill1956, 1981; Sokolov 1962). The corallites of Rugosa aregenerally characterized by complex internal features ofskeleton, mainly radial plates and tabulae (tabularium) ordissepiments (dissepimentarium) (Hill 1956). Rugosa areboth solitary and colonial. On the other hand, Tabulataoften lack both vertical and horizontal inner structuresbut are usually colonial.

Corallites of Kolihaia are extremely thin-walled andpreserved fully compressed. We were unable to observeany internal features between the compressed walls of thecorallites. The only massive structures are the radiciformprocesses in the proximal parts of the corralites. Theradiciform processes formed by carbonate are hollow andgrow directly from the wall (Fig. 1, Pl. I/1, 4, 5, 6, Pl. II).They are branching, circular in cross-section and theirdiameter distally decreases continually.

The existence of radiciform processes in the proximalparts of Kolihaia is rather puzzling. Short blunt rootletsare known in Rugosa but rugosans never lack inner struc-tures as often as Tabulata. Tabulates, on the other hand,are unknown to develop radiciform processes. In ouropinion, such processes could have developed as an ada-ptation to their epiplanktic mode of life, either in thesame way as the rootlets of Rugosa, or possibly from theconnecting structures. In any case, we suggest that theincrease of the corallites by offsets produced laterallyrelates the genus Kolihaia to the phylum Cnidaria, andan absence of internal structures such as tabulae or sep-

Fig. 1 Colony of Kolihaia eremita Prantl, 1946, from the upper Wenlock rocks (Monographus belophorus Zone) at KonÏprusy near Beroun.Specimen JK 8383, x5.

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ta and probably also mural pores demonstrates mostprobably a relationship to Tabulata, probably Aulopori-da. The absence of the internal features of the corallitein Kolihaia may be the result of the adaptation to an epi-planktic mode of life, where the maximal reduction ofthe weight/mass is one of the important features. We con-sider the presence of the conspicuous radiciform proces-ses, unknown in other Tabulata or Rugosa, as a furtheradaptation to an epiplanktic life as the processes mayhave served to attach the colony to floating or buoyantobjects, probably algae.S p e c i e s : Kolihaia eremita Prantl, 1946 (Silurian of thePrague Basin, Bohemia) and Kolihaia sardiniensis Gno-li, 1992 (Lower Silurian of the SW. Sardinia).

Kolihaia eremita Prantl, 1946Figs 1ñ3, Plates IñII

1946 Kolihaia eremita sp. nov.; Prantl, p. 3ñ7, 9ñ13, Pl. 1.1962 Kolihaia eremita Prantl, 1944; Fischer, p. W138, Fig. 78/4.1966 Kolihaia Prantl, 1944; Fischer, p. W138, Fig. 78/4.1990 Kolihaia eremita; Turek, p. 345ñ346, Pl. 8, Figs 1, 2.

H o l o t y p e : Specimen figured by Prantl, 1946 on Pl. 1, Fig. 1, re-figured by Fischer, 1962 and 1966 on Fig. 78/4 and deposited inthe National Museum, Prague under No. 22 129/9.

P a r a t y p e s : Other specimens figured by Prantl, 1946 on Pl. 1.T y p e h o r i z o n : Motol Formation.T y p e l o c a l i t y : KonÏprusy near Beroun.O c c u r r e n c e : Prague Basin, Bohemia, Praha-Mal· Chuchle

(VyskoËilka) ñ upper Motol Formation, M. riccartonensis Zone;LodÏnice (»ernidla), KonÏprusy, 200 m SW from the cemetery,

Vöeradice ñ Monograptus belophorus Zone, Motol Formation;Bykoö near Zdice, Testograptus testis Subzone; Kosov Quarry nearKr·l˘v Dv˘r ñ C. lundgreni Subzone; Praha-Stod˘lky ñ S. chimaeraZone.

D e s c r i p t i o n : The description of the species is giv-en by Prantl (1946). Only the morphological featuresconsidered important for interpretation of its taxonomicposition as well as newly observed features are men-tioned below. Characteristic radiciform processes aredeveloped in the proximal part of the conical calcareouscorallites and are more common on their convex side.The first processes occur typically about 3ñ5 mm fromthe apex of the corallite. Radiciform processes are hol-low with an inner diameter slightly more than one halfof the process width (Pl. II/3). The processes arebranched irregularly and dichotomously and their lengthreaches up to half of the corallite length. Individual cora-llites increase laterally by production of offsets.D i m e n s i o n s : Length of the corallites up to 25 mm,maximum width of the compressed corallite at distal endis up to 5 mm, width of rootlets about 0.2 mm.N o t e : Pronounced annulations of the corallite walls arethe most conspicuous feature of Kolihaia eremita. Theyresemble rejuvenations known in Rugosa but they differin their regularity. Thick annulations are usually arrangedin pairs: after the wall withdrawal toward the coralliteaxis the diameter abruptly increases into two annulationsclose to each other. It seems to contradict the cnidarianaffinities of Kolihaia.

Fig. 2 Colonies and single corallites of Kolihaia eremita covering bedding plane of the Silurian shale (Monographus belophorus Zone at KonÏ-prusy near Beroun; x5.

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R e l a t i o n s h i p s : Gnoli (1992) described the new spe-cies Kolihaia sardiniana from the lowermost Ludlow ofSW Sardinia. Its comparatively very small corallite hasa similar general shape to Kolihaia eremita but distinct-ly differs in the type of the processes which are regular-ly distributed along the corallite, probably due to adap-tation of the animal to a benthic mode of life.P a l e o e c o l o g y : Kolihaia eremita single corallitesand colonies are preserved in a black shale facies of theSilurian scattered or covering one or several beddingplanes in the sequence (Fig. 2), occuring mainly togeth-er with graptolites and rare single valves of brachiopods(Niorhynx, Valdaria). They were obviously fossilised inan abiotic environment. Prantl (1946) and Turek (1990)supported the idea of an epiplanktic life based on thepresence of unbroken branching radiciform and fragileprocesses at the proximal part of the individuals andwhole colonies and because of their occurrence in theabiotic environment of the black graptolitic shales. Thebranching processes most probably assisted in anchoringthe individuals and colonies in the algal clumps floatingin the ventilated upper parts of the water column. Therock surface in the vicinity of the radiciform processesis usually whitened by reduction processes when frag-

ments of organic matrix (algae and body softparts) on them decayed during fossilisation(Fig. 1, Pls IñII). The same whitening is welldemonstrated also on the distal ends (body softparts) of the corallites around the aperture (Fig. 1;Pl. I, Fig. 2). Another feature supporting an epi-planktic mode of life is the minimal weight of thetubes (corallites) without internal features.

Conclusion

Interpretation of the Silurian Kolihaia eremita asa member of the class Anthozoa with an epiplank-tic mode of life completes our view on the lifestrategy of the fossil members of the phylum Cni-daria. The latter phylum forms a clearly definedanimal group (by formation of their unique cni-doblast) with a very long fossil record. There isevidence for hydrozoans, pennatulid-like antho-zoans, and primitive scyphozoans even in the Pre-Cambrian strata (Glaessner 1984). Cnidarianshave had a quite unique position in the zoologi-cal system and resolution of some basic, but stillunsolved questions regarding their phylo-genywould have an influence on the interpretation ofthe phylogenetic relationships of many animalphyla. One of such questions is the symmetry ofthe first Metazoa. Radiata uniting ìlowerî animalswith radial symmetry have been considered torepresent more primitive animals than Bilateriauniting Metazoa with a bilaterally symmetricalbody. For this reason, radial symmetry is tradi-

tionally considered to be the body symmetry of the firstmetazoans. On the other hand, some analyses of symme-try in animals has suggested that radiality is a second-ary feature and so is not a symmetry of ancestral Meta-zoa (Willmer 1996 and reference therein). Cnidarians area metazoan group uniting animals with both body sym-metries, bilateral Anthozoa (disposition of mesenteries,muscles and siphonoglyphs) and radial Hydrozoa (Will-mer 1996). Resolution of the question whether evolutionhas been from hydrozoan ancestors to later anthozoansor vice versa is of crucial significance for our understand-ing of the evolution of the early Metazoa. For a solutionto the latter question we have to analyze all the availabledata on the phylogeny of the Cnidaria. The present pa-per describing an unknown life strategy of the SilurianAnthozoa represents a small contribution to this subject.

Acknowledgement. Many thanks to K. Histon, from theGeologische Bundesanstalt, Vienna for her help with thecritical reading of the manuscript and for language cor-rections. This publication is based on the work partlysponsored by grant 205/01/0143 from the Grant Agencyof the Czech Republic.

Submitted January 10, 2001

Fig. 3 Single corallite of Kolihaia eremita from the Monographus belophorus Zone atKonÏprusy near Beroun; A ñ specimen JK 8393a, x6, B ñ specimen JK 8383, x11.

A B

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EpiplanktonnÌ anthozon, Kolihaia eremita Prantl, 1946 (Cnidaria), ze siluru praûskÈ p·nve (»echy)

Nov˝ materi·l umoûÚuje p¯i¯adit epiplanktick˝ druh Kolihaia eremita Prantl, 1946, povaûovan˝ za Ëerva nebo kornulitida, ke kmeni Cnidaria apravdÏpodobnÏ k Rugosa nebo Tabulata.

References

Bather, F. A. (1923): The shell of Cornulites. ñ Geol. Mag., London, 60:542ñ545.

Fischer, D. W. (1962): Small conoidal shells of uncertain affinities, W98ñ W143. ñ In: Moore, R. C. (Ed.) Treatise on invertebrate paleontol-ogy, Part W, Miscellanea. ñ The University of Kansas and The Geo-logical Society of America, Inc., New York and Lawrence, Kansas,W1ñW259.

ñ (1966): Small conoidal shells of uncertain affinities, W98 ñ W143. ñIn: Moore, R. C. (Ed.) Treatise on invertebrate paleontology, Part W,Miscellanea. ñ The University of Kansas and The Geological Soci-ety of America, Inc., New York and Lawrence, Kansas, W1ñW259.

Glaessner, M. F. (1984): The Dawn of Animal Life. A biohistorical study.ñ Cambridge University Press.

Gnoli, M. (1992): The problematic organism Kolihaia sardiniensis sp. nov.of the latest Wenlock ñ earliest Ludlow of SW Sardinia. ñ Boll. Soc.Paleont. Ital., Modena, 31(3): 383ñ385.

Hill, D. (1956): Rugosa, F233 ñ 245. ñ In: Moore, R. C. (Ed.) Treatise oninvertebrate paleontology, Part F, Coelenterata. ñ The University of

Explanations of plates

P l a t e IKolihaia eremita Prantl, 1946, KonÏprusy near Beroun, Bohemia, Motol Formation, Wenlock, Sheinwoodian, Monograptus belophorus Biozone:1 ñ detail view of specimen JK 8369 showing branched radiciform processes, x20.2 ñ detail view of specimen JK 8383 showing whitening around the aperture, x10.3 ñ specimen JK 8376a, x25.4 ñ detail view of corallite, specimen JK 8372, x20.5 ñ detail view of specimen JK 8383a showing branched radiciform processes, x30.6 ñ specimen JK 8390a, showing an apex of the corallite, x20.

P l a t e I IKolihaia eremita Prantl, 1946, KonÏprusy near Beroun, Bohemia, Motol Formation, Wenlock, Sheinwoodian, Monograptus belophorus Biozone:1 ñ specimen JK 8388, x40.2 ñ detail view of specimen JK 8369 showing whitening around radiciform processes, x10.3 ñ detail view of branched radiciform process, specimen JK 8379a, x45.4 ñ specimen JK 8393a, x25.5 ñ specimen JK 8370, x30.

Kansas and The Geological Society of America, Inc., New York andLawrence, Kansas, F1ñF498.

ñ (1981): Rugosa and Tabulata. ñ In: Teichert, C. (Ed.) Treatise oninvertebrate paleontology, Part F, Supplement 1. ñ The GeologicalSociety of America, Inc. and The University of Kansas, Boulder,Colorado and Lawrence, Kansas, F1ñF762.

Prantl, F. (1946): Kolihaia eremita gen. et sp. nov. (Annel. Tubicola) zest¯edoËeskÈho siluru (Kolihaia eremita nov. gen., nov. spec. a newTubicolar Annelide from the Silurian of Bohemia). ñ VÏst. Kr·l. »es.SpoleËn. Nauk, T .̄ mat.-p¯ÌrodovÏd., Praha, (24): 1ñ12.

Richards, R. P. (1974): Ecology of the Cornulitidae.ñ J. Paleont., Tulsa,48(3): 514ñ 523.

Sch‰fer, W. (1972): Ecology and palaeoecology of marine environments.ñThe Univ, of Chicago Press, Chicago, 1ñ568.

Sokolov, B. S. (1962): Gubki, archeociaty, kishechnopolostnye, chervi. ñIn: Orlov, Yu. A. (1962) Osnovy paleontologii, Moskva, 1ñ430.

Turek, V. (1990): Comments to upper Wenlock zonal subdivisions in theSilurian of Central Bohemia. ñ »as. Min. Geol., Praha, 35(4): 337ñ353.

Willmer P. (1996): Invertebrates relationships ñ patterns in animal evolu-tion. ñ Cambridge University Press, 400 pp.

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1

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5 6

J . K ¯ Ì û ñ J . F r ˝ d a ñ A . G a l l e : The epiplanktic anthozoan, Kolihaia eremita Prantl, 1946 (Cnidaria), from the Silurian of the Prague Basin(Bohemia) (Pl. I)

For explanation see p. 243

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1

2

3

4 5

J . K ¯ Ì û ñ J . F r ˝ d a ñ A . G a l l e : The epiplanktic anthozoan, Kolihaia eremita Prantl, 1946 (Cnidaria), from the Silurian of the Prague Basin(Bohemia) (Pl. II)

For explanation see p. 243

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The epiplanktic anthozoan, Kolihaia eremita Prantl, 1946 (Cnidaria