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THE UNIVERSITY OF KANSAS PALEONTOLOGICAL CONTRIBUTIONS

May 15, 1970 Paper 47

SIGNIFICANCE OF SUTURES IN PHYLOGENY OF

JURGEN KULLMANN AND JOST WIEDMANN Universinit Tubingen, Germany

ABSTRACT Because of their complex structure ammonoid sutures offer best possibilities for the recognition of homologies. Sutures comprise a set of individual elements, which may be changed during the course of ontogeny and phylogeny as a result of heterotopy, hetero- morphy, and heterochrony. By means of a morphogenetic symbol terminology, sutural formulas may be established which show the composition of adult sutures as well as their ontogenetic development.

WEDEKIND ' S terminology system is preferred because it is the oldest and morphogenetically

the most consequent, whereas RUZHENTSEV ' S system seems to be inadequate because of its

usage of different symbols for homologous elements. WEDEKIND ' S system includes only five symbols: E (for external lobe), L (for lateral lobe), I (for internal lobe), A (for adventitious lobe), U (for umbilical lobe). Investigations on ontogenetic development show that all taxonomic groups of the entire superorder Ammonoidea can be compared one with another by means of their sutural development, expressed by their sutural formulas. Most of the higher and many of the lower taxa can be solely characterized and arranged in phylogenetic relationship by use of their sutural formulas.

INTRODUCTION Today very few ammonoid workers doubt the (e.g., conch shape, sculpture, growth lines) rep- importance of sutures as indication of ammonoid resent less complicated structures; therefore, phylogeny. The considerable advances in our numerous homeomorphs restrict the usefulness of knowledge of ammonoid evolution during recent these features for phylogenetic investigations. decades have been based overwhelmingly upon The exact function of the specific form of the investigations of sutures. septum and its correlation with organs of the In general, recognition of homologies is re- ammonoid are not yet known. At any garded as the principal method of phylogenetics. moment in the life of the animal the intimate Recognition of homologous similarities, and dif- relationship between the last septum and soft ferentiation of homologous from analogous struc- parts of the organism is shown by the morpho- tures of similar function, are the prime indicators logical positions of this septum, which forms the of the relationships of organisms. adapical termination of the body-chamber against Among ammonoids sutures offer best pos- the phragmocone. Because the septum is so much sibilities for the identification of homologies, for a part of the general organization of the am- they are manifestations of highly complex an- monoid animal, changes in soft parts connected atomical structures, and the linear nature of su- to the septum are always accompanied by changes tures allows comparisons between them to be in the septum. Important changes of the suture made with relative ease. Other characteristics during ontogeny and in the phylogeny of any 2 The University of Kansas Paleontological Contributions—Paper 47 ammonoid group thus reflect changes of the FURNISH, 1957, p. L23, fig. 11; RUZHENTSEV, 1956, other organs of the ammonoid animal. fig. 29) seems to be rare. In the course of phy- The great overall stability and phylogenetic logeny, however, simplification of sutures is not irreversibility of suture development strongly im- at all uncommon as, for example, in the clymeniids ply an extensive system of pleiotropic genes and where forms with few sutural elements are derived polygenic characters (see Kosswio, 1959, p. 218). from others with many lobes; also certain Creta- This indicates that the suture represents a large cous ammonoid taxa with quadrilobate primary amount of the genetic material of the organism. sutures stem from forms with five-lobed primary From the genetical point of view, then, importance sutures. of the suture in ammonoid phylogeny and sys- Morphological features observed in the onto- tematics is obvious. genetic development of sutures in numerous am- The recognition of homologous structures in monoid genera, together with symbols adopted the suture is based upon two kinds of information for such parts as their lobes and saddles, are —stratigraphie succession of the ammonoids and illustrated in Figures 1 to 10, inclusive, distributed their ontogenetic development. The first depends through most of the discussions given in the upon favorable stratigraphie sections, and the present paper. Summaries of suture development second upon favorable preservation of the . contained in Figures 11 to 16 follow other illustra- The methodological prerequisites for recogni- tions in a group toward the end of the paper. tion of homologies and their role in the identifica- Such an arrangement should be most convenient tion of phylogenetic relationships were treated in for readers and it is hoped will facilitate their a precise way by REMANE (1952). Also, applica- understanding. Attention may be called here to tion of homology criteria to ammonoid sutures the editorial practice of printing symbols for suture has been discussed by SCHINDEWOLF in numerous parts in italics throughout the text, for it is judged publications (e.g., 1961-69). that simplicity and clarity are furthered by this Lobes are individual elements of the suture. typographic distinction. Lettering on diagrams In ontogeny they are formed first after conclusion speaks for itself. of the larval stage (i.e., after formation of the prosuture). Individual lobes are designated in ACKNOWLEDGMENTS to accordance with their original position relative We wish to thank Professor O. H. SCHINDE- other elements. During ontogeny and phylogeny WOLF for his active interest and helpful discus- three types of changes can occur: the position of sions. Appreciation is extended to Professors R. lobes may change (heterotopy); the lobe itself C. MOORE and CURT TLICHLRT, Lawrence, Kansas, may change (heteromorphy); or the ontogenetic for critical reading of the manuscript and offering change time of appearance of a lobe may many valuable suggestions. J. KULLMANN is (heterochrony). responsible for the introductory part, explanation In the course of ontogeny individual elements of WEDEKIND ' S terminology and discussion of may be reduced as, for example, in Spiroceras Paleozoic ammonoids; he expresses his thanks to bifurcati (QuENsTEDT) as reported by SCHINDE- WOLF (1951, p. 29, fig. 17; 1961, p. 92, fig. 45). Dr. R. A. DAVIS, formerly Iowa City, Iowa, and Disappearance of elements during ontogeny, as now at Swansea, Wales, for stimulating discussion for example in Paracuariceras incisum (see of the introductory parts of the present work. SCHINDEWOLF, 1965, p. 496, fig. 296) and perhaps J. WIEDMANN is responsible for the discussion of Agathiceras uralicum KARPINSKY (see MILLER & Mesozoic ammonoids.

DESIGNATION OF INDIVIDUAL PARTS OF SUTURES For purpose of communication among paleon- dividual elements in the adult suture is by far tologists a terminology for individual elements simpler and serves well for communication of of complex ammonoid sutures is necessary. Two morphological data. The latter, based on onto- types of terminology, not necessarily mutually genetic development, seems to be more useful exclusive, have been used: a morphographic- in communicating information about homologies. descriptive system and one based on morpho- When a strictly morphogenetic suture terminology genesis. The former, based on position of in- is used, each element of the mature suture is

Kullmann & Wiedmann—Sutures in Phylogeny of Ammonoidea 3 designated by a symbol which records the entire ing to their position by supplementary letters ti ontogeny of that element. Such symbols can be (ventral), m (medial), and d (dorsal) (e.g., gathered together into groups which record the Ei-->E ivEi „,Eid ). Single lobes formed in this ontogeny of the adult suture. These groupings way may be subdivided repeatedly (e.g., Evi--) of symbols have been called sutural formulas. EidvEid.Eie2; E142 ---> EteuvEid2mEids—with un- It is possible to symbolize each element of derstanding that the latter numeral in E1d2, E 1d21), the adult ammonoid suture solely by its position Ems, etc., indicates the quantity of the preceding in the suture. Such symbols can be grouped into letter). On the other hand, if a lobe divides into what resembles the sutural formula as described a series of incisions (e.g., Crimites, I 4 =S, Fig. above but indicates only the relative position of 12,e), it is designated by the letter S (Suturallobus, the sutural elements in the adult suture. WEDEKIND, 1916; SCHINDEWOLF, 1961, p. 63-78, NOETLING (1905, 1906) first used a morpho- fig. 31). Since the incisions do not represent genetic symbol terminology. WEDEKIND (1913) homologous units, their detailed notation is super- improved this and later applied his system to fluous. A morphological description of the in- ammonoids in general. His is the oldest practica- cisions would unnecessarily complicate formula ble morphogenetic system of terminology. characterization of the sutures, and any gain SCHMIDT (1921) added morphographical sym- would be small, as such sutural characteristics are bols to the morphogenetical terminology of neither generic, nor even specific, peculiarities, WEDEKIND. RUZHENTSEV (e.g., 1949, 1957), using and often occur asymmetrically. what is essentially WEDEKIND ' S system, proposed Brackets and colons, introduced by RUZHEN- a number of new names and symbols for sutural TSEV, serve to classify parts of the suture. Lobes elements and his lead has been followed by several in the process of division are placed in brackets American authors (e.g., FURNISH, GLENISTER, so long as their intermediary saddles have not MCCALEB, NASSICHUK). Soviet colleagues (notably reached the height of the neighboring saddles. Popov, 1965) have made additional terminological It is difficult to decide at what point a lobe in modifications. the process of development becomes independent, however. A colon is used to represent an um- WEDEKIND'S TERMINOLOGY bilical seam and can be used with adequate ac- curacy only when a saddle lies on an umbilical WEDEKIND ' S terminology was applied primarily seam. If a to the primary suture, which is the second suture lobe is halved by the umbilical seam the use of a colon becomes meaningless. to be formed. The so-called prosuture, which Successive stages in sutural precedes the primary suture, belongs to the pro- development are indicated by arrows between septum and is part of the protoconch. The first- sutural formulas or corresponding parts of suture formulas. formed suture shows in earliest ammonoids a In the simple circle, in later representatives a prominent order of their phylogenetic appearance the external saddle, thus differing basically from all five symbols proposed by WEDEKIND are: following sutures. The majority of all Paleozoic E (Externlobus), external lobe; abdominal lobe sensu HYATT (1867) ammonoids have a primary suture with three (synonyms: V=ventral lobe, lobes ("protolobes"), for which three symbols RUZHENTSEV, 1949, M=median lobe, WEDEKIND, are required and two more for lobes ("metalobes") 1913, which signified the medial parts of a bipartite which arise from the two saddles of the primary external lobe). This is the suture. Should several lobes originate in succes- first lobe, already found in bactritids and almost all sion in the same place, then the symbols are ammonoids, which appears in the vicinity of the designated by numbers according to their order siphuncle, and only in a few groups (Cly- of origin. In this way we can describe the onto- meniida, Pseudohaloritidae) does it lose its spatial genetic development of the suture from the connection with the latter. The external primary suture to the last septum quite adequately, lobe is cut by the median plane and lies on the using five symbols only. outer side of the spiral shell, spanning the The simple procedure stated can become com- space between the two external saddles. Loss plicated by the subdivision of single lobes, if these of the external lobe is restricted to the cly- divisions become independent lobes. If the newly meniids. formed part-lobes or whole lobes are clearly sep- L, lateral lobe (synonyms: G=Briickenlobus, arated by saddles, they can be designated accord- SCHMIDT, 1921; P=pleural lobe, SCHMIDT, 1952; 4 The University of Kansas Paleontological Contributions—Paper 47

0=omnilateral lobe, RUZHENTSEV, 1957; ex L parte: U=umbilical lobe sensu RUZHENTSEV, 1949, not U1 , U 2 , etc. sensu RUZHENTSEV; not L=lateral lobe sensu RUZHENTSEV, 1949). The second lobe to appear in the history of the ammonoids is similarly present in some bactri- tids and practically all ammonoids. In forms where the suture consists of E and L the latter is broad and occupies the area between the ex- ternal and internal saddles. A narrowing of the lateral lobe is already perceivable in Early forms, and this comes about either a as result of introduction of a broad internal lobe with wide saddles between I and L (e.g., Mimosphinctes), or as result of expansion of the external saddle and increase in involution with slow corresponding increase in the height of the whorls (e.g., Anarcestes, with Latanar- cestes as a transition, see Fig. 1). RUZHENTSEV L opposed this view and in his opinion the lobe is, in the first case, the omnilateral lobe and, in the second case, the umbilical lobe. How- ever, we believe that no morphogenetic founda- tion supports this view. I, internal lobe (synonyms: D=dorsal lobe, RUZHENTSEV, 1949; not /=internal lateral lobe sensu RUZHENTSEV, 1949). The third lobe in phylogenetic order of appearance is that cut by the median plane and which lies in the inner zone of the whorl. We encounter I for the first a time in advolute coiled forms (Mimosphinctes

can tabricus KULLMANN) ; in some it is replaced 2 by a later internal saddle. The following two symbols introduced by WIEDMANN (1966b), represent special cases of I and occur mainly in Mesozoic forms: b=lituid I, a simple, un- F frilled, double-pointed internal lobe (restricted to Phylloceratida); /,=septal lobe, bifid ex- tension of the internal lobe climbing up the face of the preceding septum (restricted to ).

2. Latanarcestes noeggerati (voN BucH), Middle De- vonian (Eifelian), Germany.-3. Anarcestes lateseptatus (BEyRicti), Middle Devonian (Eifelian), Germany. (All after Schindewolf, 1933, fig. 19, 25, 26; sutures drawn to same length). [The sutural development of Latanarcestes presents an intermediate stage between those of Mimagoniatites and Anarcestes. The lateral lobe of Latanarcestes is located laterally like that of Mimagoniatites whereas the corre- sponding lobe of Anarcestes lies more on the flank close to

FIG. 1. Ontogenetic development of sutures in genera of the umbilical seam. In RUZHENTSEV ' S terminology, the Anarcestina (Goniatitida). 1. Mimagoniatites zorgensis lateral lobe of Mimagoniatites is indicated by the symbol (RoEmER), Lower Devonian (upper Emsian), Germany. 0, and those of Latanarcestes and Anarcestes by U.] Kullmann & Wiedmann—Sutures in Phylogeny of Ammonoidea 5

The two lobes described next are those which A, adventitious lobe (Advent!' vlobus), WEDEKIND, originate through division of saddles. 1916 (synonyms: L=lateral lobe sensu Ru- U (Umschlaglobus), umbilical lobe, WEDEKIND ZHENTSEV, 1949; N=neolateral lobe, POPOV, (1913) (synonyms: K=Kehllobus, SCHMIDT, 1965). A lobe (or series of lobes) which de- 1921, and U=Umschlaglobus sensu SCHMIDT, velops from the external saddle. The formation 1921; (=internal lateral lobe sensu RUZHEN- of adventitious lobes pushes the lateral lobe TSEV, 1949, and LP, U 2 , etc., sensu RUZHENTSEV, back into an umbilical position, provided that 1949; not U=umbilical lobe without super- no further umbilical lobes are developed. script sensu RUZEIENTSEV, 1949). A lobe (or series of lobes) which originates from the saddle COMPARISON OF TERMINOLOGIES area between internal and lateral lobes, and USED BY WEDEKIND which then occurs in the vicinity of the um- AND RUZHENTSEV bilicus either on the outer or inner part of the

shell is defined as the umbilical lobe. The de- RUZHENTSEV ' S declared aim, in contrast to that velopment of umbilical lobes pushes the lateral of SCHMIDT, is a morphogenetical terminology. lobe from its umbilical position out onto flanks In some respects, however, RUZHENTSEV provided of the shell, provided that no adventitious lobes a morphographic-descriptive terminology, a fact are formed. All Mesozoic ammonoids possess that is evident only after careful analysis of his at least one umbilical lobe in their primary system (see SCHINDEWOLF, 1968, p. 835ff.). Un-

suture. In the course of their ontogeny a fortunately, scrutiny of RUZHENTSEV ' S viewpoint small number of Mesozoic ammonoids develop is made more difficult because of the polemics one or more lobes on the saddle between U, in his works (especially 1960, 1964). In the and I. Since these originate in the umbilical following we shall only touch upon some

area they are designated Un . more important points in dealing with the con- SCHMIDT and RUZHENTSEV have related the troversy between WEDEKIND and RUZHENTSEV. umbilical lobe to shell morphology. SCHMIDT RUZHENTSEV ' S bias toward a descriptive view- differentiated between lobes which originate point of sutures is evident from the importance from the internal saddle and develop externally attached by him to the mere names of symbols. (U) or internally (K). The latter can shift to He substantiated his use of the symbols L, U, and I the flanks of the shell during ontogeny. (of WEDEKIND) in a different context in that the RuzHENTsEv renamed WEDEKIND 'S L as U, but symbols chosen by WEDEKIND are much better distinguished between U sensu WEDEKIND suited for other lobes. In doing so RUZHENTSEV which he called 1, which always lies internally, failed to perceive that the importance of names and all other umbilical lobes. The sequence of from which the symbols are derived is irrelevant origin continues in RUZHENTSEV 'S terminology in a nomenclature based on morphogenetical sym- with U1 , U2 , etc., but in some cases with 11 , P, bols. Quite the opposite is the case as we assume, etc. WEDEKIND 'S terminology is advantageous from the outset, that the individual elements will in that the lobal sequence can be expressed change their position and shape in the course of clearly in a formula. For example, RUZHENTSEV ontogeny. For example, it is not significant expressed the formula of Protocanites as: whether a lobe is called dorsal, external, or ventral. VUU'/D, whereas WEDEKIND 'S formula is What is decisive is which part of the adult suture ELL12 11,1. The latter formula makes it possible can be traced back to the E lobe of the primary to read off fRuzHENTsEy's symbols in paren- suture. From a comparison of E in various forms these] that U,(/) is developed before U 2 (U,). we can work out its ability to change in the course The difference is even more noticeable in the of ontogeny and phylogeny, which, in turn, is case of some Mesozoic ammonoids. RUZEIEN- important for the systematics and phylogeny of TSEV described the suture of Normannites sp. as the form in question. follows: The principle of comparison of homologous (V,V,) (U 2 U,U 2 ) W/2 /2 ,1 3 , : /2 7 12 7 / 3 ,PD, structures in ammonite sutures was developed by whereas WEDEKIND 'S formula is: NOETLING and WEDEKIND half a century ago. ELU 2 U3 U 7 : U Ç U,Un /. The nomenclature is that of WEDEKIND. Ru- In this case the latter is not only shorter and zHENITsEv opposed this terminology, arguing that clearer but contains more information. the main three designations were incorrectly used:

6 The University of Kansas Paleontological Contributions—Paper 47

1) external lobe instead of "ventral lobe"; 2) in- "Oxygenium"=0, contrary to LAVOISIER, not all

ternal lobe in place of "dorsal lobe," and 3) lateral acids contain oxygen. RUZHENTSEV ' S objection lobe for "umbilical lobe" and "omnilateral lobe." that SCHINDEWOLF ignored the rules of priority In addition to what has already been said, that the in respect of U is unfounded. WEDEKIND derived name is unimportant for the meaning of a symbol, the symbol U from the word Umschlaglobus;

the following pertinent objections to RUZHENTSEV ' S SCHINDEWOLF later correctly latinized the German opinions may be raised: word for the sake of international clarity (Um- 1) The terms internal and external are un- bilikallobus). In so doing he did not alter the

related to the orientation of the ammonite animal, symbol. All in all, it follows that RUZHENTSEV ' S whereas the terms dorsal and ventral, when ap- renaming is in no way "a new type of nomen- plied to position of the last septum, can refer to clature which is substantially different from the various parts of the septum depending on length former one" (RUZHENTSEV, 1962, p. 271), but of the body chamber. For this reason HYATT rather a retrogression to earlier descriptive designa- (1867) proposed the name "abdominal lobe" for tions. Superfluous and homologous terms con- the external lobe. Having regard to MuryEl's tribute to disorder rather than aid in clarifying opinions regarding the orientation of Nautilus the modes of development of sutures. Future a renaming of V and D as anterior and posterior research in the field of computer-calculation of lobe ought to be considered. septal-area composition will be considerably handi- 2) As to the lateral lobe, RUZHENTSEV advo- capped by such terms.

cated that the lobe defined by WEDEKIND as lying In the following use is made of WEDEKIND ' S between internal and external lobes, called lateral terminology, because it is the older and, in our by him, is, in fact, an umbilical lobe (U) when opinion, the more logical system. In the interests it is small and situated near the umbilicus, or an of simplicity in scientific work the use of various omnilateral lobe (0) when it is broad and occupies symbols for one and the same object should be the entire lateral side. As RUZHENTSEV expressly avoided. Since, however, the younger terminology stated, 0 is later replaced by U, which means that of RUZHENTSEV has been used in some important they are homologous (see Fig. 1). Conversely, publications a conversion table of WEDEKIND ' S, U can develop into 0, as in Foordites, Pinacites, RUZHENTSEV ' S, and Popov's symbols is here in- and other genera (see SCHINDEWOLF, 1968, p. 841, cluded (Table 1). fig. 474-477). This demonstrates that the "am- TABLE 1. Comparison of some ammonoid suture- nilateral lobe" and the "umbilical lobe" sensu symbol terminologies. RUZHENTSEV represent homologous, heteromorph ISctimarrs terminology is not included because it is specifically variations of the same lobe. A renaming of L morphographicl as 0 and U is therefore not justified. WEDEKIND RUZHENTSEV POPOV E(Es ) V V 3) The priority of WEDEKIND ' S symbols is Es, Es, . indisputable, because in a morphogenetic termi- L 0 or U (not LP, U2 , . . .) nology, as stated above, it can only be a question I D D lz (D,10,) of symbols, since these represent homologous j , D units. The appropriateness of a name does not A (As) L As, A,,. L', I.,2 , . . . affect its priority: chemical symbols are not al- U (Us) I tered when the name of a substance is found to U2, v„ ti', r.P, . . . be inappropriate. For example, in the case of 1.1, 1' (rarely: P or following)

SUTURAL DEVELOPMENT IN LARGER SYSTEMATIC UNITS PALEOZOIC AMMONOIDS Anarcestina). These occur typically in the upper part of the Lower Devonian. The most primitive ORDER GONIATITIDA Lower Devonian genera exhibit EL, whereas the The starting point of sutural development in advanced Middle Devonian forms have the four the superorder Ammonoidea is a suture character- elements ELUI. Maenioceras (upper Givetian) ized by the elements ELI, which can be found has the most complicated suture EALU2 U 1 I (Fig. in the adult mature suture of forms of the super- 11,a) and in contrast to the tornoceratids only L family Anarcestaceae (order Goniatitida, suborder is on the flank of the whorl, since U 1 (and U2?) Kullmann & W iedmann—Sutures in Phylogeny of Ammonoidea 7 are formed before A. In a side branch (Augurites, ontogeny is valid for all later members of the Fig. 11,b) the external lobe appears to be split up suborder: Primary suture ELI, then (usually 2nd into E 1 E2 EmE2 E 1 . On the basis of their sutures suture) EAU, and later EALUI. The formula RUZHENTSEV considered the Agoniatitidae (4- EALUI occurs most commonly within the super- Mimoceratidae) and Anarcestidae to be separate family Cheilocerataceae (including Tornoceratidae suborders. In his terminology the Agoniatitina and Pseudohaloritidae), i.e., the most primitive have an "omnilateral lobe" and the Anarcestina . An increase in the number of ele- an "umbilical lobe." The sole difference between ments follows from an increase in the adventitious "0" and "U" is that the questionable lobe L in lobes (e.g., Discoclymenia: EA 3A 2 A 5 L:U1), Anarcestes and younger members of the family (Fig. 11,f). lies near the umbilicus, in contrast to the Agonia- titidae (Fig. 1). The oldest anarcestids are inter- mediary forms where, in all stages (as in Mima- h ) L is broadly developed on the whorl flank. In younger anarcestids L is restricted by the external saddle so that, combined with the greater involution of the shell, L lies in most Anarcestidae near the umbilicus. Therefore, evi- dently no genetic difference distinguishes "0" from "U" sensu RUZHENTSEV. The superfatnily Pharcicerataceae also belongs to the suborder Anarcestina, its characteristic difference from the Anarcestaceae being the de- velopment of a median saddle and lobe in the external lobe. The basic sutural pattern is d (E,E„E l )LI, as in Ponticeras. Within both of the Gephuroceratidae and Pharciceratidae one or more umbilical lobes occur: Manticoceras (E I EE 1 )L:U1 (Fig. 11,c); Timanites: b • LU2 U3 :U,I; Pharciceras: (E,Em EOLU 2 U: U3 U,I; Synpharciceras: EmEILU2UU6U8U10: U9 U7 U5 U3 U 1 1; Neopharciceras up to U24. In the a family Beloceratidae there is, in addition, a FIG. 2. Ontogenetic development of suture in Marathonites marked division of the external lobe: (Almites) invariabilis RUZHENTSEV, Lower (lower ExE„,B,LU,I (e.g., Mesobeloceras: Artinskian), USSR (southern Urals). (After Ruzhentsev, Em E3 E2 E 1 LU2 U3 U5 U 7 :U8 U 6 U 4 U i l 1956, fig. 89a-f, i, 1). [Explanation: a, prosuture; b, pri- d-g, intermediate-stage sutures; (Fig. 11,d), Beloceras up to E7 and U 18 ). The mary suture; c, 2nd suture; primary suture is trilobate; the 2nd use of formulas in these cases is somewhat re- I,, adult suture. The suture has an adventitious lobe that during ontogeny, like stricted owing to asymmetry and irregular bifurca- the umbilical lobe, divides into 3 lobes.] tion of the suture. The Prolobitaceae represent the third super- Other superfamilies differ from the Cheilocera- family of the suborder Anarcestina. Their suture taceae principally in that the external lobe has a appears to be made up of ELI and ELU / (/,/,) median saddle and lobe. Their sutural formula (Fig. 11,e); however, the group is poorly known. is (Ei E„„E i )ALUI (Fig. 11,g). The superfamily In the Anarcestina lobai increase usually takes Goniatitaceae has the basic suture from which place in the umbilical area, if not also in the area sutures of other Goniatitina superfamilies differ- of the external lobe. The suborder Goniatitina, entiate. By and large, the families , however, is characterized by the appearance of a Gastrioceratidae, and Paragastrioceratidae retain first adventitious lobe (A) prior to formation of the formula (E I E„,EJALUI. A few families an umbilical lobe (U). This means that the lateral (Metalegoceratidae and Schistoceratidae) show a lobe remains in the umbilical area (Fig. 2). The tendency toward trifid division of some lobes. simplest sutural formula is, therefore, EAUI This is typical of the superfamilies Agathicera- (, Cheiloceras). The following lobai taceae and Cyclolobaceae. The Metalegoceratidae 8 The University of Kansas Paleontological Contributions—Paper 47 subdivide the lateral lobe L into L,,L,„Ld (Fig. 12,d). The family Thalassoceratidae has no addi- 11,h) [the end form being Pseudoschistoceras tional lobal elements. In contrast the family (Ei EmEi )AL,, (L,,,,,Lm :Lmd)LdU (Fig. 11,i) ac- Dimorphoceratidae often exhibits irregular sub- cording to RUZHENTSEV, 1962 1 . The Schistocera- divisions of the external lobe and on occasion also tidae subdivide U2 into U2v U2 .U 2 d (end form of the adventitious lobe. Examples are Asturoceras Schistoceras EmEi ALU 2,,U2 m :U2aU JO. (EidElvdEn2 EmEi .,,,E lydEld)AL:U1 (Fig. 12,c) The Agathicerataceae similarly exhibit a ten- and Paradimorphoceras dency toward trifurcation, in this case of the (EidEp,,EmEn„E ld )(A vA d)L:UI. adventitious lobe and partly of the external lobe: The superfamily Adrianitaceae, in form and 1) the Delepinoceratidae display incipient trifur- lobai arrangement very similar to the Agathicera- cation of E l and A [Delepinoceras taceae, increases its lobes by the formation of (E i dElmEi „E„,Eft,ElmEld)(A,A„„A d)L:Ull (Fig. additional umbilical lobes. Characteristic of many 11,k); 2) in the Agathiceratidae only A trifurcates Adrianitaceae is the breaking down of a lobe in [Proshumardites (E l Em Ei)(A vA mAd)LUI; the the region of the umbilicus into numerous in- end form is Agathiceras (El EmEi )A,,A„,A dLUI] cisions, to which we can give no names. This (Fig. 12,a). is referred to as Suturallobus (S) by WEDEKIND. The superfamily Cyclolobaceae is characterized Examples are Emilites (E 2 E„,,E1 )AL(U2 =S)U 1l by trifurcation of all lobes except E. Examples and Crimites (E i E„,EJALUd (U 4 =S)U d UiI of the family Shumarditidae are Shumardites (Fig. 12,e). [EmEi A,A m A dL : (Uvt LnU (111m11)] and Pro- In RuzHENTSEV'S terminology Suturalloben perrinites 1(ElEmEi)ily24.(ilavAa2) LvL. La (S), which are lobes appearing simultaneously, are (U,, 2 U,.d)Um Udg given identical symbols [e.g., Cri in The same scheme of lobai development pre- ,V OLUWWW:U 2 U'ID]. This means, in vails in the families Marathonitidae and Cyclo- the cited example, that U4 represents two differ- lobidae. Lobai multiplication results from re- ent lobes. This method of notation breaks down peated trifurcation of A and U, while the lateral in complicated forms where differences can occur lobe remains single trifid. Examples of the family within a species or even in an individual in one Cyclolobidae are Vidrioceras 1(E 1E mE 1) particular asymmetrical septum (sec RUZHENTSEV, AvAmAaLvLmLd:UrUmUd(111a 1)1, Stacheoceras 1956, fig. 94-95). Such irregularities are evidence [(EiEmEr)AvilmilarAa2vAa3rAav of abnormal lobai increase and are expressed best (14d5vAd)LvLtnLd:UvUr3dUv2dUrrlUniUd by use of the additional symbol (S), used in con-

(1 1 1 77111)1, junction with the usual lobe symbol. and Waagenoceras The oldest forms of the suborder Prolecanitina (EiEmE1)AvAmAdvA d2vAd3vAdiv(A d5rAdfi) are similar to the . The suture of L,Lm :Ld U d U v2d U dr_1 mU 2(1 11.1 1)1• Protocanites possesses the same number of ele- The family Popanoceratidae is basically similar ments as that of the contemporaneous Gattendor- except for the insertion of further adventitious fia. The origin of their five lobes is different, and umbilical lobes as in Popanoceras however: in Gattendorfia EALUI and in Proto- (ElEmEl)A.tvAimAiar(Ala2vAnts)242LU2: canites ELU 2 U 1 I. It is by no means certain that U3 (U1v2U ivd)U ImUidg lobai increase occurs uniformly in the entire sub- (Fig. 12,b); in P. sobolewskyanum, teste Ru- order through insertion of umbilical lobes. ZFIENTSEV (1956), the sequence U2 and U, cannot ScHtivnEwoLF (1951, p. 22, fig. 14) reported the be verified with certainty. These similarities are presence of an adventitious lobe in Neopronorites of special importance for the systematics of these permicus—an interpretation which cannot be three families as they prove their relationship to proved, since the trilobate primary suture in this one another. form is immediately followed by a quinquelobate In the superfamily Dimorphocerataceae we suture. The absence of an adventitious lobe has find a totally different lobai configuration. The been demonstrated clearly for Merocanites asiati- family Neodimorphoceratidae is similar at first cus (KARPiNsky) which has the sutural formula to the family Goniatitidae, but differs in that the ELL 2 U ,I (Fig. 3), but the possibility exists number of lobes of its suture increases through that the lobe taken to be L in other Prolecanitina differentiation of the external lobe. The end form is in fact the lobe A. is Neodimorphoceras (E 1E2E,,,,E 2E1 )AL:UI (Fig. The basic family Prolecanitidae shows an in- Kullmann & W iedmann—Sutures in Phylogeny of Ammonoidea 9 crease in U from Prolecanites [(ELU 2 U 3 L :11,1) (E1E.Ei)(Lv2Lv4L4) U2U3U4U5 • • • U6U1 (MI) (Fig. 12,f)] up to U 7 in Acrocanites. In the (Fig. 12,g) ], Uddenites [(E 1 EmE1 )(-L4) Daraelitidae and Pronoritidae the number of um- U2 Us LI 4 U 5 U 7 . . 1, and Medlicottia [(E1 E,,,E 1 ) bilical lobes is increased even more and, in addi- (-L4U2 U3 U 4 U 5 U 7 . . . 1. tion, E is further developed in Daraelites The superfamily Sagecerataceae is character- [(E 1 E,,,E 1 )LU2 Us U 4 (1 5 ...Us ti :1 (111 i ) up to U9 ] ized by elaboration of the external lobe, as in and Neopronorites [(E I E„,E 1 )(LyL4)U 9 U3 U 4 ... Pseudosageceras [E.E 4E3 E2EILU 2 U3 U 4 U 6 • • • 1.1 3 U 1 (1 1 1 1 ), up to U.to]. U5 L1 1 (1 1 1 1 ) (Fig. 12,h)] (see B000sLovsKrY et al., 1962). The lobai development of some isolated groups systematically close to Praeglyphioceras, Kara- gandoceras, and Prodromites remains unexplained. In these a median lobe develops in the external saddle (E7 E.E7 ). The first two genera belong in the suborder Goniatitina, since the lobe lying laterally appears to be adventitious. The suture of Prodromites, on the other hand, has the formula E,,,E,LU2 U3 . . . and, therefore, probably belongs in the suborder Prolecanitina. It is possible that h these are independent, restricted groups which existed for a short time and left no successors.

ORDER \fmr—isik-v- Members of the order Clymeniida are restricted to the uppermost Devonian and are close to the e Anarcestina. The sutural formula ELI occurs here, too, and further, as in the Anarcestina, lobai d increase occurs primarily in the umbilical lobe and secondarily in the adventitious lobe. c The greatest similarities with the Anarcestina are found in the suborder Gonioclymeniina. Lobai b development proceeds from ELI via ELUI and EALUI to EA 2A 1LU 1 U2 I (Sphenoclymenia) a (Fig. 12,i). Some families develop a median saddle in the external lobe (e.g., Biloclymenia) FIG. 3. Ontogenetic development of the suture in Mero- (EI E„,E 1 )LU2 U 1 I. Among the Parawocklume- canites asiaticus (KARPovsKiv), Lower (lower Visean), USSR (from Karpinskiy, 1896, P. 187). The riaceae a lobai reduction occurs from (E I EOL: primary suture is trilobate; later on a 4th lobe is developed (I i l i ) to a single L. (umbilical lobe), which pushes the lateral lobe from its The suborder Clymeniina includes the typical original umbilical position out onto the flanks. [a, pro- clymeniids where the external lobe is reduced in suture; b, primary suture; c-i, further stages of develop- the adult stage. The basic suture is L:I, and ad- ment; k, adult suture.] vanced forms have either AL! or LUI. Cymacly- menia (AL/U!) (Fig. 12,k) is the most compli- The superfamily Medlicottiaceae retains the cated genus of the suborder. sutural development of the Prolecanitaceae with modification of the external saddle in that parts of the lateral lobe lying near the external lobe MESOZOIC AMMONOIDS become included in the external saddle. The Viewed broadly, Mesozoic ammonoids are incisions which result cannot be traced mor- characterized by a progressive development of the phogenetically and commonly are not generically, suture. This involves both denticulation of the or even specifically, typical. Furthermore, they adult suture and number of lobes in the primary may be asymmetrical. RUZHENTSEV ' S notation, suture. The first, as is well known, proceeds therefore, would appear to be superfluous. from the unipolar "ceratitic" frilling of the lobes Examples are Prouddenites of most ammonites to the bipolar "am-

10 The University of Kansas Paleontological Contributions—Paper 47

trilobate quadrilobate quinquelobate sexlobate

E L U I E L U, U, I E L U, U, U, I

A \:.> E L U, U, I

E L U I .77

; E L I 1 .5 0 f

Fie. 4. Progressive and regressive evolution of primary sutures in ammonoids. monitic" frilling of the lobes and saddles of as quinquelobate, forms (all - Jurassic and Cretaceous ammonites. The primary phylloceratids), perhaps even trilobate forms (the suture (Fig. 4) of the Triassic ammonites has Permian ). Whether in this stem gained an extra element over its Paleozoic fore- group of Mesozoic ammonites the change of the runners, and thus has become quadrilobate primary suture coincides with the system bound- (ELL 1,1). In Jurassic and Cretaceous ammonoids aries is unknown, since the Triassic phylloceratids, the primary sutures are quinquelobate like all Triassic ammonites, have not been investi- (ELU 2 U 1 1), while in the progressive lytoceratids gated sufficiently. Since, however, the unity of of the Late Cretaceous, the tetragonitids, a sexlo- these forms is indicated by other characters, such bate primary suture is present (SCHINDEWOLF, as form of the internal lobe and phylloid saddles, 1968). it would be unwise to overrate this continuous Within both trends reversions appear, espe- transition within the phylloceratids. cially in the Cretaceous, and thus a return to The same is true for the sexlobate tetragonitids, "ceratitic" or even "goniatitic" sutures (e.g., Neo- which are connected with the lytoceratids not only lobites), as well as reduction of the primary suture by a continuous transition (WIEDMANN, 1962a), of all Cretaceous heteromorphs and "false hop- but also by a major character (e.g., form of the litids" (WIEDMANN, 1966a,b) to the quadrilobate internal septal lobe) (WIEDMANN, 1966a,b; 1968). form of the Triassic ammonites (WIEDMANN, Moreover, since the sexlobate primary suture is 1963, 1965, 1966a). As a basic rule, augmentation apparently first formed within the tetragonitids of the lobes of Mesozoic ammonites occurs, when themselves, it seems more reasonable to regard not simply by lobai splitting, through the building this group, too, as a superfamily within the of umbilical lobes ("U type" of SCHINDEWOLF). Lytoceratina (WIEDMANN, 1962a) rather than as Adventitious lobes (A lobes) are only formed, a separate suborder (SCHINDEWOLF, 1967b). if ever, at maturity (Arcestidae, Sphenodiscidae). While the Ceratitina are generally admitted The consequent advantage of using configura- to be a natural unit, which SCHINDEWOLF (1968) tion of the primary suture for large-scale system- believed could be linked confidently with the atic division of the Mesozoic ammonites deserves , the derivation of the Jurassic- close attention (Fig. 4). If we treat the Phyl- Cretaceous from the Lytoceratina loceratina in the manner of SCHINDEWOLF (1968) or Phylloceratina still presents great difficulty. as the root forms of all Triassic ceratitids, as well The previous view supported by Luppov & as the Jurassic-Cretaceous ammonites, and accept DRUSHCHITS (1958) that the bifid or trifid form their stratigraphie range as Late Permian to the of the lateral lobes indicates derivation from the end of the Cretaceous, then only this suborder lytoceratids or phylloceratids has become untenable includes quadrilobate (e.g., Leiophyllites), as well following the proof of both ontogenetic (WIED- Kullmann & W iedmann—Sutures in Phylogeny of Ammonoidea 11

MANN, 1962a, fig. 53) and phylogenetic transition terizes the Ammonitina and , which (CossAN, 1952, fig. 3) from one lobe form to the therefore occupy a kind of intermediate position other, and the discovery of both lobe types on between phylloceratids and lytoceratids (Fig. 7). either flank of one suture ( WIEDMANN & DIENI, If, as suggested by SCHINDEWOLF (1968), the 1968, fig. 45). This is similarly true for the so- Ammonitina and Ancyloceratina are included called Suturallobenbildungl (WEDEKIND, 1916), with the Lytoceratina in a second order, Lyto- which, according to early opinion of SCHINDEWOLF ceratida (although the phylogenetic connection (1923, 1929, 1961) favored assignment to the of these groups is not yet clarified), then the phylloceratids s.l., but which in the meantime has sutural features mentioned above lead to the been found in both the lytoceratids (WIEDMANN, following diagnoses of the Mesozoic orders and 1963) and desmoceratids ( WIEDMANN, 1966b). suborders. [Here, and in following parts of the In place of Suturallobenbildung, the internal text, primary sutures are marked by an asterisk suture, in particular the internal lobe, has assumed (*)-] importance in this question. SALFELD (1920, 1924) drew attention to presence in the phylloceratids Diagnoses of Mesozoic Orders and Suborders of a simple ( unfrilled), double-pointed lituid I, 1. Internal lobe lituid (II) Order PHYLLOCERATIDA but himself doubted the importance of this feature 1) Primary sutures: *ELI (Permian)—> because he believed to have observed an iterative (Triassic)-9 *ELU2U// (Jurassic- transition between the lituid / of the phyllocera- Cretaceous) Suborder PHYLLOCERATINA tids and the normal / of the Ammonitina. This 2) Primary suture *ELL/il .... Suborder CERATITINA was rejected by SCHINDEWOLF (1962) for Jurassic II. Internal lobe not lituid Order LYTOCERATIDA forms and by WIEDMANN (19626, 1966a,b) for 1) Primary suture *ELLI,U,I (Lytocerataceae) Cretaceous ones, so that today the lituid internal —> *ELU2U3U1/ (Tetragonitaceae). Septal lobe (//) can be regarded as the most reliable lobe (10 Suborder LYTOCERATINA 2) Primary suture *ELU2iIil. Internal lobe phylloceratid suture and also of character of the simple (I) Suborder AMMONITINA that of the bulk of the Ceratitina. It is thus 3) Primary suture *ELUL Internal lobe reasonable to unite both these form groups in the simple (I) Suborder ANCYLOCERATINA order Phylloceratida as recommended by SCHINDE- WOLF. Above all, such a procedure simplifies Since in Mesozoic ammonoids the external classification. lobe of the adult suture is always bifid, in Jurassic

SCHINDEWOLF ' S (1968) recommendation to as- and Cretaceous ammonoids already that of the sign subordinal rank to the pinacoceratids, arces- primary suture, it is simpler to leave out the tids, and lobitids is not accepted by us, pending formula E i E i , although this may usefully be more detailed investigations of the ontogenetic applied for Paleozoic forms. The differences in lobe development of Triassic ammonites. ontogenetic lobe development are of importance Exceptions, with a denticulated /, occur within for the further division of the Mesozoic am- the thus defined Phylloceratida only in the arces- monoids. tids, ptychitids, and tragophylloceratids, but all are phylogenetic end forms which contribute noth- ORDER PHYLLOCERATIDA ing to the development of the Ammonitina. The The Otocerataceae, the root stock of the lecanitids, celtitids, and choristoceratids with a Mesozoic ammonoids, exhibit a lobe develop- single-pointed / also do not completely fit into the ment of the formula *EL(U / )/—>ELU 2 :U / / i normal pattern of the Phylloceratida. (Fig. 13,a) ---> ELU 2 U3 U5 : UJJ,/, (Fig. 13,c). In contrast to the above forms in the lytocera- Therefore, they are not true phylloceratids in tids s.str. the septal lobe appears in evolutionary SCHINDEWOLF ' S (1961) previous interpretation history very early (i.e., in the earliest Liassic). since they lack the so-called Suturallobenbildung. This feature, according to our present knowledge, SHEVYREV (1968), possibly as a result of similar is exclusively characteristic of the Lytoceratina, observations, suggested a new suborder Paracel- in which the 1, can be contrasted with the titina. The internal lobe of these forms is already of the phylloceratids. On the other hand, a lituid, however, so that their inclusion in the denticulated, single- or double-pointed / charac- Phylloceratina is justified. 'This is the more or less symmetrical lobe splitting at From their lobe formula and the little informa- the umbilical seam (see Fig. 5). tion available on Triassic ammonoids (BRANco, 12 The University of Kansas Paleontological Contributions—Paper 47

h

U2 U3U4 U7 I

d

U2U3 Ul

E L U2 U3UI I

E L U2 LII I

-\,..T\/-s-I'W

FIG. 5. (Explanation on facing page.) Kullmann & Wiedmann—Sutures in Phylogeny of Ammonoidea 13

1879; HYATT & SMITH, 1905; SMITH, 1914, 1927, late I (without septal lobe) and the greater degree 1932; SCHINDEWOLF, 1929, 1961, 1962, 1968; of suture frilling (especially of the saddles) of SHEVYREV, 1961, 1962, 1968; RUZFIENTSEV, 1962; Trachyphyllites are different from the suture of ZAKHAROV, 1967; WIEDMANN, 19696) the following palaeophyllitids. groups can be attached to the Otocerataceae: Suturallobenbildung in Us or U4 (Fig. 5), Ussuritidae within the phylloceratids (Fig. 13,e); regarded by SCHINDEWOLF as typical of the phyl- Meekocerataceae including the bulk of the former loceratids, first occurs in younger phylloceratids, "Noritaceae"; Clydonitaceae (Fig. 13,f); Trop- the Discophyllitidae (incl. ) and itaceae; Proptychitidae within the "Proptychita- , and these families continue un- ceae" of SHEVYREV; and finally also the Ceratita- changed to the end of the Cretaceous. The lobe ceae including the "Hungaritaceae" of SHEVYREV formulas ELU 2 (U 3 = S) U1 11 and ELU 2 Us (Fig. 13,h, 14,a). In the last group an intensive (1/=_S)U 1 /c (Fig. 15,a) are varied only slightly proliferation of the umbilical lobes as far as U, in through the weak frilling of or even U,, takes place, which makes a separation I as mentioned above. On the whole, the phyl- of these "Ceratitina" from the Phylloceratitina, loceratids represent an exceptionally conservative as suggested by SCHINDEWOLF, possible but not group extending from the Carnian to the Maas- necessary. Also the Triassic heteromorphs (i.e., trichtian (except for progressive frilling of the Choristoceratidae, incl. Cochloceratidae) with the phylloid saddles). Where exactly the transition end formula ELU / / (Fig. 13,g) attach themselves from a quadrilobate to a quinquelobate primary easily to this group. suture took place is not known, although it must From this type of pure umbilical lobe prolifera- be in the neighborhood of the Triassic-Jurassic tion a second mode of differentiation of the um- boundary. bilical suture can be derived easily, and during It is noteworthy from their lobe formula the course of lobe ontogeny in this only one ele- ELU 2 (Us =S, 10:10)U / L (Fig. 14,c) that the ment is formed at the umbilical seam. However, Megaphyllitidae lie very close to the phylloceratids. this element does not, as in the above-discussed The Suturallobenbildung within U3 shows an ex- forms, originate as a new lobe in the saddle LU treme frilling with more than 20 incisions sym- (or U2 U,) but is always inserted on the ventral metrically distributed on either side of the umbili- lobe shoulder of U,. It is therefore reasonable to cal seam. This does not apply to Procarnites label this element U,,, (not U2 or Us ) and to which shows asymmetrical Suturallobenbildung contrast it with the earlier U m. The resulting within U 2 (e.g., Ptychophylloceras in SCHINDE- lobe formula EL(U,„:U /d )L (Fig. 13,i) character- WOLF, 1961, fig. 29, 30). This asymmetry can izes the Tirolitidae, Dinaritidae, and Hellenitidae be expressed in the formula ELU2 (Us =S, 7: within the "Ceratitaceae," and the Kashmiritidae 4)U,1, (Fig. 13,k). Procarnites, selected as type and Sibiritidae of the "Noritaceae," grouped as of a new family by CHAo (1959), was placed in the Dinaritaceae by SHEVYREV (1968). the Proptychitaceae by SHEVYREV (1968). Turning back to the Ussuritidae, some dif- The Ptychitaceae with their lobe formula ficulty arises in the interpretation of Leiophyllites ELU 2 (U 3 =S , 8:8)U,! (Fig. 14,e) can also be and the "Palaeophyllitidae" of Popov. Their lobe placed close to the forms mentioned above, espe- formula is somewhat intermediate and may be cially the Megaphyllitidae with their similarly interpreted as ELU2 Us sU I L as well as ELU2U/v: symmetrical Suturallobenbildung, differing dis- U m!, (Fig. 13,d). Probably here, or within the tinctly in the stronger frilling of the saddles and Dinaritaceae, is the point where Lytoceratida with internal lobe. These characters associate the their Triassic forerunners, the Trachyphyllitidae ptychitids with the Arcestidae, in which, however, (WILDNIANN, 1966a, 1968), were derived. They no Suturallobus appears to be present. Instead, portray the lobe formula ELU2 (//,,,:U m )1 (Fig. a quinquelobate primary suture (SexitynEwOLF, 14,1) and thus in the Triassic, the morphological 1929) and the formation of adventitious (A) and difference between the lytoceratids and the primi- internal umbilical lobes (U3 ) in the saddle U,/, tive phylloceratids is very small: only the denticu- otherwise unknown in Triassic forms, occurs

FIG. 5. Suture ontogeny and Stiturallobenhildung in U, of phylloceratids: Sotverbyceras (Holcophylloceras) calypso

(DORBIGNY), Lower Cretaceous (), France (from Wiedmann, 1968). [a, primary suture; j , adult suture at whorl height of 3 mm.] 14 The University of Kansas Paleontological Contributions—Paper 47

(SCHINDEWOLF, 1968). The arcestids, with their 1957, 1958, 1960, 1963; BEZNOSOV, 1960; SCHINDE- resulting lobe development WOLF, 1961-68; WIEDMANN, 1962a, 1963, 1965, *ELU 2 L/ 1 J--->EALU2 U 3 U / U„,,Un d/ 1966a,b, 1968, 1969a) were already present in (Fig. 14,d), stand markedly apart from the other- Triassic forms. wise very uniform character of the Triassic am- monoids. ORDER LYTOCERATIDA Similarly, the Lobitidae, with lobe-splitting in As already mentioned, the formation of the the elements L, U2 and U/ according to the septal lobe in I (Is ) is a qualitatively new feature formula E (L„Ld) (U2 ,U 2a) Us : U/ v Uidb (Fig. in the lytoceratids which is found occurring as 14,b) also occupy a special position. early as the early Liassic. The lobe formula of phylogenetically The lobe development of the the , ELU 2 (U 1v : U1 d) Is (Fig. 6, unimportant pinacoceratids unfortunately has not 14,g), can be derived easily from the Triassic been sufficiently investigated. Trachyphyllitidae. The same is true for the sex- Thus the most essential types of lobe formation lobate primary suture of the Tetragonitaceae, found among Jurassic-Cretaceous ammonites and direct derivatives of the Lytoceratidae. In this about which much new information has recently case the formation of a massive suspensive lobe been published (especially by MIKHAILOVA, in U1 occurs first in the (Fig.

E L U2U1 C

FIG. 6. Suture ontogeny of true lytoceratids with I, and subdivided U,: juilleti (D'ORBIGNy), Lower Cret- aceous (Valanginian), France (after Wiedmann, 1968). [a, primary suture; e, adult suture a whorl height of 2 mm.; stippled, septal lobe.] Kullmann & W iedmann—Sutures in Phylogeny of Ammonoidea 15

14,h), and proceeds to a Suturallobenbildung of the Aegoceratidae. The Hammatoceratidae, in U4 in the (Fig. 14,i), indicating Paroniceratidae, and Dumortieriidae with the lobe convergence with the phylloceratids (WIEDMANN, formulas ELU 2 L 31-11 :(UiU 1 4)1 (Fig. 15,f) 1963): ELU 2 U 1 /8-3ELU2 U3 (U 4 .=S, 2:2)U,/8. —>ELU 2 U 3 U 4 :U 1 / were regarded as the root of

According to SCHINDEWOLF ' S (1962) most re- the Hammatocerataceae, derived from Dactylio- cent investigations, the psiloceratids, which repre- ceratidae still having a split U 1 . These opinions sent the root stock of the Jurassic and Cretaceous of SCHINDEWOLF resulted, moreover, in the sub- Ammonitina with quinquelobate primary suture, division of the previously homogeneous Haplo- have no Suturallobenbildung. Instead, a very early cerataceae: the oppeliids with an undivided U, division of U 1 occurs. The resulting lobe formulas being placed in the Hammatocerataceae while ELU 2 Us :U,,,U, d/—>ELU 2 U3 UL/ 5 :U,,,U, d/ were forms with (according to SCHINDEWOLF) a "sec- derived by SCHINDEWOLF (1962) from an un- ondarily undivided" U ., were grouped as Haplo- known lytoceratid root. More important than cerataceae. These comprise the bulk of the grapho- the as yet not fully clarified question of the root ceratids, strigoceratids, and sonniniids, as well of the psiloceratids—at or near the point of di- as the true haploceratids and a large part of the vergence of the lytoceratids from the phylloceratid Cretaceous oppeliids. Perhaps a simpler classifica- main stock—is the fact that all younger Ammo- tion would be based on the assumption that a nitina can be easily linked to the psiloceratids. repeated oscillation between an undivided and

SCHINDEWOLF ' S studies of lobe ontogeny thus in- divided U, occurred (Fig. 7). Certainly, oppeliid dicate that the Ammonitina form a monophyletic and haploceratid sutures are closely similar in unit. their strong proliferation of lobes, but the differ- The lobe formula of the Eoderocerataceae, ing fate of U, seems to be a radical point of dif- ELU 2 U1 :U,,U, 4 (Fig. 15,c), is so closely similar ference which can be seen in the lobe formulas to that of the Psilocerataceae that SCHINDEWOLF which for are ELU 2 Us Ur-7 5 (.7 7 : even considered uniting these two superfamilies, U8U6U1rUl41->ELU2USUUSU7U9U1 IU18: thought by SPATH (1938) and ARKELL (1950, U, 2 U,J.J 8 U 6 U 1 ,U, d/ (Fig. 15,g), and for Hap- 1957) to be diphyletic. Perfect identity of sutures loceratidae ELU2UsU5E17(18:U6UU11 (Fig. is found among the , Polymorphiti- dae, Amaltheidae, and Dactylioceratidae. While the suture of the Phlycticeratidae In addition, fusion of the ventral and dorsal (ELU2 L/ 3 :U,,U,,d) can be compared closely with part of U, may occur in both superfamilies (in that of the oppeliids, those of the Strigoceratidae,

SCHINDEWOLF ' S opinion always secondary fusion, Mazapilitinae, and Aconeceratinae are identical i.e., in the , Echioceratidae, Oxynoti- with that of the haploceratids: ELU 2 Us Us :U 4 U,/ ceratidae, Phricodoceratidae, and Aegoceratidae (Strigoceratidae), ELLI2U3U5U7(19118U6LIE111 [including Androgynoceras and Oistocerasl), all (Aconeceratinae). of which have the common lobe development Whereas the oppeliids also were traced directly *ELU,U,/—>ELU 2 U3 :U,/—*ELU 2 :U,1 (Fig. back to the hammatoceratids by SCHINDEWOLF, 15,d). Additionally, as a slight modification of the haploceratids were derived from the dumor-

SCHINDEWOLF ' S views, Suturallobenbildung may tieriids already discussed, via the Graphoceratidae occur in U,, according to the formula and Sonniniidae with the lobe formulas ELU,(U,„=S)U, d/ in the psiloceratids and ELU2 Us U 5 :(7 6 UU,/ eoderoceratids (i.e., in the Cymbitidae and the ELU 2 U 3 U 5 U 7 U 9 U,,U, 0 U 8 U 6 UU 1 / , for which SPATH, 1923, probably (see Fig. 15,i). The boundary between the two erroneously, assumed a phylogenetic relationship). superfamilies divides the graphoceratids with Basing his opinion on the yet unproven assump- "primary divided" U, (Dumortieriidae) from tion that a secondarily fused U, may not split the graphoceratids and sonniniids with "secondary again, SCHINDEWOLF (1964) believed the Hildo- undivided" U,. In contrast the Tmetoceratidae cerataceae to be diphyletic. He restricted the with a probable lobe formula ELU2 U,..1 (see Fig. superfamily to include only the Arieticeratidae, 15,h) were regarded as end members (with un- , and Harpoceratidae having the divided U,) of the Hammatocerataceae, compar- lobe development ELU 2 U1 :U i /—>ELU2 U 3 U5 : able to the Arietitidae, Echioceratidae, or Aego- U U,./—>ELU 2 U 3 U 5 U 7 U 9 U 11 U 12 :U 10 U 8 U 6 UU,/ ceratidae of the Psilocerataceae. (Fig. 15,e) and placed them as the descendants The Otoitidae, earliest representatives of the

16 The University of Kansas Paleontological Contributions—Paper 47

LYTOCE RATI NA ANCYLOCE RAT I NA AM MON ITINA PHYLLOCERATI NA

U I

Dol1111//e/O. u

FIG. 7. Phylogeny of elements / and Ui within Jurassic-Cretaceous ammonites (after Wiedmann, 19666). [Psiloc.= Psilocerataceae, Hap/oc.=Haplocerataceae, Hop/itac.=Hoplitaceae, Ancy/oc.=Ancylocerataceae, Donvilleic.=Douvillei-

ceratidae, Astieric.=- Astiericeratidae.]

stephanoceratids, also show a division of U / , in- that from Hammatocerataceae (SCHINDEWOLF, dicating their origin in the Hammatocerataceae. 1966). This division is already reversed in the almost In contrast to the very homogenous Peri- simultaneously appearing Stephanoceratidae and sphinctaceae, the Stephanocerataceae show a nota- Sphaeroceratidae, just as in the bulk of the bly large variation in con figuration of the suture. Stephanocerataceae. Nevertheless, all these forms The original otoitid suture ELU 2 U3 :(U /n U 1 4U„/ are united by the possession of a Un unique to (Fig. 16,a) —> ELU2 U3 U:U 5 (U /y :U id )U„/ the stephanoceratids. Consideration of only the ELU 2 U 3 U1 :UU 1 Un/ is replaced in the majority adult suture is insufficient for the understanding of the stephanoceratids (i.e., Stephanoceratidae, of these sutures (Fig. 8). Sphaeroceratidae, Macrocephalitidae, Pachycerati- In labelling the lobe peculiarly situated in the dae, Oecoptychiidae, Kosmoceratidae, Cardio- saddle U 1 /, as "Un " SCHINDEWOLF (1965) retreated ceratidae, Morphoceratidae) by the lobe formulas from his earlier (1923) interpretation of this ele- ELU 2 U3 U,:U i U 1 U,/ (Fig. 16,b)-->ELU 2 :Us U 1 1 ment as a "heterochronous U./ ." Study of the nu- (Fig. 16,d). In the pachyceratids and the oecopty- merous lobe pictures published by SCHINDEWOLF chiids the division of U 1 , as well as the U„, are raises the suspicion that this element, almost always reduced. Also, for the Parkinsoniidae a lobe (except in the Otoitidae) found on the lobe formula ELU 2 U ,3 1 ; :t1 (Fig. 16,e) was shoulder of an undivided U 1 , does not correspond assigned and thus supported their attachment to to a heterochronous U 1 , but may perhaps be a the Stephanocerataceae (SCHINDEWOLF, 1965), al- heterochronous U id. This would explain why a though in this case the identity of Un and U ld U„ is never found in the closely related Peri- (e.g., op.cit., fig. 288) and thus the possibility of sphinctaceae, with an almost universally present a perisphinctid formula ELU2 U8 1/ 5 :UU / „U id/ divided U / . Thus derivation of Perisphinctaceae cannot always be denied confidently. The Spiro- from Stephanocerataceae seems more likely than ceratidae (incl. Acuariceratidae) with their curious Kullmann & Wiedmann—Sutures in Phylogeny of Ammonoidea 17 ontogenetic lobe reduction restricted to this hetero- undivided Li b and with weak Suturallobenbildung morph group *ELU 2 U 1 /—>ELU 2/ or EL 121 (Fig. in U. They demonstrate once again the restricted 16,f) were correctly placed as descendants of the phylogenetic importance of this feature: parkinsoniids. A special development within the ELU s (U 3 =- S)U,/ (Fig. 16,c). Following their Stephanocerataceae are the Tulitidae without U„, suture formulas, the Tulitidae, Pachyceratidae

/

FIG. 8. Suture ontogeny of stephanoceratids with U,, and subdivided Otoites sp. cf. O. ttuntilostts WESTERMANN, (middle ), England (from Schindewolf, 1965). [a, primary suture; g, adult suture at whorl height of 2.5 mm.; i, at height of 5.8 mm.] 18 The University of Kansas Paleontological Contributions—Paper 47

and "Oecoptychiidae" may be placed likewise required is the fact that the Cretaceous Ammo- within the Perisphinctaceae. New investigations nitina as a whole can now be traced back to their about this point are needed. For the majority of Jurassic forerunners and not linked to the phyl- the Perisphinctaceae the lobe formula loceratids or lytoceratids, as accepted by WRIGHT ELL I 2 U 3 (U :S) UIVUIdI is applicable (Fig. 16,g), (1955, 1957) and CASEY (1957, 1961). Also im- but in the Simoceratidae (ELU2 U 3 :U 1 ,U 1 ,,,/) the portant for this conclusion was the observation Suturallobenbildung is reduced, and in the Ende- that no transition exists between the lituid / of moceratidae, Aspidoceratidae, Peltoceratidae, the phylloceratids and the denticulate / of the and also the splitting of U 1 : Cretaceous Ammonitina ( WIEDMANN, 1962b, ELU 2 U 3 1 1 1 (Fig. 16,h). In their suture for- 1966a). mula ELU 2 (1/3 =S)U,/ the Holcodiscidae show A subject of special interest which illustrates greater affinity with the Perisphinctaceae than the overriding importance of lobe ontogeny is with the Desmocerataceae (WanmANN, 19660. that of the so-called Cretaceous heteromorphs and Within these desmocerataceans, forms with their derivatives. Contrary to their present wide- and without division of U, appear simulta- spread interpretation as a polyphyletic collection neously. Those with an undivided U and with of phylogenetic end forms, recently all these het- the lobe formula ELU 2 U3 U 5 U 7 :U 6 /../U,/ (Fig. eromorphs have been shown to be characterized 16,i) are here the older (Eodesmoceratinae), by a quadrilobate primary suture (WIEDMANN, which led to the suggestion of the derivation of 1963, 1965, 1966b, 1969a). This makes a mono- the Desmocerataceae from the haploceratids phyletic origin probable, at least for the Ancylo- (WIEDMANN, 1966a). The Pulchelliidae with their cerataceae (incl. Turrilitaceae): *ELUI-->ELUI suture development *ELU2 U,/—>ELU 2 U3 U5 U7 : (Fig. 9,1 ). Since the change from a quinquelo- U8 L/6 UU,/ appear to follow directly, and sim- bate to a quadrilobate primary suture took place ilarly, all of the Acanthocerataceae for which the suddenly, transitional forms are hardly to be ex- analogous formula ELU 2 U3 U1 :U6 L/U,I (Fig. pected. From their general evolute shell habit 16,/) is characteristic. The formation of adventi- and scarcity of suture elements the root forms are tious lobes (A) within the Sphenodiscidae is a most likely to be present among the lytoceratids, slight deviation from this scheme. although origin in the Jurassic Ammonitina, as All younger desmoceratids (Desmoceratinae, derivatives of Lytoceratina, can no longer be ruled Puzosiinae), as well as the Silesitidae, Kossmati- out today. ceratidae, Pachydiscidae, and above all most of The Scaphitaceae also exhibit a quadrilobate the Hoplitaceae, differ from the above in the primary suture (WiEomANN, 1965). From phylog- Suturallobenbildung in U and a splitting of U eny as well as lobe ontogeny of this group it has The extraordinarily constant lobe formula common become clear (op.cit., fig. 14, 15) that the re- to all these forms is ELU2 U,(U=S)U,X, d/ generated lobes in the saddle LU observed by (Fig. 16,k), which demonstrates the necessity to SCHINDEWOLF (1961, 1968) are in reality homol- unite the "Desmocerataceae" and Hoplitaceae ogous with the saddle frills of the root form (WIEDMANN, 19666). While WIEDMANN, on the Eoscaphites. Therefore they were called pseudo- basis of continuous transition between eodesmo- lobes by WIEDMANN (1965) and labelled p. ceratids and desmoceratids, supported the view Thus the lobe development of the Scaphitaceae that all Hoplitaceae were rooted in the haplo- fits without difficulty into the general pattern of the ceratids and that therefore a new transition from Cretaceous heteromorphs: *ELUI-4ELp,p,p 3 : an undivided to a divided U, occurred (Fig. 7), U U d i (Fig. 9,2). This view is supported by the SCHINDEWOLF (1966) again held such a regenera- open initial whorl of Eoscaphites. tion or reversion to be impossible. He derived More surprising was the observation that a all younger desmoceratids and hoplitids from quadrilobate primary suture is also present in perisphinctid forms with a "still original" U,. some Hoplitaceae, the so-called "false hoplitids" The Acanthocerataceae, however, represent, in (i.e., Douvilleiceratidae, Cheloniceratidae, Para- SCHINDEWOLF ' S opinion, phylogenetic end forms hoplitidae, Acanthohoplitidae, Astiericeratidae, of the desmoceratids or direct descendants of the Trochleiceratidae, Mathoceratidae, Deshayesiti- haploceratids (SCHINDEWOLF, 1967a). dae). Thus in the Trochleiceratidae, which were More significant than these discrepancies, for classified with the Pulchellidae by WRIGHT (1957), the solution of which further investigations are a lobe development according to the "heteromorph

Kullmann & W iedmann—Sutures in Phylogeny of Ammonoidea 19

Scaphitaceae

primary suture line

FIG. 9. Different types of suture ontogeny in Ancyloceratina. 1. Standard evolution of Cretaceous heteromorphs (Ancylocerataceae).-2. Suture development by lengthening of saddle LU, insertion of "psetniolobes" (p) and sub- division of U, (Scaphitaceae).-3. Suture development by subdivision of the lobes L and U (Douvilleicerataceae). —4. Suture development by subdivision of saddle Ut?!=lobe regeneration (Deshayesitaceae).

formula" *ELUI—>ELUI was found (WIEDMANN, WIEDMANN ' S (1966b, p. 45) assumption of identity 1963); in the Douvilleiceratidae the suture of these two genera could be proved. Since all development is expressed by the formulas these "false hoplitids" show an extraordinary *ELUI—>EL,LdUv :UdI (Fig. 9,3), and in the convergence to the genuine hoplitids in their in- Parahoplitidae by the analogous formulas volution, sculpture, and even in apparent com- *ELUI—>ELU„:U dI (Fig. 10) or ELU„Urd :Ud/ plexity of the adult suture, their evolutionary role (WIF.DMANN, 1966b). Final proof that these probably would not have been recognized without Douvilleicerataceae also are derivatives of the het- consideration of lobe ontogeny. Figure 9 gives eromorphs, probably the leptoceratids, was dem- an impression of the identity, as well as diversity, onstration of an open first whorl in the oldest of the well-defined heteromorph suture within douvilleiceratid genus Paraspiticeras by WIED- the four superfamilies now included here. MANN (1966b, pl. 4, fig. 2a). At this point the inadequacy of the Soviet lobe The Deshayesitidae, which belong to the same terminology becomes evident. Because of its com- form group of "false hoplitids," even show a plexity and inability to bring out homologies, genuine lobe regeneration according to the for- relationships of the Douvilleicerataceae remained mulas *ELU 2 ?/—>ELU 2 ?U,?: U 1 ?/ (Fig. 9,4). obscure even though thorough investigations of Thus a separate origin in late heter- lobe patterns had been made by MIKHAILOVA °morphs (Hemihoplites) can be assumed for this (1957, 1958, 1960, 1963). A short comparison of form group proposed as distinct superfamily lobe formulas selected from recent Soviet (MI- Deshayesitaceae (WIEDMANN, 1966b). Indepen- KHAILOVA, 1960; RUZHENTSEV, 1960, 1962; SHEVY- dently, TOVBINA (1965) considered the hetero- REV, 1960) and German papers (SclinsIDEwoLF, ceratid genus Colchidites to be a possible ancestor. 1961-68; WIEDMANN, 1963, 1965, 1966a,b) should This apparent discrepancy could be resolved, if make this clear (Table 2). 20 The University of Kansas Paleontological Contributions—Paper 47

0

o

E LUI

FIG. 10. Suture ontogeny of a "false hoplitid": Gargasiceras gargasense (n'ORBIGNY), Lower Cretaceous (), France (from Wiedmann, 19666). [a, primary suture; h, adult suture at whorl height of 4.5 mm.]

Kullmann & Wiedmann—Sutures in Phylogeny of Ammonoidea 21

TABLE 2. Comparison of Suture Formulas

WEDEKIND'S terminology RUZHENTSEV ' S terminology

Phylloceras ELU2U, (Uj=S)Uiii (V ,1/ i)UUVLI 4 UVII 911"U"U":11"UVW ID Lytoceras ELL I f(U 1,)1. (V 1)ULP I 1:1 Schlotheirnia ELI 1,11,L 4 1.15:U ,d1 ( V ,VI)UUVOL1 4 :121,D Ham matoceras ELLIE( 1,U 4 :( U,U,1)I (VA/ OLICIVLP ELL EU ,U 4 U ,U 7:U sU ,,.0 ,d1 (V Ai OUIPUVOU 6 :U sldsD Cadoc eras ELU,LI,U,sU (V IV 1)UUVU :U ;sir D ELU2U,U 4 :1I 5U ,U,.1 (V IV OUIPLIV D Polyplectites ELLI.1.13:U,U „I ( 1 /IV ,)ULP 1,s1,PD Kosmoc.eras ELUtU,U 4 U (V IV i)UU'llIT :1 1 1XD Strenoceras ELU2U,U 4 11,,U,d1 ( V Y :U'ID Perisphinctes ELL 4.=S )U,,,U (V ,V )UU'Ll; :LI,' ID "Pseudo perisphinctes" ELL 1,U,(11 4 =-S) U,.U,,d ( V ,VOULIVW ....0 , 31,1eD Grossouvria ELU,U,(LI 4 =S)U,,,U (V,V OWL 1 /U, s tI EL11,11.4U 4 =S)U (V ,1/ 1)U UT , 2 t1 ,q1 !ID ChelonicerasELrLaUr:UaI (V IV i)UU2012:1,D Colon; biseras ELU :Ujl ( V ,V1)1JU I :ID Acanthohoplites ELU,U dUdI (V ,V DOM' :ID Hypacanthoplites ELUrUrd:U ai (V A/ i)UIPU,':U1 21D Des hayesites ELWIL?:11,?1 ( V ,V i)DIP:PPD

As indicated by Table 2 the Soviet terminology acanthoplites shows perfect agreement with that for the closely related genera "Pseudoperisphinc- of Perisphinctes. tes" and Grossouvria shows neither mutual agree- This short review should show that only a ment nor formulas comparable with Perisphinctes. completely morphogenetically based lobe termi- "Pseudoperisphinctes," however, can be compared nology, such as that proposed by WEDEKIND, can with Cadoceras, and Grossouvria with Craspedites. differentiate homologous and convergent elements. Craspedites itself shows more agreement with Only from a scheme of this kind can new phylo- than with any of the closely related genetic knowledge be expected. We must, of genera Macrocephalites, Strenoceras, or Kosmo- course, realize that any terminology is no more ceras. Within the Stephanocerataceae the Soviet than an aid to facilitate communication concern- terminology shows extraordinary convergence, ing an often extremely complicated set of facts. while on the other hand much greater agreement The system which is simplest and above all most appears to exist with certain "false hoplitids" accurate will always be preferable. The ambition (Macrocephalites-Colombiceras, Polyplectites-Che- to produce new and more complicated terminol- loniceras, or Deshayesites). The lytoceratid suture ogies should not be allowed to disturb what after shows most affinity with the polyplectid suture, all must be the goal of every terminology: inter- and that of the phylloceratids with the oppeliids. national understanding. It would be very regret- For example, particular attention should be di- table if the continuing controversy over lobe ter- rected to the difference in Soviet terminology be- minologies should damage the reputation of lobe- tween the sutures of the "false hoplitids" Acan- ontogeny studies without bringing nearer realiza- thohoplites and Hypacanthoplites which in reality tion of mutual wishes for a natural system of are identical. Instead, the lobe formula of Hyp- ammonoid classification.

SURVEY OF SUTURAL DEVELOPMENT IN SUPERFAMILIES The presently understood sutural development in ammonoid superfamilies is outlined in the following tabulation. Sutural formula of primary suture is marked by an asterisk (*).

I. Order Goniatitida Suborder Anarcestina Families 1. Anarcestaceae a. EL, ELI,—>ELUI,-->EALU2U (Fig. 11,a) Mimoceratidae, Anarcestidae, a.o.

b. ELI,-3(E1 E2 E„i E2 E 1 )LI (Fig. 11,b) Auguritidae

22 The University of Kansas Paleontological Contributions—Paper 47

2. Pharcicerataceae a. *ELI--->(ElEmEl )L1,-->(El E„,E 1 )LUI (Fig. 11,c) -->(EiEniEJLUR Us . . . Gephuroceratidae Pharciceratidae b. *ELI--->E,E„,E,LU „I [Eœ = up to E7, Uy=llp to U18] (Fig. 11 4) Beloceratidae 3. EL/,--->ELU(/ i /i ) (Fig. 11,e) Prolobitaceae Suborder Goniatitina 4. *ELI—>EALI,-->EALUI,—>EA 3A 2 A 1 LUI (Fig. 11,1) Cheilocerataceae 5. Goniatitaceae a. *ELI-4(El E„,E1 )ALUI (Fig. 11,g) Goniatitidae, Gastrioceratidae, Paragastrioceratidae b. Same as a, L-3(t„.L,,,L4),-3L„L.L4 (Fig. 11,h) Metalegoceratidae

c. Same as a, U2 —>(U 2vU2mU2d) 2vU2mU2d (Fig. 11,i) Schistoceratidae 6. Agathicerataceae a. *ELI--->(E l EmEi )ALUI—>trifurcation of E l and A (Fig. 11,k) ...-Delepinoceratidae b. Same as a, trifurcation of (E1 E„,E1 )A„A niA 4L:UI (Fig. 12,a) Agathiceratidae 7. Shumarditaceae a. *ELI(E i E,„Ei )ALUI—>simple trifurcation in A, L and I, repeated trifurcation in A and U (Fig. 2) Shumarditidae, Marathonitidae b. Same as a, repeated trifurcation of A, L, and U Cyclolobidae c. Same as a, trifurcation of A and U, furthermore insertion of other A and U (Fig. 12,b) Popanoceratidae 8. Dimorphocerataceae a. *ELI--->(E / E,,,E0ALU/-->denticulation of individual lobes Thalassoceratidae b. Same as a, however more or less irregular bifurcation of E and A (Fig. 12,c) Dimorphoceratidae c. Same as a,--->(E1E2E„,E2 E0AL:U1 (Fig. 12,d) Neodimorphoceratidae 9. *ELI—>(E,E,,,E 1 )ALUI--->(E i EmE1 )AL(U2 =S)U1 1, -->further insertion of U, the last U=S (Fig. 12,e) Adrianitaceae Suborder Prolecanitina 10. Prolecanitaceae a. *EL/-4ELU 2 U / /,—>increase of U up to 7 (Fig. 12,1) Prolecanitidae b. Same as a,-->(E 1 EmEi )LU 2 U 1 /, increase of U up to U9, denticulation of individual lobes Daraelitidae c. Same as a,-->(E / EmEi )(LrL4)U2 U / /, increase of U up to U10, bifurcation of individual lobes Pronoritidae 11. Same as 10e, but L,, becomes part of the external saddle, which becomes incised (Fig. 12,g) Medlicottiaceae 12. *ELI(?)—>E„,E,LU,L U0=U2-111 Sagecerataceae II. Order Clymeniida 13. ELI,-->ELUI,—)EA 2 A I LU 1 U 21 (Fig. 12,i) or---->(E / En.„Ei )LU 2 U i / Gonioclymeniaceae 14. *ELI—>(El EOL(I I I,),—>L Parawocklumeriaceae 15. *ELI--->LI,—>ALI, or LUI, or ALUI (Fig. 12,k) Clymeniaceae III. Order Phylloceratida 16. Otocerataceae a. *ELI?—>ELU 2 :U i l i (Fig. 13,a) Xenodiscidae b. *ELU / /--->ELU2 L/3 :U / / / (Fig. 13,b) Ophiceratidae c. *ELU 1 ELU2 U3 U5 :U 4 U1 /1 (Fig. 13,c) Otoceratidae Kullmann & Wiedmann—Sutures in Phylogeny of Ammonoidea 23

17. Phyllocerataceae a *ELU / /—>ELU2 Us :U / / / (or ELU2U/v:Ulaii and therefore may be included in Meekocerataceae) Ussuritidae, (Fig. 13,d)—>ELU 2 /./ 3 UJ U 6 :UU,/ / (?) "Palaeophyllitidae" b. *ELL7 / /? and *ELU 2 L/ i /--->ELU2 U3 U / / /--> ELU2 (U 3 =S)U 1 / 1-->ELU 2 (U 3 _–_–_S)U 1 I (I frilled) Discophyllitidae c. *ELU 2 U 1/--->ELU 2 (U3 =S)U 1 i 1—> ELU2 U3 (U 4 =S)U 1 .1 1 (Fig. 5, 15,a) Phylloceratidae 18. Meekocerataceae *ELU / /---)ELU 2 :U / / i--->ELU 2 U s :U 111 19. Clydonitaceae a. *ELU 1 l—>ELU 2 :U,1—>ELU2 I. 3 :U,I (Fig. 13,f) Clydonitidae, Clionitidae, Arpaditidae, Tibetitidae, Thisbitidae, Trachyceratidae b. *ELU,1--->ELU2 U / :/ (I single-pointed) Lecanitidae, Nannitidae c. *ELUI—>ELU:I (I single-pointed) (Fig. 13,g) Choristoceratidae 20. Tropitaceae *ELU / /—>ELU 2 :U i / 21. Ceratitaceae a. *ELU,1--->ELU 2 /./ 3 :U,1 /—>ELU 2 U 3 U 1 U 7 :U 6 UU / /i (Fig. 13,h) Ceratitidae b. *ELU 1 /—>ELU 2 UJJ 6 U 8 U 10 U H U 9 U 7 U7U 3 U // 1 (Fig. 14,a) Hungaritidae

c. *ELU / /—>ELU 2 U:Us U 1 i 1 Aplococeratidae d. *ELU I /--->ELU2(Uiv:Vi4/i Prionitidae 22. Dinaritaceae *ELU / (--->EL(U /y :U /d )/ / (Fig. 13,i) 23. Lobitaceae *ELU,/--->E(LyLd)(U2vU2d)U3:U/rU/d// (Fig. 1 4,b) 24. Arcestaceae ?a. *ELU I /---->ELU 2 (U3 =S)Ud 1 (Fig. 13,k) Procarnitidae h. *ELUI—>ELU 2 (t.1 3 =S)U,/, (Fig. 14,c) Megaphyllitidae c. *ELU 2 U,EALU 2 U3 U,U,,,U„d/ (Fig. 14,d) -Arcestidae 25. Ptychitaceae *ELU,I-->ELU 2 (U 3 =S)U,1 (Fig. 14,e) 26. Pinacocerataceae Suture unknown IV. Order Lytoceratida Suborder Lytoceratina 27. Lytocerataceae a. *ELU,/(?)—>ELU 2 (U R,:U / d)/ (Fig. 14,1) Trachyphyllitidae b. *ELU 2 U / /—>ELU 2 (U iv :U / d)/8 (Fig. 6, 14,g) —> ELU2U3:(U/vUid)/8 Lytoceratidae c. *ELU 2 1./ i /—>ELU2 U / /8 Ectocentritidae 28. Tetragonitaceae a. *ELU2 (/ 1 /—>ELU2 U 1 /8 (U 1 suspensive) (Fig. 14,h) Gaudryceratidae b. *ELU 2 U / / and ELU 2 U 3 L/ 1 /---> ELU 2 Us (U-S)U 1 I, (Fig. 14,i) Tetragonitidae Suborder Arnmonitina 29. Psilocerataceae a. *ELU 2 U / /—>ELU2 U 3 :U/vUld (up to 4-=S) (Fig. 15,b) Psiloceratidae,

24 The University of Kansas Paleontological Contributions—Paper 47

Schlotheimiidae, Arietitidae (p.p.), Polymorphitidae, Amaltheidae, Dactylioceratidae b. *ELU 2 1/ 1/—>ELU 2 U3 U /2 :U id,Uidd/—> ELU2 U 3 :(/ / U1d/ (Fig. 15,c) Eoderoceratidae c. ELL/2 (U 1v =S)U 1d/ Cymbitidae, Liparoceratidae d. *ELU2 L/ //--->ELU 2 U3 :U i /—>ELU 2 :U i/ (Fig. 15,d) Arietitidae (p.p.), Echioceratidae, , Phricodoceratidae, Aegoceratidae 30. Hildocerataceae *ELU 2 L/ i /ELU 2 U3 :L1 1 / ELU2 U3 U 5 U,U 3 U n t/ 22 :U /0 U 8 U 6 UU,/ (Fig. 15,e) 31. Hammatocerataceae a. *ELU2L/ / /---->ELU 2 U3 U:(U /2 U / a)/ (Fig. 15,f) Hammatoceratinae b. *ELU2 U 1 /—>ELU 2 U3 U 1 / Phymatoceratinae c. ? ELU2 U 1 :1 (Fig. 15,h) Tmetoceratidae d. *ELU2 L/ / /-->ELU 2 U3 1J:L/,/ -Dumortieriidae e. *ELU2 U/ /—>ELU2 U 3 U 4 U 3 U 2 U 8 U 6 UU, d1 (to U 13 ) (Fig. 15,g) f. *ELU2 U 1 /--->ELU 2 U.7 :U 1 , Um! Phlycticeratinae g. *ELUI--->ELU 2 U 5 :U 4 U3 U j U 1 I Paroniceratidae 32. Haplocerataceae a. *ELU2 U / /-->ELU 2 U3 U 3 :U 6 UU / / Graphoceratidae, Strigoceratidae b. *ELU2 U / /—>ELU2 U 3 U 5 UU / /—> ELU2U1U5U7U 3 // 11 U 10 U 8 U 6 U 4 L/ 1/ (Fig. 15,i) Sonniniidae c. *ELU2L///--->ELU 2 U3 U 5 U 2 U 8 :U 6 U 4 U 1 / (Fig. 15,k) d. *ELU2L11/—>ELU 2 U 3 U 5 :U 6 U 4 1/ 1 / Mazapilitinae e. *ELU2U//--->ELU 2 U3 1/ 3 U 7 U2 :U s U G UE/ 7 / Aconeceratinae 33. Stephanocerataceae a. *ELU21/2/—>ELU2U3:(U /2 U 1 d)Un/ (Fig. 8, 16,a) —>ELU2 U3 U 5 :UU 1 t/./ Otoitidae b. *ELU2U,/--->ELU2U3U 5 :UU,U„/ (Fig. 16,b) Stephanoceratidae, Sphaeroceratidae c. *ELU2 U 1 /—>ELU2 U 3 U 4 :U 5 U 1 Un/ Macrocephalitidae d. *ELL/2U1/—>ELU 2 L/ 3 :U 1 Lin/ Kosmoceratidae e. *ELU2 U i-->ELU2 Us U4 U1 U ni Cardioceratidae f. *ELU2U 1 1--->ELU 2 U 3 U 5 :1/ 4 U 1 U„/ (Fig. 16,e) Morphoceratidae, Parkinsoniidae g. *ELU2 U 1 /--->EU2 / (Fig. 16,f) or ELU2/ Spiroceratidae, Acuariceratidae h. *ELU 2 U,/-->ELU2 (1/3 =S)U Ï / (Fig. 16,c) Tulitidae j. *ELU2 L/ / /—>ELU2 :U 1 Ui / (Fig. 16,d) Pachyceratidae k. *ELU2 L/1/—>ELU 2 U3 U:U 1 1 "Oecoptychiidae" 34. Perisphinctaceae a. *ELU2U1/—>ELU2U 3 (5)1-1 1 U 1 4/ (Fig. 16,g) Perisphinctidae, Berriasellidae, Reineckeiidae,

Kullmann & W iedmann—Sutures in Phylogeny of Ammonoidea 25

Craspeditidae, Olcostephanidae b. *ELU 2 L/ 1 /-->ELU 2 U3 U 4 U 1 / (to U7) (Fig. 16,h) Oosterellidae, Endemoceratidae, Aspidoceratidae c. *ELU 2 U 1 1—>ELU 2U 3:1J 11 Peltoceratidae d. *ELU 2 U,1—*ELU 2 UU iv U / d/ Simoceratidae e. *ELU 2 U / 1—>ELU 2 (U 3 =S)U 1 /—> ELU 2 11 3 (U=S)U,,,U id/—>ELL/ 2 (U 2 =S)U1/ Holcodiscidae 35. Hoplitaceae a. *ELU 2 U 1 /--->ELU 2 U 3 U 5 U 7 :U 6 U 4 U,1 (Fig. 16,i) Eodesmoceratidae b. *ELU 2 U / 1—>ELU 2 U 3 U 5 U 7 :U 8 U 6 UU 1 1 Pulchelliidae c. *ELU 2 /./ / /—>ELU 2 Us (Uc=S)U,,U,d/ (Fig. 16,k) , Kossmaticeratidae, Pachydiscidae, Hoplitidae, Schloenbachiidae, Placenticeratidae (with trifurcation in L) d. *ELU 2 11 1 /—>ELU 4 U 3 UU 1t,U 1 d1 Silesitidae, Leymeriellidae 36. Acanthocerataceae a. *ELU 2 1/ 1 /—>ELU 2 U 3 U 5 :U 6 UU3/ (Fig. 16,1) Brancoceratidae, Lyelliceratidae b. without U6 Acanthoceratidae, Vascoceratidae c. *ELU 2 U 1 /--->ELU 2 U 2 U 1 / Collignoniceratidae d. *ELU 2 U,1—>EALU 2 U 3 U 5 U 7 U 9 U 1 1:U10U8U6UU1/ Sphenodiscidae Suborder Ancyloceratina 37. Ancylocerataceae *ELUI—>ELUI (Fig. 9,1) 38. Scaphitaceae *ELUI—>ELUI—>ELp i p,p3 :Uv Udl (Fig. 9,2) 39. Douvilleicerataceae a. *ELUI—>ELUI—>EL,,Ld U,,:UdI (Fig. 9,3) Douvilleiceratidae b. *ELUI—>EL,L d Ul Astiericeratidae c. *ELUI—>ELL4:U dI (Fig. 10) or ELUUrd:Ud/ Parahoplitidae d. *ELUI--->ELUI Trochleiceratidae 40. Deshayesitaceae *ELU 2 ?/—>ELU 2 ?U 3 ?:U,?/ (Fig. 9,4) 16 The University of Kansas Paleontological Contributions—Paper 47

REFERENCES

ARKLLL, W. J., 1950, A classification of the Jurassic ammonites: Jour. , v. 24, p. 354-364, fig. 1-2. KUMMEL, BERNHARD, & WRIGHT, C. W., 1957, Mesozoic Ammonoidea: in Treatise on Invertebrate Paleontology (R. C. Moore, cd.), Part L: Ammonoidea, p.L80-L490, fig. 124-558, Geol. Soc. America & Univ. Kansas Press (New York). BEZNOSOV, N. V., 1960, K sistematike yurskikh Ammoni- tida [To the systematics of the Jurassic Ammonitidae]: Paleont. Zhurnal, 1960, no. I, p. 29-44, fig. 1-13. BOGOSLOVSKIY, B. I., 1961, Eifelskie ammonoidei Urala i vo prosy klassifikatsii agoniatitov [Eifelian ammonoids from the Urals and the problem of agoniatite classifica- tion]: Paleont. Zhurnal, 1961, no. 4, p. 60-70, fig. 1-7, pl. 7. LIBROVICH, L. S., & RUZHENTSEV, V. E., 1962, Nadotryad Ammonoidea, Ammonoidei. Sistematiches- kaya chast [Subclass Ammonoidea, Ammonoids. Sys- tematical part]: in Yu. A. Orlov (ed.), Osnovy Paleontologii, Mollyuski-Golovonogie, 1, p. 335-425, fig. 93-187 (Moscow). BRANCO, W., 1879, Beitrdge zur Entwickelungsgeschichte der fossilen Cephalopoden. Theil I. Die Ammoniten: Palaeontographica, v. 26, p. 15-50, pl. 4-13. CASEY, RAYMOND, 1957, The Cretaceous ammonite genus Leymeriella, with a systematic account of its British occurrences: Palaeontology, v. 1, p. 29-59, fig. 1-6, pl. 7-10.

FIG. 11. Adult sutures of Paleozoic Anarcestina and Goniatitina.

a. Maenioceras terebratum (SANDBERGER), Middle Devon- ian (Givetian), Germany (from Bogoslovskiy, et al., 1962, after Holzapfel, 1895). b. Augurites mirandus BOCOSLOVSKIY, Middle Devonian (Eifelian), USSR (from d Bogoslovskiy, 1961). c. Manticoceras sinuosum (HALL), Upper Devonian (Eras- Man), USA (from Miller & Furnish, 1957, after Clarke, 1899). d. Mesobeloceras thomasi GLENISTER, Upper Devonian (Frasnian), Australia (from Bogoslovskiy, et al., 1962, after Glenister, 1958). e. Clymenoceras insolitum SCHINDEWOLF, Upper Devonian (Famennian), Germany (from Bogoslovskiy, et al., 1962, after Schindewolf, 1938). f. Discoclymenia cucullata (vox BucH), Upper Devonian (Famennian), Germany (from Schindewolf, 1951). g. Goniatites choctawensis SHUMARD, Mississippian, USA (from Miller & Furnish, 1957). h. Eothinites kargalensis kargalensis RUZHENTSEV, Lower Permian (Artinskian), USSR (from Ruzhentsev, 1956). z. Pseudoschistoceras simile TEICHERT, Lower Permian, a Australia (from Bogoslovskiy, et al., 1962, after Teichert, 1944). k. Delepinoceras bressoni cantabricus KULLMANN, Upper Carboniferous (lower Namurian), Spain (from Kull- rnann, 1962). Kullmann & Wiedmann—Sutures in Phylogeny of Ammonoidea 27

, 1961, The Ammonoidea of the Lower Greensand. l'art Ill: Palaeontograph. Soc., v. 115, P. 119-216, fig. 39-68, pl. 26-35. CHAO, KING-KOO, 1959, Lower Triassic ammonoids from western Kwangsi, : Palaeont. Sinica, new ser. B,

v. 9, ii -Fp. 155-355, 45 pl. COBBAN, W. A., 1952, Scaphitoid of the Colo- rado Group: U. S. Geol. Survey, Prof. Paper 239, 42 p., 4 fig., 21 pl. HYATT, ALPHEUS, 1867, The cephalopods of the Museum of Comparative Zoology: Harvard Univ., Museum Comp. Zoology, Bull., v. 1, p. 71-102. , 6c SMITH, J. P., 1905, The Triassic genera of America: U. S. Geol. Survey, Prof. Paper 40, 214 p., 85 pl. KARPINSKIY, A., 1896, 0 nakhozhdenii Azii Prolecanitcs i o raz vitii etogo roda [On the record of Prolecanites in Asia and the development of that genus]: Imper. Akad. Nauk, Izvestiya, v. 4 (no. 2), p. 179-194, St. Petersbourg. !Russian, with French résumé: Sur l'ex- istence du genre Prolecanites en Asie et sur son développement.1 KOSSWIG, C., 1959, Phylogenetische Trends, genetisch betrachtet: Zool. Anzeiger, v. 162 (7/8), p. 208-221. KULLMANN, jÜRGEN, 1962, Die Goniatiten der Namur- Stufe (Oberkarbon) im Kantabrischen Gebirge, Nord- spanien: Akad. Wiss. Lit. Mainz, Abhandl., Math.-Nat. KI., 1962, no. 6, 119 p., 17 fig., 7 pl. , 1963, Die Goniatiten des Unterkarbons im Kanta- brischen Gebirge (Nordspanien). Il. Pationtologie der U. O. Prolecanitina MILLER & FURNISH. Die Altersstel- lung der Faunen: Neues Jahrb. Geologic Palaontologie, Abhandl., v. 116 (no. 3), p. 269-324, 11 fig., pl. 17-20.

FIG. 12. Adult sutures of Paleozoic Goniatitina, Prolecan- itina and Clymeniida.

a. Agathiceras uralicum KARPINSKIY, Lower Permian (Artinskian), USSR (from Miller & Furnish, 1957). b. Popanoceras soboleivskyantem (DE VERNEUIL), Lower Permian (Artinskian), USSR (from Ruzhentsev, 1956). c. Asturoceras subdivisum (Kum..mANN), Upper Carbon- iferous (lower Namurian), Spain (from Kullmann, 1962). d. Neodimorphoceras texanum (Smrar); Upper Pennsyl- vanian, USA (from Bogoslovskiy, et al., 1962, after Miller 6c Downs, 1950). U1,2 e. Critnites sribkrotorvi RUZHENTSEV, Lower Permi.in fo, El) A ti,4 tin Atch, Z 02 14 Ufre(-4,(2;-4/f) (Artinskian), USSR (from Ruzhentsev, 1956). 6'4/2f,612 III I f. Prolecanites (Cantabricanites) postapplanatus KuLL- MANN, Lower Carboniferous (upper Visean), Spain (from Kullmann, 1963). g. Prouddenites primas MILLER, Upper Pennsylvanian, USA (from Miller & Furnish, 1957). Ii. Pseudosageceras tnultilobatum NOETLING, Lower Trias- a sic, USSR (from Bogoslovskiy, et al., 1962, after Kiparisova, 1947). i. Sphenoclymenia maxima (MUNSTER), Upper Devonian (Famennian), Germany (from Schindewolf, 1957). k. Cymaclymenia striata (MONSTER), Upper Devonian (Famennian), Germany (from Schindewolf, 1957).

28 The University of Kansas Paleontological Contributions—Paper 47

LUPPOV, N. P., & DRUSHCHITS, V. V., eds., 1958, Am- monoidei (Tseratity i Ammonity), Vnutrennerakovin- Lit It 112 nye. Prilozhenie: Konikonkhii [Ammonoids (ceratites and ammonites), coleoids. Appendix: coniconchs]: in Yu. A. Orlov (ed.), Osnovy Paleontologii, Mollyuski- Golovonogie, 2, 190 p., 78 pl. (Moscow). MIKHAILOVA, I. A., 1957, 0 sistematike semeistv Parahop- litidae SPATH i Deshayesitidae SToyANow [On the systematics of the families Parahoplitidae SPATH and Deshayesitidae STOYANOW] : Moskov. Univ., Vestnik, Ser. Biol., Pochuoved, Geol., Geogr., 1957 (no. 3), p. 173-182, 9 fig. (Moscow). , 1958, Nekotorie dannye o rode Acanthohoplites SINZOW i Hypacanthoplites SPATH [Some dates on the genera Acanthohoplites SINZOW and Hypacanthoplites SPATH] : Same, 1958 (no. 1), p. 101-108, 4 fig. h , 1960, Ontogenez i sistematicheskoe polozhenie roda Colombiceras SPATH [Ontogeny and systematic position of the genus Colombiceras SPATH] : Moskov. Obshch. Ispyt. Prirody, Byull., new ser. 65, Otdel. Geol. _r 35 (no. 2), p. 116-120, 4 fig. , 1963, 0 sistematicheskom polozhenii i obcheme roda Diadochoceras [On the systematic position and the scope of the genus Diadochoceras]: Paleont. Zhurnal, 1963 (no. 3), p. 65-77, 10 fig., pl. 7. MILLER, A. K., FURNISH, W. M., & SCHINDEWOLF, O. H., 1957, Paleozoic Ammonoidea: in Treatise on Inverte- brate Paleontology (R. C. Moore, ed.), Part L, 4, p. LI-L79, fig. 1-123, Geol. Soc. America & Univ. Kansas Press (New York). NoETLING, FRITZ, 1905, Über die Ontogenie von Incloceras baluchistanense NOETLING: Neues Jahrb. Mineralogie, Geologic, Pal5ontologie, 1905 (no. 1), p. 1-14, fig. 1, pl. 1. , 1906, Die Enttvicklung von Indoceras baluchis- tanense NoETLING. Em n Beitrag zur Ontogenie der Ammoniten: Geol. Palaeont., Abhandl., new ser., v. 8, no. 12, p. 1-96, 22 fig., 7 pl.

FIG. 13. Phylloceratid adult sutures of the Upper Permian and Triassic. a. sp., Upper Permian, Salt Range. b Ophiceras sakuntala DIENER, Lower Triassic (lower Scythian), Himalayas. c. Otoc eras sp. cf. O. woodwardi GRIESBACH, Lower Trias- sic (lower Scythian), Himalayas. d. Leiophyllites taramellii (MARTEEE1), Middle Triassic (Anisian), Bosnia. e. Monophyllites sphaerophyllus (HAuER), Middle Triassic (Anisian), northern Alps. f. Trachyceras sp. cf. busiris (MONSTER), Upper Triassic (lower Carnian), Tyrol. g. Choristoceras marshi (HAuER), Upper Triasstic (Rhaet- ian), northern Alps. h. Ceratites nodosus (BRuGuiRRE), Middle Triassic, Ger- many. i. Dinarites asiaticus SHEVYREV, Lower Triassic (upper Scythian), USSR. k. Procarnites kokeni (ARTHABER), Lower Triassic (upper Scythian), USSR. (a-f,h from Schindewolf, 1968; g from Wiedmann, 19696; from Shevyrev, 1968.) Kullmann & Wiedmann—Sutures in Phylogeny of Ammonoidea 29

POPOV, Yu. N., 1965, Terminologiya elementov septalnoi snuffy ammonoidei [Terminology of the sutural ele- ments of ammonoids]: Paleont. Obshch., Ezhegodnik, v. 17, P. 106-114, 1 fig. REMANE, A., 1952, Die Grundlagen des Natiirlichen Sys- tems, der Vergleichenden Anatomie und der Phylogene- tik.—Theoretische Morphologie und Systematik I: 400 p., 82 fig., Geest & Portig (Leipzig). RUZHENTSEV, V. E., 1949, Osnovnye tipy evolyutsionnykh izmenenii lopastnoi linii verkhnepaleozoiskikh am- monirov [Fundamental types of evolutionary changes in suture lines of Late Paleozoic ammonoids]: Akad. Nauk SSSR, Paleont. Inst., Trudy, v. 20, Pamyati BoRisvAx, p. 183-198, 12 fig. , 1956, Nizhnepermskie Arnmonity yuzhnogo Urala. Ammonity artinskogo yarusa [Lower Permian am- monites of the southern Urals. 11. Ammonites of the Artinskian Stage]: Same, v. 60, 275 p., 97 fig., 39 pl. , 1957, Filogeneticheskaya sistema paleozoiskikh amonoidei [Phylogenetic system of the Paleozoic ammonoids]: Moskov. Obshch. Ispyt. Prirody, Byull., Otdcl. Geol., 32 (no. 2), p. 49-64 (Moscow). , 1960, Ammonoid classification problems: Jour. Paleontology, v. 34, p. 609-619, 4 fig. , 1962, Nadotryad Ammonoidea. Ammonoidei. Obshchaya chast [Subclass Ammonoidea. Ammonoids. General part]: in Yu. A. Orlov (ed.), Osnovy Paleon- tologii, Mollyuski-Golovonogie, 1, p. 243-334, fig. 1-92, pl. 1-3 (Moscow). , 1964, K roprosti o terminologii lopastnoi linii amn2onoidei [To the terminology of the ammonoid suture line]: Palcont. Zhurnal, 1964, no. 3, p. 3-14, 2 fig. SALFELD, HANS, 1920, Über die Ausgestaltung der Loben- hei Jura- und Kreide-Ammonoideen: Gesellsch. Wiss. Göttingen, Nachricht., Math.-phys. KI., 1919, Heft 3, p. 449-467, 7 fig., 3 pl.

Fin. 14. Adult sutures of Triassic Phylloceratida and Triassic-Cretaceous Lytoceratina. a. Longobardites caucasicus (SHEvvitEv), Middle Triassic (upper Anisian), USSR. h. Paralohites nautilinus (MONSTER), Upper Triassic (lower Carnian), Tyrol. c. Megaphyllites prometheus SHEVYREV, Middle Triassic (upper Anisian), USSR. d. Arcestes bicarinatus (MONSTER), Upper Triassic (lower Carman), Tyrol. e. Sturia sansovinii (MojsisovIcs), Middle Triassic (Anisian), USSR. f. Trachyphyllites costatus ARTFIABER, Upper Triassic (Norian), Timor. g. Derolytoceras toi-tom (QuElvs-runT), Lower Jurassic (upper ), Germany. h. Gaudryceras tenuiliratum YABE, Upper Cretaceous (Senonian), Hokkaido. WIEDMANN, Lower Cretaceous a 1. Tetragonites subbeticus (upper Aptian), Balearics. (a,c,e from Shevyrev, 1968; b,d,h from Schindewolf, 1968; 1,g from Wiedmann, 19696; i from Wiedmann, I962c.)

30 The University of Kansas Paleontological Contributions-Paper 47

, 1924, Die Bedeutung der Konservativstiimme liir die Stammesentivicblung der Ammoniten: 16 p., 16 pl., Max Weg (Leipzig). SCHINDEWOLF, O. H., 1923, Über die Ausgestaltung der Lobenlinie be: den Neoammonoidea W (Ad.: Centralbl. Mineralogie, Geologic, Palaontologie, 1923, p. 337-350, 359-370, 5 fig. 1929, Vergleichende Studien zur Phylogenie, Morphogenie und Terminologie der Ammoneenloben- linie: Preuss. Geol. Landesanst., Abhandl., new ser., v. 115, 102 p., 40 fig., 1 pl. , 1933, Vergleichende Morphologie und Phylogenie der Anlangsbammern tetrabranchiater Cephalopoden: Same, new ser., v. 148, 119 p., 34 fig., 4 pl., Berlin. , 1951, Zur Morphogenie und Terminologie der Ammoneen-Lobenlinie: Palaont. Zeitschr., v. 25, p. 11-34, 19 fig., pl. 1. , 1957, Clymeniina: in A. K. Miller, W. M. Furnish, & O. H. Schindewolf, Treatise on Invertebrate Paleon- tology (R. C. Moore, ed.), Part L, Ammonoidea, p. L37-L47, Geol. Soc. America & Univ. Kansas Press (New York). , 1961-68, Studien zur Stammesgeschichte der Am- moniten, Parts 1-7: Akad. Wiss. Lit. Mainz, Abhandl., Math.-nat. KI., Part 1 (1961), 1960, no. 10, p. 1-109, fig. 1-58, pl. 1, 2; Part 2 (1962), 1962, no. 8, p. 111- 257, fig. 59-149, pl. 3; Part 3 (1964), 1963, no. 6, p. 259-406, fig. 150-243; Part 4 (1965), 1965, no. 2, p. 407-508, fig. 244-301; Part 5 (1966), 1966, no. 3, p. 509-640, fig. 302-396; Part 6 (1967a), 1966, no. 8, p. 641-730, fig. 397-439; Part 7 (1968), 1968, no. 3, p. 731-901, fig. 440-478. , 196713, Die Paliiontologie im Rahmen der Natur- ivissenschalten: in S. Lauffer (ed.), Festgabe f. Dr. WALTER WILL, p. 183-199, 3 fig., C. Heymann (Kiiln).

FIG. 15. Adult sutures of Jurassic-Cretaceous Phyllo- ceratina and Ammonitina. a. onoense (STANTON), Lower Cretaceous (Aptian), USA. h. Psiloceras psilonottim (QuENsTED-r), Lower Cretaceous (lower Hettangian), Germany. c. Eoderoceras armatum (SowtRay), Lower Jurassic (upper ), England. d. Phricodoceras taylort (J. DE C. SOWERBY), Lower Jurassic (lower Pliensbachian), Germany. e. Pseudolioceras compactile (Simpson1), Lower Jurassic (upper Toarcian), Germany. 1. Hammatoceras insigne (ZIETEN), Lower Jurassic (upper Toarcian), France. g. Paroecotraustes tenuistriattis (GROSSOUVRE), Middle Jurassic (upper Bathonian), Germany. h. Tmetoceras scissum (BerxEcxE), Middle Jurassic (lower Bajocian), . i. Dorsetensia sp., Middle Jurassic (middle Bajocian), Germany. k. Haploceras grasianum (D'ORBIGNy), Lower Cretaceous (lower Valanginian), France. a (a from Wieclmann, 196213, b from Wieclmann, 196913; c,d from Schindewolf, 1962; e-i from Schindewolf, 1964; k from Wiedmann, 1966a.) Kullmann & W iedmann—Sutures in Phylogeny of Ammonoidea 31

, 1969, Homologic' und Taxonomie. Morphologische Grundlegung und phylogenetische Auslegung: Acta Biotheoretica, v. 18 (1968), p. 235-283, fig. 1-14.

1/2 Us t Up U, I SCHMIDT, HERMANN, 1921, fiber Goniatiten, eine Revision ihrer Systematik mit Beiffigung neuer Beobachtungen: Centralbl. Mineralogie, Geologic, Paliiontologic, v. 17, 1 p. 538-544, l fig. , 1952, Prolobites und die Lobenentwicblung der Goniatiten: Palaont. Zeitschr., v. 26, p. 205-217, 4 fig., pl. 13. SHEVYREV, A. A., 1960, Ontogenetichesboe razvitie neko- torykh verbhneyurskikh ammonitov [Ontogenetic de- velopment of some Late Jurassic ammonites]: Moskov. Obshch. Ispyt. Prirody, Byull., Otdel. Geol., v. 35 (no. 1), p. 69-78, 6 fig. , 1961, Ontogenetichesboe razvitie nebotorybh anizisbibh tseratitov Kavbaza [Ontogenetic develop- £17 ue ment of some Anisian ceratites from the Caucasus]: \I Paleont. Zhurnal, 1961, no. 4, p. 71-85, 11 fig. , 1962, Razvitie lopastnoi linii i terminologiya ego elementov u mezozoisbibh ammonoidei [Development of the suture line and terminology of its elements in Mesozoic ammonoids]: Same, 1962, no. 2, P. 21-33, 7 fig. , 1968, Triasovye ammonoidei yuga SSSR [Triassic ammonoids of southern USSR]: Akad. Nauk SSSR, UrvUni Palcont. Inst., Trudy, v. 119, 272 p., 92 fig., 21 Pl. U2 U3 Uy-rS SMITH, J. P., 1914, The Middle Triassic marine inverte- brate faunas of North America: U. S. Geol. Survey, Prof. Paper 83, 148 p., 99 pl. , 1927, Upper Triassic marine invertebrate faunas of North America: Same, Prof. Paper 141, iv+262 p., 121 pl. (Li Liz , 1932, Lower Triassic ammonoids of North Amer- ica: Same, Prof. Paper 167, 111 p., 1 fig., 81 pl.

FIG. 16. Adult sutures of Jurassic-Cretaceous Ammonitina.

a. 01011CS Sp. Cf. O. 1111721110SW WESTERMANN, Middle Jurassic (middle Bajocian), England. b. Chondroceras tenue WESTERMANN, Middle Jurassic (middle I3ajocian), Germany. c. Bullatimorphiles sp. cf. B. microstoma uhligi (PoPovici- HATZEG), Middle Jurassic (upper liathonian), Germany. d. Pachyceras lalandeanum (D'ORBicNy), Upper Jurassic (lower ), France. e. Parkinsonia sp. cf. P. parkinsoni (SowERBY), Middle Jurassic (upper Bajocian), Germany. f. Spiroceras bifurcati (QuENsTEDT), Middle Jurassic (upper Bajocian), Germany. g. Grossottvria stdcifera (OppEL), Upper Jurassic (Cal- lovian), Germany. h. stevenini (1•11cm.i,․), Lower Cretaceous (upper Valanginian), Spain. Eodesmoceras celestini (Ptcxxx & CAMPICHE), Lower Cretaceous (Valanginian), Switzerland. k. Neosilesites bal earensis (FALLor), Lower Cretaccous (upper Aptian), Balearics. I. lyelli (LEYMERIE in D'ORBIGNY), Lower Cretaceous (middle ), France. (a-e from Schindewolf, 1965; f from Schindewolf, 1961; g from Schindewolf, 1966; h,i from Wiedmann, 1966a; k,/ from Wiedmann, 196611.) 32 The University of Kansas Paleontological Contributions-Paper 47

SPATH, L. F., 1923-43, A monograph of the Ammonoidea Scaphites: Palaeontology, v. 8, p. 397-453, 16 fig., pl. of the Gault. Part 1-16: Palaeontograph. Soc., v. 75-97, 53-60. xiv+787 p., 248 fig., 72 pl. , 1966, Stammesgeschichte und System der post- , 1938, A catalogue of the ammonites of the Liassic triadischen Ammonoideen. Ein Überblick. 2 Teile.: family Liparoceratidae in the British Museum (Natural Neues Jahrb. Geologic u. Paldontologie, Abhandl., v. History): ix+191 p., 17 fig., 26 pl., British Museum 125 (Festb. SCHINDEWOLF), p. 49-79, fig. 1-13, pl. 1,2 (Nat. History) (London). (1966a); Same, v. 127, p. 13-81, fig. 14-47, pl. 3-6 TOVBINA, S. Z., 1965, Ob ontogeneze ammonitov roda (1966b). Colchidites [On the ontogenesis of the ammonite genus 1968, Evoluci6n y clasificaci6n de los ammonites Colchidites]; Paleont. Zhurnal, 1965, no. 3, p. 40-48, del Cretdcico: Santander, Univ. Indust., Bol. Geologia, 5 fig. v. 24, p. 23-49, 15 fig., 2 pl. WEDEKIND, R., 1913, Die Goniatitenkalke des unteren , 1969a, The heteromorphs and ammonoid extinc- Oberdevon von Martenberg bei Adorf: Gesellsch. tion: Biol. Reviews, v. 44, p. 563-602, 23 fig., 3 pl. Naturforsch. Freunde Berlin, Sitzungsber., 1913, p. 1969b, Über den Ursprung der Neoammonoideen: 23-77, 14 fig., pl. 4-7. Eclogae Geol. Helvetiae (in press). , 1916, Über Lobus, Suturallobus und Inzision: WIEDMANN, JOST, & DIENI, 1., 1968, Centralbl. Mineralogie, Geologic, Palontologie, 1916 Die Kreide Sardiniens und ihre Cephalopoden: Palaeont. Italica, v. 64, p. (8), p. 185-195, 6 fig. 1-171, 101 fig., pl. 1-18. WIEDMANN, JOST, 1962a, Ammoniten aus der Vascogoti- schen Kreide (Nordspanien). I. Phylloceratina, Lyto- WRIGHT, C. W., 1955, Notes on Cretaceous ammonites. 11. ceratina: Palaeontographica, v. 118A, p. 119-237, The phylogeny of the Desmocerataceae and the Hop- 58 fig., pl. 8-14. litaceae: Ann. Mag. Nat. History, ser. 12, v. 8, p. , 19626, Die systematische Stellung von Hypophyl- 561-575. loceras SALFELD: Neues Jahrb. Geologic u. Paliionto- , 1957, in W. J. ARKELL, B. KUMMEL, & C. W. logic, Abhandl., v. 115, p. 243-262, 5 fig., pl. 16. WRIGHT. , Mesozoic Ammonoidea, in Treatise on Inverte- , 1962e, Unterkreide-Ammoniten von Mallorca. 1. brate Paleontology (R. C. Moore, ed.), Part L, Am- Lfrg.: Lytoceratina, Aptychi: Akad. Wiss. Lit. Mainz, monoidea, p. L80-L490, fig. 124-558, Geol. Soc. Amer- Math.-nat. KI., Abhandl., 1962, no. 1, 148 p., 36 ica & Univ. Kansas Press (New York). fig, 10 pL ZAKHAROV, Yu. D., 1967, Novye vidy aniziiskikh am- 1963, Entwicklungsprinzipien der Kreideammoni- monoidei yuzhnogo Prim orja [New Anisian ammonoid ten: Paliiont. Zeitschr., v. 37, p. 103-121, 6 fig., pl. 1. species from southern Primorye]: Paleont. Zhurnal, , 1965, Origin, limits, and systematic position of 1967, no. 3, p. 39-47, 4 fig., pl. 6.