Devonian Carrier Shells (Euomphalidae) from North America and Germany GEOLOGICAL SURVEY PROFESSIONAL PAPER 824 Devonian Carrier Shells (Euomphalidae) from North America and Germany By ROBERT M. LINSLEY and ELLIS L. YOCHELSON GEOLOGICAL SURVEY PROFESSIONAL PAPER 824 A study of a behavior pattern in which foreign matter is attached to the shell of living and Devonian gastropods UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1973 UNITED STATES DEPARTMENT OF THE INTERIOR ROGERS C. B. MORTON, Secretary GEOLOGICAL SURVEY V. E. McKelvey, Director Library of Congress catalog-card No. 73-600142 For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402 Price $1.15 (paper cover) Stock Number 2401-02370 CONTENTS Page Page Abstract------------------------------------------------------------------------------------- 1 Systematic paleontology- Continued Introduction and acknowledgments-------------------------------------- 1 Superfamily Euomphalacea- Continued Implantation among the recent fauna _________________________________ _ 2 Family Euomphalidae- Continued . The Xenophoridae ___________________________________________________________________ _ 3 Genus Straparollus Montfort- Continued Habits of Xenophora conchyl·iophora ___________________________ , 3 Subgenus Euomphalus J. Sowerby______ 12 Habits of Xenophora neozelanica_________________________________ _ 4 Stratigraphic distribution______________________________________________________ 14 Functional significance of implanted material in Positioning of shell during implantation_____________________________ 15 the Xenop horidae .. __________________________________________ . ___________ _ 6 Distribution and nature of implanted materiaL________________ 17 Implantation in fossil gastropod shells _______________________________ _ 6 Functional significance of implanted materiaL_________________ 18 Systematic paleontology-------------------------------·------------------------- 7 Presence and absence of implanted materials_____________________ 19 Superfamily Euomphalacea __________________________________________ _ 7 The status of Philoxene___________________________________________________________ 19 Family Euomphalidae ____________________________________________ _ 7 Evolutionary groups of Devonian carrier shells________________ 19 Genus Straparollus Montfort ________________________ _ 7 Conclusions-------------------------------------------------------------------------------- 21 Subgenus Serpulospira Cossman __________ _ 7 Selected references------------------------------------------------------------------- 22 Subgenus Straparollus Montfort __________ _ 8 IndeX------------------------------------------------------------------------------------------- 25 ILLUSTRATIONS [Plates follow index] PLATE 1. Xenophora pallidula Reeve. 2. Xenophora neozelanica Suter. 3. Straparollus (Strapm·ollus) and S. (Serpulospira). 4. Straparollus (Strapat·ollus). 5, 6. Straparollus (Euomphalus) and S. (Straparollus). Page FIGURE 1. Chart showing stratigraphic occurrence of incrusting euomphalids_______________________________________ 15 2. Drawings showing reconstruction of euomphalids as mobile animals__________________________________ 16 3. Drawings showing reconstruction of euomphalids as sessile animals__________________________________ 17 III DEVONIAN CARRIER SHELLS (EUOMPHALIDAE) FROM NORTH AMERICA AND GERMANY By RoBERT M. LINSLEYl and ELLIS L. YocHELSON ABSTRACT p. 241) noted in his comprehensive study of Paleo­ The modern carrier shell Xenophm·a has an elaborate be­ zoic gastropod type species, the holotype of E. laevis havior pattern in which the gastropod attaches foreign mat­ does not have any foreign material attached to its ter to its shell. The process is a long and deliberate one that has selective significance in terms of visual and olfactory shell. Other specimens, which are presumably con­ camouflage. Besides the Xenophoridae, first certainly known specific, to some degree do cement shell material to from the Cretaceous, one turritellid in the Miocene-Pliocene their shells. and one Miocene modulid mastered this peculiar art. Various Examination of topotypic material of the type members of the Euomphalacea from the Ordovician, Silurian, species of Philoxene and of all the available Devo­ and Devonian implanted shell fragments. The systematic part of this report consists primarily of de­ nian species in North America that attach foreign scriptions of the known Middle and Late Devonian carrier material suggests that this peculiar habit may be shells of North America; it includes a discussion of eight indulged in by individuals within a population but species, five of which are new. The new species are Strapa­ is not necessarily followed by all members. Our study rollus (Straparollus) mortoni, S. (Straparollus) cottrelli, S. has also demonstrated that the implantation of for­ (Euomphalus) hoffmani, S. (Euomphalus) winnipegosis, and S. ( ?Euomphalus) incrustatus. Specimens of the type species eign matter on the shell is not restricted to a par­ of Philoxene and two related forms were also studied. ticular shell form but cuts across the lines of three The Devonian carrier shells are of two or more separate currently recognized subgenera. These observations evolutionary stocks having different stratigraphic ranges. raise questions concerning the taxonomic, functional, One group implants foreign matter regularly; in the other, and stratigraphic significance of this particular trait this feature is quite irregular. Both groups include species having individuals that do not implant material. as well as the biologic validity of Philoxene. In an In both living and fossil carrier shells, the animals show attempt to understand the problems presented by little preference in materials selected except on the basis of these Devonian forms, we have examined the modern size. The implanted material probably served primarily as carrier shell X enophora and the closely related Tu­ tactile camouflage as well as visual camouflage. gurium. Philoxene Kayser is based on implantation and is here re­ The present study is a fusion of interests origi­ garded as based on a spurious concept. The type species should be placed in the Straparollus senso stricto and other nally derived from two separate geographic areas. species distributed under various subgenera of Straparollus; More than a decade ago, M. H. Staatz, U.S. Geologi­ implantation alone seems a poor criterion for discriminating cal Survey, submitted for examination a collection a species. The species studied in this paper are treated under from Nevada which was later identified asPhiloxene. three subgenera: Straparollus (Straparollus), S. (Serpulo­ The effort required to transport a large block of spira), and S. (Euomphalus). Species of S. (Euomphalus) in which implantation occurs seem to indicate that the angu­ limestone for acid treatment from an area difficult lated whorl profile characteristic of this taxon evolved more of access is cordially acknowledged. than once in time. For the past 10 years, Linsley has been engaged INTRODUCTION AND ACKNOWLEDGMENTS in a study of the gastropod fauna of the Rogers City Limestone. Access to the Calcite quarry, Michigan Kayser (1889, p. 292) established the genus Phil­ Limestone Operations, U.S. Steel Corp., at Rogers oxene and designated Euomphalus laevis Archiac City, Mich., and to the Presque Isle Corporation and Verneuil (1842) as type species. The genus was quarry (formerly Lake of the Woods quarry) north judged distinct from other euomphalid gastropods of Alpena, Mich., managed by Mr. Roy Hutchison because individuals attached foreign matter, usually for a consortium of steel companies, has always been shell fragments, to their shells. As Knight (1941, graciously granted. Field investigations in 1966-69 1Department of Geology, Colgate University, Hamilton, N.Y. were supported by the ~esearch Council of Colgate 1 2 DEVONIAN CARRIER SHELLS (EUOMPHALIDAE) FROM NORTH AMERICA AND GERMANY University. In addition, during 1965, the National IMPLANTATION AMONG THE RECENT FAUNA Science Foundation provided a grant to support a The incorporation of foreign bodies into the hard collecting trip by 10 high school students under Lins­ parts of an organism is not a common trait, but it ley's supervision. Further, the Research Council of is widespread in a systematic sense. Some Forami­ Colgate University provided an Undergraduate Re­ nifera have apparently been building an agglutinated search Participation Grant which permitted John test (Towe, 1967) since the Cambrian; other Fo­ Cottrell and John Hoffman to assist with fieldwork. raminifera may imbed foreign grains in a calcium Naming of two species after these assistants is a carbonate test. A few tunicates and sponges aggluti­ partial acknowledgment of the calibre of help they nate sand grains in their flesh, in part, perhaps, provided throughout the various stages of this inves­ accidentally. Worm burrows may be formed of tigation. grains of sand bound by mucus, and many worms Specimens were generously loaned by Dr. Roger that live on the bottom construct tubes of clastic Batten, American Museum of Natural History grains. Fresh-water caddis fly larvae and terrestrial (AMNH). Large collections were lent for study by bagworms are well-known examples of makers of the Museum of Paleontology, University of Michi­ agglutinated tubes. Several animals, such as the gan (UMMP). Dr. G. M. Ehlers, University of "decorator crab," which