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Problems of functional morphology and classification in stylonuroid eurypterids (Chelicerata, Merostomata), with observations on the Scottish Silurian Stylonuroidea*

Published online by Cambridge University Press:  06 July 2012

Charles D. Waterston
Affiliation:
Royal Scottish Museum, Edinburgh

Synopsis

Eurypterids of the Superfamily Stylonuroidea Diener 1924 sensu Størmer (1974, 373) from the Pentland Hills, Midlothian, are redescribed and the evidence which these forms may give concerning the life environment of the Gutterford Burn Eurypterid Bed (Upper Llandovery), from which most of them have been obtained, is considered. Five species are recognised. Parastylonurus ornatus (Laurie) is redescribed with special reference to the organs of locomotion and reproduction. A new form from the Gutterford Burn is described as Parastylonurus hendersoni sp. nov. Stylonurus macrophthalmus Laurie is designated the type species of the new genus Hardieopterus and Stylonurus knoxae Lamont as the type species of the new genus Lamontopterus. The unique holotype of Lamontopterus knoxae is of morphological interest in showing evidence of the gut.

Stylonuroids from other Scottish Silurian localities are also considered. Two new species, Brachyopterella ritchiei sp. nov. from Seggholm and Hardieopterus (?) lanarki sp. nov. from the Logan Water are described. Stylonurella spinipes (Page) is redescribed and this species is reported for the first time from Seggholm.

The new information provided by this study has raised a number of problems of classification and the criteria upon which eurypterids are classified, and particularly those applied to the stylonuroid eurypterids, are re-assessed and a new classification of the Stylonuroidea proposed. This has required the description of new taxa; a new family, the Parastylonuridae, is proposed to accommodate Parastylonurus, Hardieopterus and probably Lamontopterus which share a unique combination of characters of the prosomal appendages and metastoma. A new genus, Kiaeropterus, is described to accommodate certain species previously assigned to Stylonurella but displaced from that genus as now emended.

A reconstruction of Parastylonurus ornatus in the walking position is attempted and has resulted in the recognition of a new generation of problems related to the functional morphology of these animals. Criteria which may appropriately be used in deciding the manner in which they walked and the posture which they adopted when doing so, are discussed. It is suggested that, in response to certain anatomical and physiological constraints which otherwise would have rendered the animals unstable in the walking position, the post-abdomen and telson have been specialised as hydrodynamic structures for monitoring water movements and vectoring the animal for maximum walking efficiency. The significance of this hypothesis is discussed in relation to the functional morphology and evolution of other members of the Stylonuroidea.

Type
Research Article
Copyright
Copyright © Royal Society of Edinburgh 1979

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References

Andrews, H. E., Brower, J. C., Gould, S. J. and Reyment, R. A., 1974. Growth and variation in Eurypterus remipes DeKay. Bull. Geol. Instn Univ. Upsala, 4, 81114.Google Scholar
Armstrong, J., Young, J. and Robertson, D., 1876. Catalogue of Western Scottish Fossils. British Association, Glasgow.Google Scholar
Brower, J. C., 1975. Silurian crinoids from the Pentland Hills, Scotland. Palaeontology, 18, 631656.Google Scholar
Caster, K. E. and Kjellesvig-Waering, E. N., 1956. Some notes on the genus Dolichopterus Hall. J. Paleont., 30, 1928.Google Scholar
Caster, K. E. and Kjellesvig-Waering, E. N., 1964. Upper Ordovician Eurypterids of Ohio. Palaeontogr. Am., 4, 301342.Google Scholar
Clarke, J. M. and Ruedemann, R., 1912. The Eurypterida of New York. Mem. N.Y. St. Mus. Nat. Hist., 14, 1439.Google Scholar
Clarkson, E. N. K., Eldredge, N. and Henry, J. L., 1977. Some Phacopina (Trilobita) from the Silurian of Scotland. Palaeontology, 20, 119142.Google Scholar
Cocks, L. R. M., 1970. Silurian brachiopods of the Superfamily Plectambonitacea. Bull. Br. Mus. Nat. Hist. (Geol.), 19, 139203.Google Scholar
Cocks, L. R. M., Holland, C. H., Rickards, R. B. and Strachan, I., 1971. A correlation of Silurian rocks in the British Isles. Jl Geol. Soc. Lond., 127, 103136.Google Scholar
Cocks, L. R. M. and Toghill, P., 1973. The Biostratigraphy of the Silurian rocks of the Girvan District, Scotland. Jl Geol. Soc. Lond., 129, 209243.Google Scholar
Copeland, D. E., 1968. Fine structures of salt and water uptake in the land crab Gecarcinus lateralis. Am. Zool., 8, 417432.Google Scholar
Copeland, D. E. and Fitzjarrill, A. T., 1968. The salt absorbing cells in the gills of the blue crab (Callinectes sapidens Rathbun) with notes on modified mitochondrea. Z. Zellforsch. Microsk. Anat., 92, 122.Google Scholar
Denison, R. H., 1956. A review of the habitat of the earliest vertebrates. Fieldiana, Geol., 11, 359457.Google Scholar
Diener, C., 1924. Fossilium Catalogus, I: Pars. 25: Eurypterida. Berlin.Google Scholar
Eldredge, N., 1971. Patterns of cephalic musculature in the Phacopina (Trilobita) and their phylogenetic significance. J. Paleont., 45, 5267.Google Scholar
Eldredge, N., 1972. Morphology and relationships of Bouleia Kozlowski, 1923 (Trilobita, Calmoniidae). J. Paleont., 46, 140151.Google Scholar
Etheridge, R., 1888. Fossils of the British Islands, vol. 1, Palaeozoic.Google Scholar
Fage, L., 1949. Classe des Mérostomacés (Merostomata, Woodward (1866)). In Grasse, P.-P., Traité de Zoologie, VI, 219262.Google Scholar
Fretter, V. and Graham, A., 1976. A Functional Anatomy of Invertebrates. London, New York and San Francisco, 589 pp.Google Scholar
Guieysse, A., 1908. Étude des Organes Digestifs chez le Scorpion. Archs Anat. Microsc., 10, 123139.Google Scholar
Hanken, N.-M. and Størmer, L., 1975. The trail of a large Silurian eurypterid. Fossils and Strata, 4, 255270.Google Scholar
Henderson, J., 1880. On some recently discovered Fossiliferous Beds in the Silurian Rocks of the Pentland Hills. Trans. Edinb. Geol. Soc., 3, 353356.CrossRefGoogle Scholar
Henderson, J. and Brown, D. J., 1867. On the Silurian Rocks of the Pentland Hills, Part I. Trans. Edinb. Geol. Soc., 1, 2333.Google Scholar
Henderson, J. and Brown, D. J., 1870. On the Silurian Rocks of the Pentland Hills, Part II. Trans. Edinb. Geol. Soc., 1, 266272.Google Scholar
Heubusch, C. A., 1962. Preservation of the Intestine in three specimens of Eurypterus. J. Paleont., 36, 222224.Google Scholar
Holm, G., 1898. Über die Organisation des Eurypterus fischeri Eichw., Zap. Imp. Akad. Nauk, 8, 2, 157.Google Scholar
Hunter, J. R. S., 1883. The Silurian rocks of the Logan Water, Lesmahagow. Trans. Geol. Soc. Glasg., 7, 5664.Google Scholar
Jennings, J. S., 1961. The geology of the eastern part of the Lesmahagow Inlier. Ph.D. thesis, Univ. of Edinburgh (unpublished).Google Scholar
Kier, P. M., 1973. A new Silurian echinoid genus from Scotland. Palaeontology, 16, 651663.Google Scholar
Kjellesvig-Waering, E. N., 1958. Some previously unknown morphological structures of Carcinosoma newlini (Claypole). J. Paleont., 32, 295303.Google Scholar
Kjellesvig-Waering, E. N., 1961 a. Eurypterids of the Devonian Holland Quarry Shale of Ohio. Fieldiana, Geol., 14, 7998.Google Scholar
Kjellesvig-Waering, E. N., 1961 b. The Silurian Eurypterida of the Welsh Borderland. J. Paleont., 35, 789835.Google Scholar
Kjellesvig-Waering, E. N., 1963. Pennsylvanian Invertebrates of the Mazon Creek Area, Illinois: Eurypterida. Fieldiana, Geol., 12, 85106.Google Scholar
Kjellesvig-Waering, E. N., 1966. A Revision of the families and genera of the Stylonuracea (Eurypterida). Fieldiana, Geol., 14, 169197.Google Scholar
Kjellesvig-Waering, E. N., 1978. Vattenfallet Eurypterida (in press).Google Scholar
Kjellesvig-Waering, E. N. and Heubusch, C. A., 1962. Some Eurypterida from the Ordovician and Silurian of New York. J. Paleont., 36, 211221.Google Scholar
Kjellesvig-Waering, E. N. and Størmer, L. 1952. The Dolichopterus-Strobilopterus Group in the Eurypterida. J. Paleont., 26, 659661.Google Scholar
Lamont, A., 1947. Gala-Tarannon Beds in the Pentland Hills, near Edinburgh. Geol. Mag., 84, 193–208. 289303.CrossRefGoogle Scholar
Lamont, A., 1952. Ecology and correlation of the Pentlandian—a new division of the Silurian system in Scotland. Rep. 18th Int. Geol. Congr. Lond., 10, 2732.Google Scholar
Lamont, A., 1955. Scottish Silurian Chelicerata. Trans. Edinb. Geol. Soc., 16, 200216.Google Scholar
Laurie, M., 1892a. Additions to the Eurypterid Fauna of the Upper Silurian. Rep. Br. Ass. Advmt Sci., 1892, 724725.Google Scholar
Laurie, M., 1892b. On Some Eurypterid Remains from the Upper Silurian Rocks of the Pentland Hills. Trans. Roy. Soc. Edinb., 37, 151161.Google Scholar
Laurie, M., 1893. The Eurypterid-bearing Deposits of the Pentland Hills. Rep. Br. Ass. Advmt Sci., 1893, 470.Google Scholar
Laurie, M., 1894. The Eurypterid-bearing Deposits of the Pentland Hills—Second Report. Rep. Br. Ass. Advmt Sci., 1894, 302303.Google Scholar
Laurie, M., 1898. The Eurypterid-bearing Rocks of the Pentland Hills—Final Report. Rep. Br. Ass. Advmt Sci., 1898, 557558.Google Scholar
Laurie, M., 1899. On a Silurian Scorpion and some additional Eurypterid Remains from the Pentland Hills. Trans Roy. Soc. Edinb., 39, 575590.Google Scholar
Lawrence, R. F., 1953. The biology of the cryptic fauna of forests, with special reference to the indigenous forests of South Africa, Balkema, A. A., Cape Town and Amsterdam, pp. 408, 160 figs.Google Scholar
Leutze, W. P., 1958. Eurypterids from the Silurian Tymochtee Dolomite of Ohio. J. Paleont., 32, 837942.Google Scholar
Lochhead, J. H., 1961. Locomotion. In The Physiology of Crustacea, ed. Waterman, T. H., vol. 2, 313364.Google Scholar
Mcmanus, J. J., 1969. Osmotic relations in the Horseshoe Crab, Limulus polyphemus. Am. Midl. Nat., 81, 569573.Google Scholar
Macmillan, D. L., 1975. A physiological analysis of walking in the American lobster (Homarus americanus). Phil. Trans. Roy. Soc. Lond. B, 270, 159.Google ScholarPubMed
Manton, S. M., 1950. The evolution of arthropodan locomotary mechanisms Part I, The locomotion of Peripatus. Zool. J. Linn. Soc., 41, 529570.Google Scholar
Manton, S. M., 1952. The evolution of arthropodan locomotary mechanisms Part II, General introduction to the locomotary mechanisms of the Arthropoda. Zool. J. Linn. Soc., 42, 93117.Google Scholar
Manton, S. M., 1958. Hydrostatic pressure and leg extension in Arthropods, with special reference to Arachnids. Ann. Mag. Nat. Hist., 1, 161182.Google Scholar
Manton, S. M., 1972. The evolution of arthropodan locomotary mechanisms. Part X, Locomotary habits, morphology and evolution of the hexapod classes. Zool. J. Linn. Soc., 51, 203400.Google Scholar
Manton, S. M., 1973 a. The evolution of arthropodan locomotary mechanisms, Part XI, Habits, morphology and evolution of the Unirama (Onychophora, Myriopoda, Hexapoda) and comparisons with the Arachnida, together with a functional review of uniramian musculature. Zool. J. Linn. Soc., 53, 257375.Google Scholar
Manton, S. M., 1973 b. Arthropod phylogeny—a modern synthesis. J. Zool. Lond., 171, 111130.Google Scholar
Miller, J., 1975. Structure and function of trilobite terrace lines. Fossils and Strata, 4, 155178.Google Scholar
Miller, J., 1976. The Sensory Fields and Life Mode of Phacops rana (Green, 1832) (Trilobita). Trans. Roy. Soc. Edinb., 69, 337367.Google Scholar
Moore, P. F., 1941. On gill-like structures in the Eurypterida. Geol. Mag., 78, 6270.Google Scholar
Mykura, W. and Smith, J. D. D., 1962. In The Geology of the Neighbourhood of Edinburgh, ed. Mitchell, G. H. and Mykura, W.. Ch. II, Ordovician and Silurian. Mem. Geol. Surv. Scotland, 1022.Google Scholar
Norford, B. S., 1973. Lower Silurian species of the trilobite Scotoharpes from Canada and Northern Greenland. Bull. Geol. Surv. Can. 222, 332.Google Scholar
Novojilov, N., 1962. Fundamentals of Palaeontology. Eurypterida, 404422. Moscow.Google Scholar
O'connell, M., 1916. The Habitat of the Eurypterida. Bull. Buffalo Soc. Nat. Sci., 11, 3, 277 pp.Google Scholar
Page, D., 1859 a. Advanced Text-Book of Geology, 2nd edn. Edinburgh and London.Google Scholar
Page, D., 1859 b. Further Contributions to the Palaeontology of the Tilestones or Siluro-Devonian Strata of Scotland. Rep. Br. Ass. Advmt Sci., 1858, 104105.Google Scholar
Page, D., 1860. On the Upper Silurian of Lesmahagow, Lanarkshire. Rep. Br. Ass. Advmt Sci., 1859, 6365.Google Scholar
Page, D., 1861. Advanced Text Book of Geology, 3rd edn. Edinburgh and London.Google Scholar
Panikkar, N. K., 1940. Influence of Temperature on Osmotic Behaviour of Some Crustacea and its Bearing on Problems of Animal Distribution. Nature, Lond., 146, 366367.CrossRefGoogle Scholar
Patten, W. and Redenbaugh, W. A., 1900. Studies on Limulus, I and II. J. Morph., 16, 1–26. 91200.Google Scholar
Peach, B. N. and Horne, J., 1899. The Silurian Rocks of Britain I. Scotland. Mem. Geol. Surv. U.K.Google Scholar
Peach, B. N., Horne, J. and Macconochie, A., 1901. In Fauna, Flora and Geology of the Clyde Area, ed. Elliot, Laurie & Murdoch, , pp. 423444. British Association.Google Scholar
Petrunkevitch, A., 1955. Chelicerata-Arachnida, In Treatise on Invertebrate Paleontology, ed. Moore, R. C., Part P. Arthropoda 2.Google Scholar
Richter, R., 1923. Von Bau und Leben der Trilobiten III. Die Beziehung von Glatze und Magen. Palaeont. Hung., 1, 7789.Google Scholar
Ritchie, A., 1963. Palaeontological studies on Scottish Silurian fish-beds. Ph.D thesis, Univ. of Edinburgh (unpublished).Google Scholar
Ritchie, A., 1968 a. New evidence on Jamoytius kerwoodi White, an important Ostracoderm from the Silurian of Lanarkshire, Scotland. Palaeontology, 11, 2139.Google Scholar
Ritchie, A., 1968 b. Lanarkopterus dolichoschelus (Størmer), gen. nov., a mixopterid eurypterid from the Upper Silurian of the Lesmahagow and Hagshaw Hills inliers, Scotland. Scott. J. Geol., 4, 317338.Google Scholar
Robertson, J. D., 1957. The habitat of the early vertebrates. Biol. Rev., 32, 156187.Google Scholar
Robertson, J. D., 1970. Osmotic and ionic regulation in the horseshoe crab Limulus polyphemus (Linnaeus). Biol. Bull. Mar. Biol. Lab., Woods Hole, 138, 157183.Google Scholar
Rolfe, W. D. I., 1973. Excursion 15: Silurian arthropod and fish assemblages from Lesmahagow, Lanarkshire. In Excursion Guide to the Geology of the Glasgow district, ed. Bluck, B. J., 119126. Geol. Soc. Glasg.Google Scholar
Ruedemann, R., 1921 a. Preservation of Alimentary Canal in an Eurypterid. Bull. N. Y. State Mus., 127/128, 9295.Google Scholar
Ruedemann, R., 1921. A recurrent Pittsford (Salina) Fauna. Bull. N.Y. State Mus., 219/220, 205215.Google Scholar
Ruedemann, R., 1935 b. The Eurypterids of Beartooth Butte, Wyoming. Proc. Amer. Phil. Soc., 75, 129141.Google Scholar
Schafer, W., 1954. Form und Funktion der Brachyuren-Schere. Abh. Senckenb. Naturforsch. Ges., 489, 165.Google Scholar
Schevill, W. E., 1936. Habits of trilobites. Natn Res. Coun. Comm. Paleoecology. Ann. Rep. 1935–1936, 2943.Google Scholar
Schmalfuss, H., 1978. Structure, Patterns, and Function of Cuticular Terraces in Recent and Fossil Arthropods. Zoomorphologie, 90, 1940.CrossRefGoogle Scholar
Smith, A. G., Briden, J. C. and Drewry, G. E., 1973. Phanerozoic World Maps. Spec. Pap. Palaeontol., 12, 139.Google Scholar
Størmer, L., 1934. Merostomata from the Downtonian Sandstone of Ringerike, Norway. Skr. Norske Vidensk-Akad., Mat-naturv Kl., 10, 125 pp.Google Scholar
Størmer, L., 1951. A new eurypterid from the Ordovician of Montgomeryshire, Wales. Geol. Mag., 88, 409422.Google Scholar
Størmer, L., 1955. Chelicerata-Merostomata. In Treatise on Invertebrate Paleontology, ed. Moore, R. C.. Part P. Arthropoda 2.Google Scholar
Størmer, L., 1969. Eurypterids from the Lower Devonian of Willwerath, Eifel. Senckenberg. Leth., 50, 2135.Google Scholar
Størmer, L., 1973. Arthropods from the Lower Devonian (Lower Emsian) of Alken an der Mosel, Germany. Part 3: Eurypterida, Hughmilleriidae. Senckenberg. Leth., 54, 119205.Google Scholar
Størmer, L., 1974. Arthropods from the Lower Devonian (Lower Emsian) of Alken an der Mosel, Germany. Part 4: Eurypterida, Drepanopteridae, and other groups. Senckenberg. Leth., 54, 359451.Google Scholar
Størmer, L., 1976. Arthropods from the Lower Devonian (Lower Emsian) of Alken an der Mosel, Germany. Part 5: Myriapoda and additional forms, with general remarks on fauna and problems regarding invasion of land by arthropods. Senckenberg. Leth., 57, 87183.Google Scholar
Størmer, L. and Kjellesvig-Waering, E. N., 1969. Sexual Dimorphism in Eurypterids. In Sexual Dimorphism in Fossil Metazoa and Taxonomic Implications, ed. Westermann, G. E. G.. International Union of Geological Sciences, A/1, 201214.Google Scholar
Størmer, L. and Waterston, C. D., 1968. Cyrtoctenus gen. nov., a large late Palaeozoic Arthropod with pectinate Appendages. Trans. Roy. Soc. Edinb., 68, 63104.Google Scholar
Tipper, J. C., 1976. The stratigraphy of the North Esk Inlier, Midlothian, Scotland. Scott. J. Geol., 12, 1522.Google Scholar
Waterston, C. D., 1957. The Scottish Carboniferous Eurypterida. Trans. Roy. Soc. Edinb., 63, 265288.Google Scholar
Waterston, C. D., 1962. Pagea sturrocki gen. et sp. nov., a new eurypterid from the Old Red Sandstone of Scotland. Palaeontology, 5, 137148.Google Scholar
Waterston, C. D., 1964. Observations on Pterygotid Eurypterids. Trans. Roy. Soc. Edinb., 66, 933.Google Scholar
Waterston, C. D., 1975. Gill structures in the Lower Devonian eurypterid Tarsopterella scotica. Fossils and Strata, 4, 241254.Google Scholar
Wills, L. J., 1965. A supplement to Gerhard Holm's Über die Organisation des Eurypterus Fischeri Eichw, with special reference to the organs of sight, respiration and reproduction. Ark. Zool., 18, 93145.Google Scholar
Woodward, H., 1864. Descriptions of some new Palaeozoic Crustacea. Geol. Mag., 1, 196200.Google Scholar
Woodward, H., 1872. A Monograph of the British Fossil Crustacea belonging to the Order Merostomata. Palaeontogr. Soc. (Monogr.), pt iv, 121180.Google Scholar