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A Quantitative Study of Variation Within The A QUANTITATIVE STUDY OF VARIATION WITHIN THE MESOZOIC ECHINOID NUCLEOLITES. David Gordon Attilio Scurry, B.Sc Department of Geology Bedford College, London. Submitted for the degree of Doctor of Philosophy. 1 9 7 9 ProQuest Number: 10098370 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest. ProQuest 10098370 Published by ProQuest LLC(2016). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code. Microform Edition © ProQuest LLC. ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106-1346 A quantitative study of variation within the Mesozoic echinoid Nucleolites. D,G.A.Scurry. ABSTRACT Bivariate (reduced major axes) and multivariate (discriminant functions,canonical variâtes,principal components,cluster analysis) computer techniques are used to analyse quantitative measurements on fifteen test dimensions of the Mesozoic cassiduloid echinoid Nucleolites and its probable living descendant Apatopygus recens. A modification to standard techniques is described for analysis of reduced major axes and the cluster analysis. Twenty" samples are analysed. They include nine species of Nucleolites from the Jurassic (N.ampTus,N,burgundiàè,N.elongatus, N.latiporus,N.micraulus,N.scutatus,N.woodwardii) of England and France and Cretaceous (N.rotundus,N.subquadratus) of France and North Africa, The genotype,Nucleolites scutatus,is represented by seven samples. Significant infraspecific variation occurs primarily in size,shape and position of the periproct and in the length of the anal sulcus. A large periproct correlates with large sediment grain s iz e . No such differences are proved between three samples of N.latiporus ( = N.clunicularis auct.). Statistical differences between N.latiporus and N.scutatus are of the same order as between infraspecific samples of N.scutatus. Samples of the seven remaining fossil species are compared to the genotype. Quantitatively,differences can be expressed in terms of two test variables,always including either the length of the periproct and/or its distance from the apical disc. The size of the periproct is facies dependent. Within the Jurassic species, the length of the anal sulcus is consistently shorter in the stratigraphically younger species. This trend is not continued in the Cretaceous species. Overall,the distance of the apical disc from the anterior of the test becomes consistently shorter in the stratigraphically younger of the nine species studied. N.scutatus shows a similar range of variation to infraspecific samples of Apatopygus reccns from New Zealand, A.recens burrows completely in a coarse substrate and injests large substrate particles. N.scutatus probably did likewise. These habits are therefore of ancient origin in the Cassiduloida, CONTENTS Abstract p./ 2 1. Introduction p. 4 2. Stratigraphy p. 11 3* S y ste m a tic s p . 22 4« Methods of Study p. 90 5* Infraspecific variation within Nucleolites scutatus p.126 6. 'Biometrics of Nucleolites latiuorus p.l$8 7* Biometrics of some other Jurassic nucleolitids p . 189 8* Bioihetrics of some Cretaceous nucleolitids p . 255 9* Biometrics of Apatopygus recens p . 282 10. Discussion p . 304 11. Summary p .321 12. References. p . 326 List of Plates Nucleolites s c u ta tu s P l. 1 Nucleolites amplus P l. 2 Nucleolites woodwardii p l . 2 Nucleolites e lo n g a tu s p l . 3 Nucleolites l a tin o r u s p l . 3 Nucleolites m ic rau lu s p l . 4 Nucleolites subouadratus p l . 4 Nucleolites ro tu n d u s p l . 5 Apatopygus recens p l . 5 CHAPTER 1 - INTRODUCTION; 1 A) The genus NUCLEOLITES Nucleol i tes is an important Mesozoic irregular echinoid genus because of its common occurrence; its numerous species; its long stratigraphie range; the fact that it is one of the earliesc known and arguably one of the most primitive of the cassiduloid e.chinoids; and because of its presumed ancestry of or clo se a f f i n i t y w ith a l l o th er members of the order & Cassiduloida, The first undoubted occurrence of Nucleolites is in the parklnsoni zone of the middle J u ra s s ic upper B.-vjocian stag e. The genus becomes diverse in number of species and locally abundant in number of individuals for the first time in the succeeding Bathonian stage. It seems to reach its acme,judged by common occurrence of individuals and extensive geographic range,in the Oxfordian,N,scutatus Lamarck especially being common in the Corallian (middle and upper Oxfordian) of England and France, Subsequently the genus is represented by less common individuals but comprises species showing a greater range in morphology. In the Cretaceous it gives rise to many new genera but also disappears from England by this time. It is then of frequent occurrence only in eastern France,Germany and Switzerland. By the upper Cretaceous it is found mainly in Morocco,Algeria and Tunisia,the last undoubted. Nucleolites species,N.rotundus (Peron & Gauthier),occurring in the Cenomanian of Algeria (Kier,1962). /' The genus is diverse with over eighty named species ( sensu Lambert & T hiery,1909). They occur most commonly in limestones and in the north west European area. The greatest 6. • number of species as opposed to individuals occurs in the Neocoralan ( f i g . 1 ,1 ), The total range of the genus from upper Bajocian to Cenomanian represents a time span of some sixty nine million years (Marland P^^yi964), This is significantly longer than most other cassiduloid genera (sensu Kier,1962). In that it occurs in the Bajocian,barely two stages later than the earliest known cassiduloid (Galeropygus Cotteau,l856) in the Toarcian (Kier,1977), Nucleolites is an early cassiduloid, Kier (1962,1966) considers it to be not only the type but ttie ancestral genus of the family Nucleolitidae. The family Nucleolitidae Agassiz & Desor,1847,is itself one of the largest and oldest families of the Cassiduloida, All other families of the Cassiduloida, with the exception of the contemporary Clypeidae Lambert,1898, and the ancestral Galeropygidae Lambert 1911,are probably derived from the Nucleolitidae (sensu Kier,1962), Nucleolites is therefore an early and long ranging genus which probably formed the root stock of its family and most of the order Cassiduloida (Kier,1962), The cassiduloids themselves constitute one of the five principal orders of irregular echinoids, 1 B) Taxonomy Because it is common,Nucleolites has been studied for 300 years. Because it is diverse,and its species widespread throughout west Europe,there has been considerable duplication of nomenclature at both generic and species level, A number of genera are synonymous, Pomel (1883),for example, erected eight new genera on slight differences in shape and position of the periproct and length of the sulcus. All are now 5, if ) C O c u (D "O CJ (L> - (0 o n m C < a - c/i 0) CL - 4—' I— a < *0 CL - . cu (J TJ o u c (b m 3 a Z (U Z cn (j) ~4— > O o o I— - 4-* CD o (0 (0 CL 0» u 'E u o (D E N CL O k: 10 (0 (b X— O Z X <u o u ( b c JÛ -------- 0 Ü 1— E 1_ a o 3 O O -4-» O a CÛ , •—A a ÛÛ Oj —r— —r— D CD <r CM o o o CD CM D) Ll saiDeds ;o jequunM considered synonymous with Nucleoli tes by most authors. There has been considerable controversy related to the use of the names Nucleoli tes and Echinobris;:us and this is discussed fully below (p.22 ). Wright,Mortenson and Kier,among others,have pointed out the difficulty in distinguishing between species attributed to Nuclcolites, Cotteau figured many specimens in his works on French echinoids in the latter half of the nineteenth century. Unfortunately his artist made many inaccurate drawings and in the case of Nucleolites consistently drew specimens with single pores rather than double pores in each half ambulacrum beyond the” petals, 'This feature has subsequently been found to be of great evolutionary importance within cassiduloid echinoids; single extra-petaloid pores found in only post-Albian cassiduloids (Kier,1962),• There is need,therefore,to clarify.the taxonomy of the genus and its constituent species, Nucleolites scutatus Lamarck,1816, is here confirmed as the genotype. 1 C) Q u a n tita tiv e m ethods' In order to discriminate effectively between species, some assessment of variation is required at both infra and inter specific level. Biometrical methods are adopted here because of the particular suitability of echinoids to statistical analysis. The test of echinoderms is internal and so preserves more features suitable for study than most other invertebrates. The approach has been applied to the fossil spatangoid Micraster (e,g, Rowe,1899; Kermack,1954) and to living spatangoids (e,g, Nichols,1959,1962) but not significantly hitherto to cassiduloids, Roman's (1956,1957) attempts to differentiate species of Echinolampas on the basis of museum collections are the only comparable studies with respect to this order. ID ) Computer techniques It is shown that at least 56 different measurements may be taken in order to characterise the form of the Nucleolites test. However,15 measurements are usually adequate to define the morphology of a local species population, 'ibis study describes samples ranging in size from three * to one hundred and thirty three specimens,comprising in total twenty two samples allocated to ten species.
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