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The Evolution of Ammonites.* a Demonstration Given at the British Museum (Natural History)

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The Evolution of Ammonites.* a Demonstration Given at the British Museum (Natural History) 49 THE EVOLUTION OF AMMONITES.* A DEMONSTRATION GIVEN AT THE BRITISH MUSEUM (NATURAL HISTORY). SATURDAY, 8TH FEBRUARY, 1919. By W. D. LANG, Sc.D., F.G.S. (J:}' TI.'l'missioH (If the Trustees of tilt British Museum.) I.-INTRODUCTORY. HE Evolution of Ammonites may be described as a phrase with two faces looking in opposite directions. From theTview-point of a primitive Ammonite, one face looks back to processes whereby Ammonites have been produced from organisms which Were not Ammonites; the other looks forward to the developmental history of Ammonites as such, as they diverge along multitudes of lineages, attain importance. then dominance among marine invertebrates, and finally become extinct. The subject is manifestly immense; and on such an occasion as this only hints can be given here and there of the apparent method of evolution, and perhaps a few suggestions, little better than guesses, as to some proximate causes. It would, perhaps, be true to say that a large treatise on Ammonites could do no more than this, and that all that can be hoped for now is to indicate some of the directions such a treatise would pursue; or to create an atmosphere, or attitude of mind. in which the methods of Ammonite evolution can better be understood. It is unfortunate that Ammonites suffer from their termino­ logy. To the uninitiated, the talk and writings of Ammonite specialists tend to appear to be an unnecessary and unintelligible jargon; and the amateur is discouraged from the study of what appears to him an attractive subject demoralised into a mystery by a few keen terminologists. It is unnecessary to say that this is a mistaken view, but it is one which appears to come very naturally to the amateur. Perhaps a demonstra­ tion given by an amateur (as far as Ammonites are concerned) to amateurs may help this difficulty. It is a fact that the difficulties of Ammonite-study confront the beginner at once. "There is no royal road to sutures," was once said by an Ammonite specialist; and this dictum applies with equal force to other Ammonite matters. There is not even the pleasant *Most of the facts and many of the opinions given below, as well as a wealth of additional information, may be found in C. R. Eastman's translation of L. von Zittel's Text-book of r akvcntotogy, (1900) and inA Guide to tne Fossil Invertebrate Animals in the Department of Gcoloi!Y ami Palaotttoloey in the British. Museum (Nat. His! .}, 1907. PROC. GEOL. Assoc., VOL. XXX., PART 2., 1919. 4 50 W. D lANG ON by-path for beginners that so man y subjects afford before the foot-hills of difficulty are reached . On th e contrary, the difficulties presented by Ammonites become mountain ous immedi ately the asthetic is ex changed for the scienti fic view. Terminology is a necessary aid brought forwar d to help to surmount these di fficulti es, and, if thus accepted, is not so troublesome to mast er. And if the terminology is crabbed , it is because the study of Ammonites is complex. Hence such terms as pro toconch, oxycone, homceomorph, and tachygenesis, ceasing to terrify, may begin to help ; and from being bad masters may become good servants. Perhaps a greater dr awback to Ammonite- study than a crabbed ter min ology, is th e extent and diffusion of Ammonite literature. But this mainly concerns their syste matics, and our present theme is evolution . Now a fundamental prinriple that has hitherto guided investigat ion into Amm onit e evoluti on is the so-called law of Recapitulation. Some discredit has been thrown upon this principle by workers in Recent zoology, because they have chiefly sought for it s expression in embryonic stages . But the immediate environment of an embryo is so special, that the ancest ral traits expected to be visible in its development are wont to be masked by other features caused by adaptation to its peculiar surroundings. In a fossil, how­ ever, we cannot ord inarily have direct knowledge of any growth­ stage that preceded the shell. And , though an embryonic shell is supposed to have been formed , that is, a shell present within the egg ; yet it is on recapitulation exhibited in post­ em bryonic growth-stages tha t the palreontologist chiefly relies for eluci dat ion of an Ammonite's evolut ion . For instance, an Ammonite from the Dorset Lias, which is generally known as lY1icroceras densinodus,* shows at its centre the so-called embryonic shell, or protoconch , followed by about lour whorls which have no ornament: then follow from two to th ree whorls with ri bbed or costate ornament; an d, finally, a stage in which the ribs acquire tubercles near the periphery . Consequent ly , it is supposed that the ancestral form had a plain shell, and that, during its evolution, the stock, or lineage from which .1.1'1. densinodus arose acquired, first a ribbed , and then a tubercula te ornament. This succession of plain, costate and tubcrculate stages in Ammonite ornament is shown in nu merous group s quite unconnected with M. densin odus and with each other (for instance, in the Liparo­ ceratid lineage discussed further on), and illustrat es a second principle fundament ally affecting Ammonite evolutio n, namely . that of Orthogenesis, whereb y t he same general sequence of "Tbe particul ar spec ime n exhibited has been labelled by ~Ir. S. S . Buckman ,;\/jrf"oceras obsoletn m Blake (non Simpson) sp. \Vhile gratefully accepting Mr. Buckman's determin ation . J have, for the sake of am at eurs, used at least the trivial nam e by w hich th is form and probably other similar, but specifically different for ms, from Charm ou th are generall y known . THE EVOLUTION OF AMMONITES. 5r structures is repeated again and again in different lineages springing from one ancestral form, so that it is possible to some extent to predict the after-history of a given primitive species. Bearing in mind, then, these two principles, namely, that of Recapitulation, following which the individual during its development repeats the history of the race; and Orthogenesis, whereby it may be predicted that the lineages diverging from an ancestral form will severally run through similar consecutive stages, let us first of all briefly consider the evolution which in latest Palseozoic times culminated in the Ammonites. Il.-EvOLUTION LEADING UP TO THE AM,\WNITE. The nearest approach in appearance to an Ammonite among Recent organisms is the Pearly Nautilus. There are manifest differences between an Ammonite-shell and the shell of Nautilus (fig. r}: notably, in Nautilus, the simply-curved suture, the median siphuncle with backwardly-projecting septal 'I b Fig. I. A Nautilus :.HHl an Ammonite. a is a plaster cast of the inside of the shell of the Recent "';aulitus ponipilins, and may be compared directly with the _si m ilar iutr-rnal cast (h) of the Ammonite. .c, sururc : ni. mark left by muscle of attachment; I. !Ines left by this mark in its previous positions. The slphu.llcle WIth.ItS se~ta! necks are not shown in these views, hut the position of t~Je slphu~de In (b) IS nH!Icate(l by the peripheral keel. (From British Museum c.mdc to Fossil Invertebrate Anirnals.) necks, the comparatively unornamentcd shell, and a ventral bay or sinus at the aperture of the shell; and, in an Ammonite, the. cOJ.TIplex suture, the perip~eral siphuncle with forwardly­ proJect~ng necks, a shell often highly ornamented, and a ventral projection at the aperture of the shell. But these differences are not great enough to preclude the probable affinitv of the tw.o shells; a.nd pakeonotology not only confirms their 'relation­ ~ll1p, but ~luCldate.s Its nature. Na tilus among Recent molluscs IS a peculiar and Isolated form of the order Cephalopoda. The 52 W . D LANG ON other Recent Cephalopods are the Cuttle-fish, Squid, Oct opus and their near allies. (The shell of the so-called 'Paper Naut ilus ' is merely a brood-cha mber for the eggs of an Octopod, and ha s no homologies with the shells of Nautilus and the Ammonites .) Those Cephalopods havin g shells built on lines similar to that of N autilus compose the group N autiloidea ; while Cuttles, Squids, Octopods and such-like are the B elem­ uoidea ; and the A mm onoidea comprise the Ammonites and their near relatio ns . Now the earl iest growth-stage manifested in Ammonites is the so-called embryonic shell, or P rotoconch (fig. 2), It is reasonable, therefore, when investi gating their mutual relation­ ship, to compare the protoconchs of ammonoids and N autilus. F ig. 2 (on left). A much-en largerl secti onal view of t he apex of th e shell of th e Ammoni te P ali oplruroceras spina /11m, sho wing th e large globular pro to conch (p) with t he blind end of th e siphuncle pro ject ing into it. The posit ion of th e siphun cle is indicated and it is seen rather rap idly edging to t he periphery ' of th e whorl ; but t he sept al necks and th e di scon tinuit y of t he se pta where they meet the siphuncle are not shown , (Adapted, after Branco, fro m Zittel's Text­ book of Palzeontol ogv}. F i g. 3 (on right ). A mu ch-enlarged sectiona l view of t he apex of t he shell of Nautilus pompilills, showing the absenc e of a protoconch.
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