Sutural Simplification in Physodoceratinae (A~Pidoceratidae, Ammonitina)

Estudios geol., 43: 271-278 (1987) SUTURAL SIMPLIFICATION IN PHYSODOCERATINAE (A~PIDOCERATIDAE, AMMONITINA) A. Checa (*) RESUMEN El análisis estructural de la interrelación concha-septo en algunos ammonites del Jurá sico superior lleva a concluir que las simplificaciones suturales aparecidas a lo largo de la filogenia fueron originadas por alteraciones ocurridas en la morfología externa de la con cha. En el caso concreto de la subfamilia Physodoceratinae, la simplificación observada en la morfología de la sutura puede tener un doble origen. En primer lugar, un incremento en el tamaño de los tubérculos periumbilicales puede determinar una pérdida de profundi dad de los elementos de la sutura. siempre acompañada de una disminución en las inden taciones (frilling) de sillas y lóbulos. En otros casos el acortamiento en profundidad está determinado por una disminución de la tasa de expansión de la espira, sin que se observe un acortamiento aparente de las ramificaciones secundarias. Palabras clave: ammonites, morfología construccional, sutura septal, Physodoceratinae, Physodoceras, Benetticeras, Orthapidoceras, Schaireira, Pseudowaagenia. ABSTRACT The estructural analysis of the shell septum interrelationship in sorne Jurassic ammoni tes allows us to conclude that sutural simplifications occurred throughout the phylogeny, were originated by alterations in the external morphology of the shell. In the case of Phy sodoceratinae the simplification observed in the morphology of the septal suture may have a double origino First, an increase in the size of periumbilical tubercles may determine a shallowing of sutural elements and a shortening of saddle and ¡obe frilling. In other cases, shallowing is determined by a decrease in the whorl expansion rate, an apparent shorte ning of secondary branching not being observed. Key words: ammonites, constructional morphology, septal suture, Physodoceratinae, Phy sodoceras, Benetticeras, Orthaspidoceras, Shaireria, Pseudowaagenia. Introducción Other authors (Henderson, 1984; Ebel, 1985) claim a primary function of muscular attachment. A fascinating aspect which has attracted great In the "pneu-model" (Seilacher, 1975; Bayer, attention in ammonite paleontology is the study 1978a, 1978b) septal morphology is considered of the functionality of the suture and the inter as a result of pressure differences in the body-ca pretation of the different changes affecting it. A meral gas interphase, so that sutural morphology very commonly observed alteration is the loss of seems to be connected with the pressure distribu indentation and depth of sutural elements, that is tion along the whorl section and with locomotion to say, simplification. activity (Bayer, 1978b, p. 153), thus depending At the present moment there are several points only partly or indirectly on the depth of the orga of view as to the real function of ammonite septa. nism's life. Sorne views consider the septum as a structure In this paper we will focus on the analysis of which counteracts the hydrostatic pressure exer sorne particular cases of Physodoceratinae, where ted on the phragmocone (Westermann, 1975). a noticiable simplification and/or shallowing of (*) Departamento de Estratigrafía y Paleontología, Facultad de Ciencias, Universidad de Granada, Avda. Fuentenueva sin, 18002 Granada. 272 A. CHECA sutural elements is appreciated. We are going to coarseness of ornamentation, we may suppose work on the phylogeny proposed by Checa (1985) that the sllcessive lengthenings of the U/U, sad and on the analysis of the structural interrelations die will lead to the surpassing of a hypothetical li present in Physodoceratinae (Checa, 1986a, mit of stability; as a consequence, a morphologi 1986b). AIso we should explain that our observa cal reorganization would take place, possibly re tions refer exelusively to the morphological sim lated to the fractal pattern of growth of ammoni plification noted in individual sutural elements tic septal suture (Damiani, 1986). A considerable (always taking equivalent ontogenetic stages) loss of indentation and depth of the sutural ele considered apart from the sutural composition ex ments affected, that is to say, their simplification pressed as the number of umbilical lobes. takes also about. Given again the fractal nature of the suture line, it must maintain a homoge neous composition in any case, so that the reor Simplification due to changes in the size of ganizatíon of the rest of the suture must be indu tuberculation ced as a consequence of the partial remodelling noted in U/U]' The cases reported are the following: The case of Sch. pipini, although differing from - Benetticeras benettii (Fig. 1B): the "normal" those mentioned aboye, may be explained by re previous sutural morphology is that of Physo ference to a similar process. In this case an in doceras wolfi and the main ornamental change crease in tuberculation coarsening takes place consists of a considerable increase in the coar once again, though it is not so spectacular as in sening of the tuberculation. In both species the Orthaspidoceras and Beneuiceras. However, the suture ineludes an incipient Us and the position suture-ornamentation relatio~ship is now diffe of the periumbilical ornamental elements coin rent, the tubercles being placed on the lobe U3. cides with the saddle U/U3. As has been observed (Checa, 1986a) the lobes - Orthaspidoceras lallierianum (Fig. 1D): this could be less capable of lengthening than the sad species, a homeomorph of the aboye described, dles. In Sch. pipini this fact is accentuated by the is ineluded in a phylogenetic line where, once clearly internal position of U3 , so that a fairly more, the tendency consists of the increase in weak lengthening would give rise to a complete size of the periumbilical tubereles. The original sutural reorganization (Fig. 2). In this case we do non-simplified suture is, again, that of Ph. wol not appreciate any significant change in depth of fi, though also containig an intermediate form, sutural elements. O. ziegleri (Fig. le), whose suture line already Another interesing feature is the apparent shows a certain loss of sutural complexity. shortening of secondary f1uting (folioles), specia - Schaireria pipini (Fig. 2B): we may again consi lIy at the lateral portions of lobes and saddles. der this species as a homeomorph of the two This loss of indentation has a special interest as ones aboye discussed, although it belongs to an regards present day hypotheses on septal functio evolutive line where the "normal" original su nality, and concretely in rclation to that which ture, composed of six umbilical lobes, is that of contemplates sutural design as a mechanism of Sch. neumayri. The remaining forms included additional support of the outer shell against hi in this evolutive line show, as does Sch. pipini, drostatic pressure. In fact, second order f1uting a suture with a high degree of simplification, certainly constitutes a remarkable portion of su although there is no appreciable shallowing of tural length and their loss or reduction would the elements. considerably increase the unsupported shell area. Both O. lallierianum and Bu. benettii have Ph. Within the modifications aboye exposed it is pos wolfi as their common ancestor (Fig. 1). As has sible to consider that the intensity of the expan been stated elsewhere (Checa, 1986a) the size of sion must be proportional to the magnitude of the individual tubercles is correlated to that of the resulting tubercular projection. the saddle U/U3 , upon which the ornamentation As along a single tubercle the distance perpen grows, due to the plastic nature of sutural chan dicularly projected to the f1ank decreases from ges in these forms. In Ph. wolfi, in coarsely tu centre to peryphery according to the longitudinal berculate specimens, we already see extreme ca profil of the tuberele, it is to be supposed that the ses of stretching and subdivision of the saddle U/ required expansion follows a similar value curve. U 3 (Schindewolf anomaly). If we take into ac Thus, the intensity of the expansion will vary de count the fact that from this species till Bu. benet pending on the point considered, so that the af tii and O. lallierianum there exists an accentua fected sutural portion will also display "zones" of tion of the tendency leading to the increase in the different lengthening whose boundaries will, in 273 SUTURAL SIMPLIFICACION o x1.7 Fig. l.-Schematic representation and suture oí: A) Physodoceras wolfi (Ch.C,.8); B) Benetticeras benettii (K.QlO.24.1O); C) Orthaspidoceras ziegleri (F.GI4.6.5); D) Orrhaspidoceras lallierianum (SSI9). Arrows indicate phylogenetic connections. 274 A. CHECA B u',--:tJ" E x1.0 ~~1 SU3 I t U~ U A I ?'1If xl.O I ~'I• Fig. 2.-Schematic representation and suture of: A) Schaireria neumayri (F.G IS .14.1); B) Schaireria pipini (F.G¡¡.5.2). principIe, be continuous. On the other hand,gi homphala (upper Kimmeridgian) which shows a ven the plastic behaviour of the suture and the not;ceably simplified sutural desing (Fig. 4). Psw. existence of "active centres" of expansion, there haynaldi (lower-middle Kimmeridgian), whose is no reason why intermediate spaces between suture suffers a slight loss of complication, may two centres need to display a homogeneous com be considered as an intermediate species between pression along their entire width. In general the two. These morphological variations corres terms, it may be supposed that shortening will be ~ond to type ~ described in Checa (1986a) and maximal at points closest to "pushing" zones and fust modahty 10 Checa (1986b), consisting of a minimal at ones farthest from them. Bearing this loss of shell wall at the umbilical seam followed in mind, it is possible to modify the scheme obtai by a subsequent reorganization of the section, ned for O. lallierianum and Bu. beneuii by esta whose result could be that of maintaining the pe blishing strips of differential lengthening affecting riumbilical elements (tubercles) parallel to the saddles and differential compression along lobes. equatorial plane; in this way, sections similar to In Sch.

Load more