The genus Haplopleuroceras (Erycitidae, Ammonitina) in the Betic Cordillera, Southern Spain

THE GENUS HAPLOPLEUROCERAS (ERYCITIDAE, AMMONITINA) IN THE BETIC CORDILLERA,

SOUTHERN SPAIN

ASUNCION L I N A R E S & JOSE S A N D O V A L

LINARES A. & SANDOVAL J. 1996. The genus Haplopleuroceras (Erycitidae, Ammonitina) in the Betic Cordillera, Southern Spain. [Le genre Haplopleuroceras (Erycitidae, Ammonitina) dans la Cordill6re B6tique, Espagne m6ridiona- le]. GEOBIOS, 29, 3 : 287-305. Villeurbanne le 31.06.1996.

Manuscrit d6pos6 le 03.08.1994 ; accept6 d6finitivement le 02.05.1995.

ABSTRACT - The analysis of some thousand specimens of the genus Haplopleuroceras, many of which have been recorded in sections sampled bed-by-bed, allows us to carry out a monographic study of this typical Midde Jurassic Mediterranean taxon. We describe and figure the classical species H. subspinatum (BUCKMAN), H. mundum BUCKMAN, H. exirnium Gt~RARD, and H. inaequalicostatum GI~RARD, which occur in the Subbetic Domain. A new species, H. mouterdei, is here nominated and described, and another form, Haplopleuroceras sp. i is described with open nomenclature. H. tobleri RENZ and H. crassum GI~RARD are considered as synonymous ofH. subspinatum. We debate the possible phylogenetic relationships of Haplopleuroceras with other Erycitidae. Malladaites pertinax (VACEK) or related species (e.g. Malladaites striaturn LINARES & SANDOVAL) giving rise to Haplopleuroceras by means of a paedomorphic process (early innovation + neoteny). On the other hand, Zurcheria is a possible peramorphic of Haplopleuroceras (early innovation + acceleration).

KEYWORDS : AMMONITINA, SUBBETIC DOMAIN, AALENIAN, BAJOCIAN, PHYLOGENETIC RELATIONS, HETEROCHRONY.

RESUM]~ - L'objectif principal de cet article est de discuter los possibles relations phylog6n6tiques du genre Haplopleuroceras en rapport avec d'autres Erycitidae : Malladaites pertinax (VACEK) et d'autres esp6ces proches. Malladaites striaturn LINARES & SANDOVAL a donn6 naissance ~ Haplopleuroceras par un processus paedomorphique (innovation pr6coce plus n6ot6nie) ; d'autre part, Zurcheria est un possible p6ramorphique de Haplopleuroceras (innovation pr6coce plus acc616ration). Plus de mille ammonites appartenant au genre Haplopleuroceras ont 6t6 analys6es. Les esp6ces classiques ont 6t6 d6crites et repr6sent6es. En outre, H. mouterdei sp. nov. et Haplopleuroceras sp. 1, (pour le moment laiss6e en nomenclature ouverte) sont pr6sent6es.

MOTS-CL]~S : AMMONITINA, DOMAINE SUBBt~TIQUE, AAL]~NIEN, BAJOCIEN, RELATIONS PHYLOGt~N]~TIQUES, H]~Tt~ROCHRONIE.

INTRODUCTION Ammoni te f auna of the genus Haplopleuroceras is ve ry abundan t in late Aalenian - ear ly Bajocian mater ia ls from various s trat igraphical sections of the Subbetic Domain (Betic Cordillera), especially in the Median Subbetic subdomain (Garcia-Duefias et al. 1967 ; Busnardo 1979 ; Linares 1979 ; Linares & Sandoval 1990, 1993 ; Linares et al. 1988 ; etc.). (Fig. 1). Indeed, in some outcrops of the Montillana- Montej icar area [El Despefiadero, Sierra del Trigo, Barranco de Agua Larga (Fig. 2), etc.] Haplopleu- roceras m a y account for up to fifty percent of the ent i re ammoni te f auna in the levels corresponding to the Aalenian-Bajocian boundary.

Extens ive field sampling in different Subbetic localities carr ied out over the last few years has allowed

us to collect about a t h o u s a n d exempla r s of Haplopleuroceras, m a n y of which come from stratigraphical sections which we sampled level by level. The grea ter pa r t of the mater ia l consists of relative ly wel l -p rese rved i n t e r n a l moulds, a l though m an y are par t ia l ly crushed or f la t tened by la teral pressures. The presence of this ab u n d an t and well- preserved mater ia l has therefore allowed us to car ry out a detai led s tudy of this in te res t ing Medi te r ranean genus wi th a gua ran t ee of accuracy.

This s tudy of the genus Haplopleuroceras will enable us to clarify its taxonomical position, an in teres t ing aspect, since this genus has previously been inc luded in d i f ferent t a x a (Amal the idae , Sonniniidae, Hammatocera t idae , Tmetocera t idae , Erycitidae, etc.). We will also discuss biostrat igraphical questions, since Haplopleuroceras is a signi-

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° J FIGURE 1 - Geological sketch map of the Betic Cordillera showing the different paleogeographic domains and the geographic location of some areas where the most representative Upper Aaalenian-Lower Bajocian sections occur. Key : 1, Non folded cover (Mesozoic- Tertiary) of the Iberian Massif; 2, Prebetic ; 3, Intermediate Domain ; 4, External Subbetic ; 5, Median Subbetic ; 6, Internal Subbetic; 7, Penibetic ; 8, Miocene syntectonic deposits ; 9, Upper Miocene to Quaternary postorogenic deposits ; 10, Internal Zones ; 11, Flysch Units of the Campo de Gibraltar Complex ; A, La R~bita-Sierra de San Pedro, B, Montillana-Campillo de Arenas-Montejicar ; C, Cerro M~ndez ; D, Arroyo Vilanos ; E, Sierra de Ponce ; F, Sierra de Ricote ; G, Sierra del Ahillo ; H, Otifiar-Rio Frio ; I, Sierra Gorda ; J, Rfo Fardes olistoliths. Carte gdologique schdmatique des Cordill~res Bdtiques avec les diffdrentes domaines paldoggographiques et la locali- sation gdographique des rggions oh sont les coupes les plus reprdsentatives de l'Aaldnien supdrieur et du Bajocien infdrieur. 1, Couverture non plissge du Massif Ib~rique (Mdsozo~que et Tertiaire) 2, Prgb~tique ; 3, Domaine Interm~diaire ; 4, Subbdtique Externe ; 5, Subbdtique Moyen ; 6, Subbdtique Interne ; 7, Penibdtique ; 8, Miocene syntectonique ; 9, ddpSts postorogdniques du Miocene supdrieur et du Quaternaire ; 10, Zones Internes ; 11, Unitds du Flysch du Campo de Gibraltar ; A, rggion de La Rdbita- Sierra de San Pedro ; B, rggions de Campillo de Arenas-Montillana-Montej[car ; C, rggion de Cerro Mgndez ; D, Arroyo Vilanos ; E, Sierra de Ponce ; F, Sierra de Ricote ; G, Sierra del Ahillo ; H, rdgion de Ot~ar-Rfo Frfo ; I, Sierra Gorda ; J, olistolithes da Rfo Fardes.

ficant biostratigraphic indicator (Fig. 2) for the uppermost Aalenian-early Bajocian of the Mediterranean Region. The study of the phylogenetic connections between Haplopleuroceras and other genera of the family Erycitidae constitutes another interesting aspect of the present paper, since as we have already shown elsewhere (Linares & Sandoval 1986) this genus forms part of an important, typically Mediterranean lineage ranging from the Lower Aalenian dimorphous pair Erycites fallifax ARKELL Spinammatoceras pugnax (VACEK) to the uppermost Aalenian-Lower Bajocian genus Zurcheria DOUVILLE.

Order AMMONOIDEA Hyatt, 1889 Suborder AMMONITINA Hyatt, 1889

Superfamily HILDOCERATACEAE Hyatt , 1867 Family ERYCITIDAE Spath, 1928

Subfamily ERYCITINAE Spath, 1928

Genus Haplopleuroceras BUCKMAN, 1892 Species t y p e - By original designation Amaltheus subspinatus S. BUCKMAN, 1892

D e s c r i p t i o n - Medium-sized platicone shells, evolute with broad and relatively shallow umbilicus; the whorl section varies from almost square to slightly compressed subrectangular ; a keel, usual

289

FIGURE 2 - Lithological succession and vertical range of the Haplo- pleuroceras species and other interesting ammonites in the Barranco de Agua Larga (section 1 -JAQI-), Montejfcar area, Median Subbetic, province of Granada. According to Linares & Sandoval (1990) and Henriques et al. (1994) the Aalenian-Bajocian boundary is marked by the first appereance of Hyperlioceras, Toxolioceras, Oeda- nia and Reynesella. Succession lithologique avec la distribution biostratigraphique des espaces de Haplopleuroceras et d'autres ammonites dans la coupe du Barranco de Agua Larga (coupe JAQ1), Subbgtique moyen, rdgion de Monteficar, province de Gra- nada. La Iimite Aalgnien-Bajocien est signalde en accord avec Linares & Sandoval (1990) et Henriques et al. (1994) par la premiere appari- tion de Hyperlioceras, Toxolioceras, Oedania et Reynesella.

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ly bordered by more or less highly developed su]ci, runs along the broad venter. The ribs, which are single or divided near the umbilical edge, are fine, sharp, prominent and slightly sinuous; on the ventro-lateral shoulder they become markedly proverse and in the body chamber they can even cross the keel, forming a "chevron" pattern. The ribs support, sometimes, one or two rows of small fine tubercles: the lower ones near the umbilical edge and the upper ones, which are normally more highly developed, are situated in the ventro-lateral area, just by the rib inflexions. Alternate series of thick bituberculate ribs and thinner and non-tuberculate ones

may appear in the inner and middle whorls Cinae- qualicostatum" stage). The simple peristome is parallel to the ribbing. Septal suture is also simple with few branching elements and a very slightly retracted U2 and U3, the latter being very reduced in size.

R e m a r k s a n d h i s t o r y - According to the concept originally proposed by Buckman (Buckman 1881, 1883), Haplopleuroceras is connected with Amaltheus and Pleuroceras, and he included his subspinatum species in these genera. Some years later Buckman (Buckman 1892) defined the genus

290

Haplopleuroceras, which he then connected with some "Zurcheria" species. La te r authors (Renz 1925; G6rard 1937 ; Arkell 1957, in Treat ise ; Leli6vre 1960) considered the genus Haplopleuro- ceras as a descendant of "Zurcheria", and they included it in the family Sonniniidae. Seyed-Emami (1967) thought t ha t Haplopleuroceras could be connected wi th Tmetoceras or wi th "Zurcheria" gr. pertinax (VACEK). Donovan et al. (1981) include the genus in Hammatocera t idae , bu t they question its origin : on the one hand they feel t ha t Haplopleu- roceras m a y be connected wi th Spinammatoceras SCHINDEWOLF, bu t they also do not reject the possi- bility of a connection between Tmetoceras and Haplopleuroceras. Ure ta (1985) places Haplopleuro- ceras within Sonniniidae, but she does not offer any just if icat ion for this.

More recently, Linares & Sandoval (1986) include Haplopleuroceras in Hammatoce ra t i nae (in the sense proposed by Arkell 1957 in Treatise, and thus taking Eryci t inae as a synonym of Hammatocera- tinae). And finally, Sadtd (1990) and Henr iques (1992) place the genus w~thin Erycitidae. Linares & Sandoval (op. cir.) clarify the taxonomical position of "Zurcheria" pertinax : for this and related species we establ ished the genus Malladaites. There is no doubt t ha t this genus forms the connecting l ink between Spinammatoceras, a typical Erycit idae (see Spinammatoceras tenax (VACEK)) and Haplopleuro- ceras. I t is therefore clear t ha t Haplopleuroceras is a typical Eryci t idae and is not connected wi th Sonniniidae.

The generic affinities of Haplopleuroceras are limited to Malladaites LINARES & SANDOVAL and Zurcheria DOUVILLE. The inner and middle whorls of Malladaites are sometimes similar to those of Haplopleuroceras, but the outer whorls have finer ribbing, and especially in the body chamber lack keel and tubercles. This also applies to Zurcheria, where there is a different septal su ture and, according to Douvill6's original description, there is no keel t h r oughou t ontogeny (see also Fernandez

L6pez et al. 1988).

D i s t r i b u t i o n - Haplopleuroceras is found in the upper ha l f of the Concavum Zone (uppermost Aalenian) and in the lower pa r t of the Discites Zone (lowermost Bajocian). I t is a typical Medi t e r ranean genus, but is not inf requent ly found in Submedi- ter- r anean and Subboreal provinces. The genus has been recorded in Spain, Portugal , Italy, France, Britain, Greece, Nor th Africa (Algeria and Morocco), the Caucasus and Iran. I t has not been recorded to date in the Eas t Tethys, Pacific and Boreal realms.

Haplopleuroceras subspinatum (BUCKMAN, 1881) Fig. 3a-d ; P1.35, figs. 1-5.

"1881 Amaltheus subspinatus BUCKMAN, p. 606. "1883 Amaltheus subspinatus BUCKMAN : Buckman, pl. 1, fig.

1. 1889 Pleuroceras subspinatum BUCKMAN : Buckman, p. 657.

"1892 Haplopleuroceras subspinatum BUCKMAN : Buckman, p. 300 ; pl. 51, figs. 1-3, 6-10.

1912 Haplopleuroceras subspinaturn BUCKMAN : Roman & Gennevaux, p. 78, pl. 3, fig. 1

?1925 Haplopleuroceras tobleri RENZ, p. 28 ; pl. 2, fig. 5. 1937 Haplopleuroceras subspinatum BUCKMAN : G6rard, p.

635 ; pl. 30, figs. 1-5. 1937 Haplopleuroceras crassum GERARD, p. 626 ; pl. 31, figs.l-

5. 1967 Haplopleuroceras subspinatum (BUCKMAN) : Seyed-

Emami, p. 102, pl. 4, fig. 13 ; pl. 12, fig. 15. 1985 Haplopleuroceras subspinatum (BUCKMAN, 1881):

Ureta, p. 365, pl. 22, figs. 7,8 ; non pl. 23, fig. 2. 1990 Haplopleuroceras subspinatum (BUCKMAN) : Linares &

Sandoval, pl. 2, fig. 1. 1990 Haplopleuroceras subspinatum (BUCKMAN) : Cresta &

G~lacz, pl. 13, fig. 1. 1992 Haplopleuroceras subspinatum (Bucm~AN,

1881) : Henriques, p. 156, pl. 5, fig. 16.

Materiel - ~venty-five almost complete specimens and many fragments from different Median Subbetic localities.

D e s c r i p t i o n - Relatively large-sized plat icone and evolute (O/D = 0,5) Haplopleuroceras, with small spiral increase. The whorl section is subrectangular , with an almost vert ical umbilical wall, f la t tened or ve ry slightly convex flanks, a broad ven te r along

PLATE 35

Figs. 1-5 - Haplopleuroceras subspinatum (BUCKMAN). 1-2a, MOD.X.9, MOD.X.49, Upper Aalenian, Concavum Zone, Limitatum Subzone, or Lower Bajocian, Discites Zone, Cortijo del Despefiadero section, Montillana area, province of Granada. 3, JAQ1.(~53). 2, Upper Aalenian, Concavum Zone, Limitatum Subzone, Barranco de Agua Larga (section 1), Montejfcar area, province of Ja6n. 4, JAQl.(-71).1, Upper Aalenian, Concavum Zone and Subzone, Barranco de Agua Larga (section 1), Montejfcar area, province of Ja6n. 5, JAC. 11.2.26. Lower Bajocian, Discites Zone, La Torquilla section, Campillo de Arenas area, province of Ja6n.

Figs. 6-8 - Haplopleuroceras mundum BUCKMAN. 6, 8, MOD.X.3, MOD.X.84, Upper Aalenian, Concavum Zone, Limitatum Subzone, or Lower Bajocian, Discites Zone, Cortijo del Despefiadero section, Montillana area, province of Granada. 7, JAQ2.20.40, Upper Aalenian, Concavum Zone, Limitatum Subzone, Barranco de Agua Larga (section 2), Montejicar area, province of Ja6n.

(All figures natural size)

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P1. 35 A. L i n a r e s & J . S a n d o v a l

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292

which a well-developed keel runs, which is bordered by two shallow sulci which weaken toward the end of ontogeny in some specimens. The ribs are single, fine, sharp and spaced apart, and are almost radial or slightly sinuous. On the inner whorls, relatively thick and bituberculated ribs alternate with others which are thinner and non-tuberculate (the "inaequalicostatum" stage). Towards the end of the phragmocone the ribs become uniform, and they have two rows of small tubercles: the inner one is situated near the umbilical edge, while the upper coincides with the point of inflexion of the ribs. From the ventrolateral tubercles on, the ribs become markedly proverse and they weaken progressi- vely, though they tend to cross the keel. The single peristome lies parallel to the ribbing. Septal suture (Fig. 3d) is relatively simple with tripartite L, a bifid and broad L-U2 saddle and a slightly retracted U2 and U3, the latter especially being only very slightly developed.

R e m a r k s a n d af f in i t ies - All the species belonging to the genus Haplopleuroceras described so far show great similarity and it is difficult to distinguish one from the other. This leads us to assume that the different "species" of Haplopleuroceras may in fact be merely morphotypes of a single "superspe- cies". If we take morphological characteristics into account, however, it is possible to differentiate some of the species clearly, while others must be considered to be synonymous with other previously defined species.

The type of/-/, tobleri RENZ (1925, p. 28, pl. 2, fig. 5) is very similar to H. s u b s p i n a t u m both in its ornamentation (where we can distinguish an "inaequa- l icos ta tum" stage and another with single and spaced ribs) and in its coiling, section, keel, etc. The absence ofventrolateral sulci on the last whorl in H. tobleri and the presence of a slightly more persistent "inaequalicostatum" stage than in H. subspina ture are the only appreciable differences. Then again, H. c ras sum GI~RARD (1937, p. 625, pl. 30m fig. 1-5) is likewise very similar to H. s u b s p i n a t u m : a more clearly depressed whorl section and slightly less sharp ribs in H. c ras sum are the only appreciable differences. In both these cases, then, we consider that the differences are not significant enough to justify the separation of two morphotypes which are identical for other characters. H. m u n - d u m BUCKMAN is similar to H. s u b s p i n a t u m , and intermediate forms between both species occur. Nevertheless there are some distinguishing characteristics which allow us to consider them as separa- te species. H. m u n d u m is more involute, has denser and thinner ribbing and some of its ribs tend to join together near the umbilical edge. The "inaequalicostatum" stage is less persistent and more irregular, the tubercles are less well developed and the umbilical row is often missing. H. e x i m i u m GERARD is

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FIGURE 4 - Haplopleuroceras rnundum BUCKMAN : A, plots of the umbilical diameter (U) and whorl height (H) against diameter (D), B, number of ribs per half whorl (N/2) against diameter (D), C, adult external septal suture, MOD.X.7. A, diagrammes de variation du diam~tre de l'ombilic (U) et de la hauteur du tour (H) en fonction du diam~tre de la coquille (D) ; B, nombre de cStes sur un demi-tour (N/2) en fonction du diarn~tre de la coquille (D); C, ligne de suture externe et adulte.

m u c h m o r e i nvo lu t e t h a n H. s u b s p i n a t u m a n d h a s t h i n n e r r i bb ing w i t h m a n y d iv ided r ibs n e a r t h e u m b i l i c a l edge.

T h e i n n e r w h o r l s of H. i n a e q u a l i c o s t a t u m Gt~RARD

are s imi l a r to those of H. s u b s p i n a t u m , a n d show the s a m e k i n d of r ibbing , b u t t h e G ~ r a r d spec ies is s l ight ly m o r e invo lu te ; i t is a lso smal le r , i t s r ibs a re t h i n n e r and m o r e d e n s e l y d i s t r i bu ted , a n d s eve ra l r ibs a r e divided, espec ia l ly in t h e ou t e r whor l s . However , H. i n a e q u a l i c o s t a t u m m a y r e p r e s e n t t h e d i m o r p h o u s mic roconch o f H . s u b s p i n a t u r n .

D i s t r i b u t i o n - H. s u b s p i n a t u m is c o m m o n l y recorded in t h e l a t e A a l e n i a n ( C o n c a v u m Zone) a n d e a r l y Ba joc ian (Disci tes Zone) of M e d i t e r r a n e a n r eg ions : s o u t h - e a s t e r n F r a n c e ( D u b a r et al. 1971 ; M o u t e r d e et al. 1972 ; P a v i a 1983 ; etc.) ; t he A p e n n i n e s (Cres t a 1988 ; C r e s t a & Gal~cz 1990), Sici ly (Renz 1925) ; I r a n ( S e y e d - E m a m i 1967), Greece , Morocco (G~rard 1937 ; Lel i~vre 1960, Colo 1962 ; E l m i 1986; Sadk i 1990, 1992, 1994 ; Bensh i l i 1989, 1990, etc.), the Ba lea r i c I s l a n d s (Fal lot 1922 ; A l v a r o et al. 1988; S a n d o v a l 1994). H. s u b s p i n a t u m also occurs, a l t h o u g h less f requent ly , in t h e S u b b o r e a l a n d S u b m e d i t e r r a n e a n p rov inces : F r a n c e ( B r a n g e r & G o n n i n 1994), P o r t u g a l ( M o u t e r d e et al. 1972 ; Fe rn~ndez -L6pez et al. 1988 ; Rocha et al. 1990 ; H e n r i q u e s 1992), t he I b e r i a n Cord i l l e ra (Goy & U r e t a 1979 ; U r e t a 1985) ; t h e C a n t a b r i a n Cord i l l e ra (Su~rez-Vega 1974 ; F e r n ~ n d e z - L @ e z & Su~rez -Vega 1979 ; Goy et aL 1991 ; etc.), G r e a t B r i t a i n ( B u c k m a n 1881, 1883, 1892 ; C a l l o m o n & C h a n d l e r 1990). I n t h e Subbe t i c H. s u b s p i n a t u r n is found m o r e i n f r e q u e n t l y t h a n H. m u n d u r n , b u t it occurs in m o s t local i t ies w h e r e U p p e r A a l e n i a n and l o w e r m o s t Ba joc ian outcrop.

Haplop leuroceras m u n d u m BUCKMAN, 1892. Fig. 4a-d ; P1.35, figs. 6-8 ; P1.36, figs. 1-5.

"1892 Haplopleuroceras mundum BUCKMAN, p. 302, pl. 51, figs. 11 and 12.

1937 Haplopleuroceras mundum BUCK_MAN : G~rard, p. 626, pl. 30, figs. 6-9.

1967 Haplopleuroceras mundurn BUCKMAN : Garda Duefias et aL, p. 6, pl. 1, fig. 5.

1967 Haplopleuroceras cf. mundum BUCKMAN : Garda- Due~as et al., p. 6, pl. 1, fig. 3.

1976 Haplopleuroceras mundum BUCKMAN : Parsons, p. 59, pl. 2, fig. 2.

1983 Haplopleuroceras subspinatum BUCKMAN : Pavia, pl. 3, fig. 6.

1988 Haplopleuroceras mundum BUCKMAN : Linares et al., p]. 2, fig. 11.

1990 Haplopleuroceras mundum BUCKMAN : Linares & Sandoval, pl. 1, figs. 3 and 12.

1990 Haplopleuroceras mundum BUCKMAN : Sandoval, pl. 1, fig. 1.

1990 Haplopleuroceras mundum BUCKlVIAN : Rocha et al., pl. 1, fig. 5.

1990 Haplopleuroceras mundum BUCKMAN : Sadki, p. 208, pl. 1, figs. 4 and 5.

1992 Haplopleuroceras mundum BUCKMAN 1892 : Henriques, p. 160, pl. 5, fig. 12.

Material - One hundred and fifty complete or almost complete specimens, all well-preserved internal moulds, together with many fragments from various localities in the Subbetic.

294

D e s c r i p t i o n - Medium sized, platicone and evolute ( U/D = 0,45) shells wi th a rec tangular and slightly compressed whorl-section and f lat tened or very slightly convex flanks. A fairly low keel bordered by shallow sulci which may disappear in the body chamber runs down the venter. The single peristome is paral lel to the ribs. The ribbing is the most significant fea ture which distinguishes H. m u n d u m from other Haplopleuroceras species. The ribs are fine, sharp and very dense, especially on inner and middle whorls ; the "inaequalicostatum" stage is normal ly of reduced dimensions and may even be missing. Ribs have a radial or slightly sinuous ru n and at t imes they tend to join together nea r the umbilical edge. Ventrolateral tubercles are present th roughout ontogeny but periumbilical tubercles only appear on occasions at the body chamber end. The very simple septal su ture is ve ry similar to tha t o fH. subspinatum.

R e m a r k s a n d a f f i n i t i e s - Buckman (1892) based his original description of / / . m u n d u m on one single small specimen wi th fine dense ribbing. Later, G6rard (1937) considered tha t this species had no periumbilical tubercles and tha t the ventrola teral tubercles were not spiniform, unlike those in H. subspinaturn. Similarly, according to G6rard (op. cir.), the whorl-section in H. m u n d u m is more compressed and the sulci are deeper and nar rower t h an those of H. subspinatum. These last differences noted by G6rard are not found in the Subbetic mater ial , where we find t ha t H. mundurn has thinner and denser r ibbing and less well-developed tubercles t ha n H. subspinatum, as well as being slightly more involute. H. ex imium GERARD is not ve ry different from H. m u n d u m , but it is more involute, has a more compressed whorl-section, its ribbing is thinner, denser and commonly non-tuberculate , and m a n y ribs divide nea r the umbilical edge.

D i s t r i b u t i o n - H. mundurn is commonly recorded in the Concavum and Disci tes Zones of the Medi te r ranean province, where it is general ly more abundan t t h a n H. subspinatum : south-east France (Caloo 1970 ; Dubar et al. 1971 ; Mouterde et al.

1972 ; Pavia 1983 ; etc.), Majorca (Fallot 1922 ; Alvaro et al. 1988), Morocco (G6rard 1937 ; Leli6vre 1960 ; Elmi 1986 ; Benshili 1989, 1990 ; Sadki 1990, 1992, 1994), Sicily (Renz 1925), I ran (Seyed-Emami 1967), etc. I t is also found in Subboreal and Subme- d i te r ranean areas : France (Branger & Gonnin 1994), Portugal (Perrot 1957 ; Ruget -Perrot 1961 ; Mouterde et al. 1972 ; F e r n ~ d e z - L S p e z et al. 1988 ; Rocha et al. 1990 ; Henr iques 1992 ; Henr iques et al. 1994), Grea t Br i ta in (Buckman 1891 ; Parsons 1976), Spain, - Iber ian and Cantabr ian cordilleras- (Su~rez-Vega 1974 ; Goy & Ure ta 1979 ; Goy et al. 1991). It is abundan t in the Betic Cordillera, and in some levels of the Aalenian-Bajocian boundary (for example, the Monti l lana - Montejficar area) this species const i tu tes a lmost fifty pe rcen t of the ammoni te fauna found.

Haplopleuroceras ex imium G~,RARD, 1937 Fig. 5a-c ; P1. 36, figs. 6-10a ; P1. 37, figs. 1-8

"1937 Haplopleuroceras eximiam GERARD, p. 627, pl. 31, figs. 10 - 11, 12 (lectetype here designed) and 13.

1967 Haplopleuroceras eximittm GERARD : Garcia-Duefias et al., fig. 5, pl. 1, fig. 2.

1967 Haplopleuroceras cf. eximiurn GI~RARD : Garcfa-Duefias et al., p. 6, pl. 1, fig. 6.

1990 Haplopleuroceras eximium GI~RARD : Linares & Sandoval, pl. 2, fig. 4.

1990 Haplopleuroceras eximium GERARD : Sadki, p. 209, pl. 1, fig. 6.

1990 Haplopleuroceras inaequalicostatum Gt~RARD : Sadki, p. 209, pl. fig. 7.

Material - Fifty relatively well-preserved complete or almost complete internal moulds and several fragments from different Subbetic localities.

D e s c r i p t i o n - Shells of smaller size t h an those of the species described above. They are p]aticone and semievolute (U/D = 0,40). The whorl section is subrectangular and is more compressed t han in other congeneric species, wi th a subvert ical umbilical wall, almost f la t tened flanks and a ven te r wi th a keel which is f lanked by sulci, which are well- developed in the inner whorls bu t m a y d isappear towards the body-chamber The simple per is tome is parallel to the ribbing. The ribs are fine, sharp and

PLATE 36

Figs. 1-5 - Haplopleuroceras mundum BUCKMAN. 1-1a, 5, JAQ2.20.33 (figured by Sandoval 1990, pl. 1, fig. 1), JAQ2.20.43, Upper Aalenian, Concavum Zone, Limitatum Subzone, Barranco de Agua Larga (section 2), Montejfcar area, province of Ja6n. 2-2a, 4, MOD.X.237, MOD.X.1, Upper Aalenian, Concavum Zone, Limitatum Subzone, or Lower Bajocian, Discites Zone, Cortijo del Despefiadero section, Montillana area, province of Granada. 3, JAQl.(-53).l (figured by Linares & Sandoval 1990, pl. 1, fig. 12), Upper Aalenian, Concavum Zone, Limitatum Subzone, Barranco de Agua Larga (section 1), Montejfcar area, province of Ja6n.

Figs. 6-10a - Haplopleuroceras eximium GI~RARD. MOD.X.187, MOD.X.164, MOD.X.165 (microconchs?) ; MOD.X.73, MOD.X.5 (macroconchs?), Upper Aalenian, Concavum Zone, Limitatum Subzone or Lower Bajocian, Discites Zone, Cortijo del Despefiadero section, Montillana area, province of Granada.

(All figures natural size).

G e o b i o s n ° 29, fasc . 3

P1. 36 /~ L i n a r e s & J. S a n d o v a l

la

~a

8

296

dense throughout ontogeny. Ribs are radial or slightly concave, many are bifurcated near the 40 umbilical edge and have no periumbilical tubercles. Sometimes, however, small ventrolateral spines can 30 appear on the points ofinflexion of the ribs. The septal suture is very simple and similar to those of other Haplopleuroceras species. In the populations 20- ofH. e x i m i u m there are bigger forms (P1. 36, fig. 9- 10a ; P1.37, fig. 1-1a, 6-6a) which can be the macro- 10- conchs together with smaller forms which may represent the dimorphous microconchs.

0 R e m a r k s - H. e x i m i um is a species which is frequently cited but rarely figured. It is therefore difficult to determine its intraspecific variability. In the Subbetic material finely-ribbed forms which have abundant divided ribs occur together with other 60 forms where the ribbing is thicker and which have few bifurcated ribs. These latter forms can be very 50 similar to some involute specimens of H. m u n d u m . On the other hand finely-ribbed specimens ofH. exi- 40 m i u m may closely resemble Malladai tes s t r ia tum 30- LINARES & SANDOVAL (1986, p. 216, pl. 1, figs. 9-12), however these latter have thinner ribbing, the 20 tubercles are limited to the inner whorls, and the keel disappears on the last whorl, where the venter 10- is flatter and broader than in H. ex imium. The specimen figured by Sadld (1990, pl. 1, fig. 7) as H. inaequal icostatum has very fine ribs, some of them divided, which leads us to think that this specimen falls within the intraspecific variability of H. ex imium.

D i s t r i b u t i o n - H. ex im ium is commonly recorded in the upper part of the Concavum Zone (Limitatum Subzone) and more frequently from the Discites Zone of Mediterranean areas, Morocco (G~rard 1937 ; Leli~vre 1960 ; Colo 1962 ; Benshili 1989, 1990 ; Sadki 1990, 1992, 1994), South-East France (Caloo 1970 ; Dubar et al. 1971 ; Mouterde et al. 1972 ; etc.). Beyond the Mediterranean province H. ex im ium is very rare, but it has been cited from the Iberian Cordillera (Fern~ndez-L6pez 1979) and the Cape Mondego area, Portugal (Mouterde et al. 1972; Fern~ndez-L6pez et al. 1988 ; Rocha et al. 1990)

Haplopleuroceras inaequalicostatum GI~RARD, 1937 Fig. 6a-c ; P1.37, figs. 9-12 ; P1. 38, figs. 1-4

"1937 Haplopleuroceras inaequalicostatum GI~RARD, p. 627, figs. 6, 7 (lectotype, here designated), 8 and 9.

1967 Haplopleuroceras inaequalicostaturn GI~RARD : Garda-Duef ias et al., p. 5, pl. 1, fig. 4.

1974 Haplopleuroceras sp. nov. Sugrez-Vega, p. 169, pl. 14A, fig. 6.

1985 Haplopleuroceras c£ inaequalicostatum GI~RARD : Ureta , p. 369, pl. 23, fig. 1.

non 1990 Haplopleuroceras inaequalicostatum GI~RARD : Sadki, p. 209, pl.1, fig. 7 = H. eximium GI~RARD.

1992 Haplopleuroceras sp. 1, Henriques , p. 162, p]. 5, fig. 17.

M a t e r i a l - Th i r ty complete in te rna l moulds and several frag-

U & H (mm)

# o

A A

D (mm 10 20 3'0 40 50 60 70 80

I, • U/D H/D 1A

N/2

• . . = , : • i m • mmm •

_.'.: ".|

0 10 2'0 30 40 50 6'0 7'0 80 D (mm)

• D vs. N/2 B

FIGURE 5 - Haplopleuroceras eximium GI~RARD : A, plots of the umbilical d iamete r (U) and whorl he igh t (H) aga ins t d i amete r (D) ; B, n u m b e r of ribs per ha l f whorl (N/2) aga ins t d i ame te r (D). ,4, diagrammes de variation du diam~tre de l'ombilic (U) et de la hauteur du tour en fonction du diam~tre de la coquille (D) ; B, nombre de c6tes sur un demi-tour (N / 2) en fonction du diam~tre de la coquille.

ment s from diverse Median Subbet ic localities, par t icu lar ly from the Mont i l lana-Montej icar area.

Descr ip t ion - Platicone, rather small and relatively evolute (U/D = 0.43) shells. The whorl-section is subrectangular or subovate in the phragmocone and rectangular with flattened flanks in the body chamber. The keel is bordered by two shallow sulci which may disappear on the last half whorl. There is a simple peristome parallel to the ribs. The most significant feature of H. inaequal icostatum is its ornamentation. In the inner and middle whorls thick and bituberculate (periumbilical and ventrolateral tubercles) alternate with thinner and non- tuberculate ribs. A transitional stage appears later in which single ribs coexist with ribs which divide near the umbilical edge. Finally, single, slightly sinuous ribs with fme - and usually only ventrola- term - tubercles are dominant in the body chamber.

297

R e m a r k s - Within the H. inaequalicostatum populations from the Subbetic typical forms which are 40- very similar to the ]ectotype coexist with specimens which have a more reduced "inaequalicostatum" stage. These latter forms bear a slight resemblance 30 to H. mundurn, whereas the forms with fine ribbing are similar to H. eximiurn. Only H. subspinatum 20 also has a well-developed "inaequalicostatum" stage ; however, it is larger, more evohite and has thicker and more widely-spaced ribs which are not 10 divided on the outer whorls. Due to this difference in size and the similarity between the inner whorls, 0 with the "inaequalicostatum" stage almost equally developed in both species, we are of the opinion that H. inaequalicostatum may represent the microconch of/-/, subspinatum.

D i s t r i b u t i o n - This type comes from Hamraoua (Morocco). The species has been found on the Aalenian-Bajocian boundary in Mediterranean and 50 Subboreal regions • Morocco (G~rard 1937 ; Colo 1962 ; Leli~vre 1960 ; Sadki 1992, 1994), Portugal 40 (Henriques 1992), the Cantabrian and Iberian 30 Cordilleras (Su~rez-Vega 1974 ; Fernandez LSpez 1979 ; Ureta 1985 ; Goy et al. 1991). In the Betic 20 Cordillera it is found especially abundantly in the Montillana-Montej~car area, in the provinces of 10 Ja~n and Granada, but has also been recorded in the Cerro MSndez area (provinces of Ja~n and Granada), the Sierra de Ricote (province of Murcia), etc.

Haplopleuroceras mouterdei nov. sp. Fig. 7 ; P1. 38, figs. 5-8

71988 Haplopleuroceras sp. nov., Fernfindez-LSpez et al., p. 306.

Derivatio nominis - Dedicated to Prof. Dr. R. Mouterde, French paleontologist, outstanding investigator of Lias and Dogger ammonites.

t t o lo type - JAQl.(-34).l, deposited in the Museum of Depart- ment of Stratigraphy and Paleontology, Granada University.

Locus typicus - Barranco de Agua Larga section, Montejicar area, province of Ja~n.

Stratum typicum - Lower Bajocian, Discites Zone.

D i a g n o s i s - Platicone, evolute, with a subrectangular whorl-section finely ornamented with sinuous ribs, a low keel and very shallow sulci, which may also be absent altogether.

Material - Four specimens from the Barranco de Agua larga section and one from the Sierra del Trigo section.

M e a s u r e m e n t s (mm)

D U H u h

JAQ1.(-34).l 37.5 15.5 13.0 41.3 34.7 32.0 12.0 11.2 37.5 35.0

JAQl.(-34).2 41.0 16.0 14.3 39.0 34.9 JAQl.(-35).l 53.0 22.8 17.3 43.0 32.6

U & H (mm)

, | o

D (mm)

10 20 30 40 50 6'0 70 80

I • U/D A H/D ]A

N/2 60

• ~ l I i

0 10 20

[ 30 40 50 6'0 7'0 80

D (mm)

[] D vs. N/2 ] B

FIGURE 6 - Haplopleuroceras inaequalicostatum G]~RARD : A, plots of the umbilical diameter (U) and whorl height (H) against diameter (D) ; B, number of ribs per half whorl (N/2) against diameter (D). 2t, diagrammes de variation du diam~tre de l'ombilic (U) et de Ia hauteur du tour (H) en fonction du diam~tre de la coquille (D) ; B, nombre de cStes sur un demi-tour (N/2) en lone- tion du diam~tre de la coquille (D).

40.0 18.6 15.0 41.5 37.5 JAQI.(-40).2 28.5 10.5 10.0 37.5 35.7 JST33.1 37.0 16.5 12.8 44.6 34.6

D e s c r i p t i o n - The only complete specimen avai- lable has a maximum diameter of 53 ram, but some of the fragments found clearly belong to larger specimens. The shells are platicone and evolute, with a wide and relatively shallow umbilicus. The whorl- section is subrectangular and compressed with a vertical umbilical wall and parallel or very slightly convex flanks. A low keel bordered by two very shallow sulci - which may also be absent - runs along the venter, which is relatively broad. The ribs vary slightly throughout ontogeny but are usually very fine and not so sharp as in other congeneric species. They have no tubercles, rise just on the umbilical edge and run in a sinuous line, at first in a clearly proverse direction and later almost radially, before

298

becoming proverse again on the ven t ro l a t e ra l shoulder. Fine ribs - sometimes reduced to mere riblets- cross the ven te r in the body chamber forming chevrons. Sometimes ribs join together nea r the umbilical edge. The septal suture, which is only par t ia l ly preserved in one specimen (Fig. 7) is very simple and is s imilar to tha t of other species of Haplopleuroceras.

R e m a r k s a n d a f f i n i t i e s - Mouterde et al. (1972) and Fern~ndez-L6pez et al. (1988) note the presence of Haplopleuroceras with fine and sinuous ribbing and only slightly marked sulci, or indeed an absence of such snlci, in the Discites Zone of Cabo Mondego (Portugal). These forms m a y belong to this new species. H. mouterdei differs from other congeneric species in its freer and more sinuous ribbing and in tha t there are no tubercles th roughout ontogeny. Some species of Zurcheria, such as Z. striata FERNANDEZ-LOPEZ et al. (1988, p. 228, pl. 1, figs. 1- 6) from the Discites Zone of the Cabo Mondego area (Portugal) show a ribbing which is similar to tha t of H. mouterdei, but they are more involute and have no keel, at least on the last whorl.

D i s t r i b u t i o n - The only mater ia l known, tha t from the Subbetic, was found in the Discites Zone of the Barranco de Agua Larga section, in the area of Montejfcar in the province of Ja~n and from the Sierra del Trigo section nea r Frailes, also in the province of Ja6n.

Haplopleuroceras sp.1 P1. 38, figs. 8-11

Material - Two complete internal moulds and several fragments, some of which include the end of the body chamber with the aperture.

Measurements (ram)

D U H u h N/2

JAQ2.19.1 61.0 29.5 18.5 46.7 30.1 45

50.0 23.0 15.0 46.0 30.0

~ E U2

I I •

FIGURE 7 - Haplopleuroceras mouterdei nov. sp. : adult septal suture, JAQ1.(-34).1, (holotype). Ligne de suture externe et adulte (holotype).

MOD.X.149 72.0 35.0 21.0 48.6 29.2 46 59.0 28.0 18.0 47.5 30.1

D e s c r i p t i o n - Platicone and evolute re la t ively large-sized shells wi th a m a x i m u m d iamete r of near ly 70 mm. The whorl-section is subrec tangula r and slightly compressed, wi th an almost vert ical umbilical wall, sl ightly convex flanks on the inner whorls which become fiat in the body chamber. There is a well-developed keel and shallow sulci persis t th roughout ontogeny. The single aper tu re is parallel to the ribbing. The whorls have a relat ively well-developed "inaequal icostatum" stage, which is la ter gradual ly replaced by fine, densely ranged slightly concave or sinuous ribs wi th ven t ro la te ra l tubercles coinciding wi th the inflexion point of these ribs. Some ribs tend to join together n ea r the umbilical edge, and all become ve ry free in the last third of the body chamber.

R e m a r k s a n d a f f i n i t i e s - The forms described above share some features wi th H. m u n d u m and H. eximium. Compared to H. mundurn, Haplopleuro- ceras sp. i has a more highly-developed "inaequalicostatum" stage, its whorl section is more compressed and the ribbing is t h inne r and denser. H. exi- m i u m is smaller, has a more reduced "inaequalico-

PLATE 37

Figs. 1-8 - Haplopleuroceras eximium GI~RARD. 1-1a, 2, 4, 5, 6-6a, MOD.X.156 (macroconch?), MOD.X.188, MOD.X.191, MOD.X. 161 and MOD.X.38, Upper Aalenian, Concavum Zone, Limitatum Subzone or Lower Bajocian, Discites Zone, Cortijo del Despefiadero section, Montillana area, province of Granada. 3, JAQJ.(-53). 15, Upper Aalenian, Concavum Zone, Limitatum Subzone, Barranco de Agua Larga (section 1), Montejicar area, province of J~ten. 7, JST1.22.4, Upper Aalenian, Concavum Zone, Limitatum Subzone, Sierra del Trigo section, province of Ja~n. 8, JAQ2.20.34, Upper Aalenian, Concavum Zone, Limitatum Subzone, Barranco de Agua Larga (section 2), Montejicar area, province of Ja~n.

Figs. 9-12 - Haplopleuroceras inaequalicostatum Gt~RARD. 9, JAQl.(-52).l, Upper Aalenian, Concavum Zone, Limitatum Subzone, Barranco de Agua Larga (section 1), Montejfcar area, province of Ja~n. 10, 11, 12, MOD.X.170, MOD.X.16, MOD.X.212, Upper Aalenian, Concavum Zone, Limitatum Subzone or Lower Bajocian, Discites Zone, Cortijo del Despefiadero section, Montillana area, province of Granada.

(All figures na tu ra l size).

G e o b i o s PI. 37 n ° 29, fasc . 3 A. L i n a r e s & J. S a n d o v a l

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Haplopleuroceras subspinatum (BUCKMAN)

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? J

Haplopleuroceras inaequalicostatum GERARD

G Mal~adaites ~ ) pertinax (VACEK)

Malladaites (m) vaceki LINARES & SANDOVAL

Spinammatoceras (M) tenax (VACEIO

Erycites fallifax ARKELL

. 6 Splnammatoceras (m) schindewoIfi LINARES & SANDOVAL

G Splnammatoceras (m) pugnax (VACEK)

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FIGURE 8 - Simplified sketch showing the possible relationships of Haplopleuroceras with other Erycitidae. Modified from Linares & Sandoval 1986. Schdma simplifig des relations phylogdndtiques possibles de Haplopleuroceras avec d'autres Erycitidae.

301

stature" stage, thinner and denser ribs and its tubercles -where they exist- are also smaller.

D i s t r i b u t i o n - Haplopleuroceras sp. 1 is found in the Concavum Zone (Limitatum Subzone) and the Discites Zone of the Despefiadero section near Montillana in the province of Granada, and the Barranco de Agua Larga (section 2) in the Montejicar area, in the provinces of Ja4n and Granada.

H A P L O P L E U R O C E R A S A N D ITS P H Y L O G E N E T I C C O N N E C T I O N S

The studies which we have carried out on the ammonite fauna and biostrat igraphy of the Subbetic Aalenian (Linares & Sandoval 1986, 1993; Sandoval and Linares 1993) have allowed us to fol- low the evolutionary patterns of a group of forms belonging to the family Erycitidae whose presence extends from the Opalinum Zone (Comptum Subzone) to the Discites Zone. This phylogenetic sequence (fig. 8) is made up of : Erycites (M) fallifax ARKELL - Spinamrnatoceras (m)pugnax (VACEK), Sp. (M) tenax (VACEK) - Sp. (m) schindewolfi LINARES & SANDOVAL, Malladaites (M)pertinax (VACEK) - M. (m) vaceki LINARES & SANDOVAL, Malladaites (M) sp. 1 (in Linares & Sandoval 1986, p. 118, pl. 1, figs. 13-15) Haplopleuroceras spp and Zurcheria spp.

A characteristic feature of all the forms mentioned above is that their septal suture is of the erycitid type, with a tripartite L and more or less retracted auxiliary U2-U3 lobes. However, according to Geczy (1966), this characteristic may be more or less accentuated depending on whether the specimen is a macroconch or a microconch (see also Linares & Sandoval 1986, Figs. 7a, 9a & l l a in text). Another common feature for most forms of this lineage is the similarity of their juvenile stages, which are cha- racterized by the presence of a more or less well- developed "inaequalicostatum" stage. In some forms (M. pertinax, "H. tobleri" RENZ, H. subspinaturn and especially H. inaequalicostatum) this stage can be exceptionally highly developed. However, in other forms (Erycites fallifax, H. mouterdei and Zurcheria spp., with the exception of Z. (Parazur- cheria) tuberculata FERNi~NDEZ-LOPEZ et al. 1988, p. 287, pl. 1, figs. 7-8) this stage is very reduced.

Within this phylogenetic sequence the evolutive step from Spinammatoceras (ej. S. (M) tenax - S. (m) schindewolfi) to Malladaites (ej. M. (M)pertinax - M. (m) vaceki is marked by the appearance of a well-developed "inaequalicostatum" stage in the inner and middle whorls. This stage, with its une- qual ribs, may have originated in the looped ribs of Spinammatoceras, since in some specimens the cen-

tral rib hendidure is only partially realized. The gradual appearance of the keel from the middle towards the outer whorls together with the existen- ce of more or less well-developed ventral sulci give evidence of the evolutive step from Malladaites to Haplopleuroceras.

We have already focused (Linares & Sandoval 1986) that the ancestor of Haplopleuroceras must have been M. pertinax or a related species such as Malladaites sp. 1 (in Linares & Sandoval 1986). H. subspinatum, H. tobleri (seen as a possible synonym for H. subspinatum ) or H. inaequalicostatum, which have a well-developed "inaequalicostaturn" stage, in the same way than the latest Malladaites, may represent earlier primitive forms of the genus, whereas H. mouterdei is the more highly evolved Haplopleuroceras species.

However, the relationships we have established can give rise to problems : 1. H. subspinatum, H. mundum, H. inaequalicostatum and H. eximium are first recorded almost exactly at the same time in the different stratigraphical sections sampled in the Subbetic (Fig. 2), so it is impossible to establish their temporal relations and, even, they could represent a single biospecies. 2. Some Malladaites species are very similar to others of Haplopleuroce- ras (e.g. Malladaites sp. 1 compared to "H. tobleri" on the one hand and M. striatum to H. exirnium on the other). This may lead us to conclude tha t different Haplopleuroceras species have their origin in other Malladaites species (a hypothesis which is not testable) and therefore to consider that Haplopleu- roceras and Malladaites are simply morphological genera of one single phylogenetic genus.

Linares & Sandoval (1986) and, in more detail, Fern~_ndez-L6pez et al. (1988) have discussed the feasible relationships between the genera Haplopleuroceras and Zurcheria DOUVILLE. Linares & Sandoval (op. cit. p. 222, fig. 13 in text) consider Zurcheria to be a possible descendant of Malladaites or Haplopleuroceras. Similarly, we included Spinammatoceras, Malladaites and Haplopleuroceras among the Hammatoceratinae, because at the time of writing we were using Hammatoceratinae in the sense proposed by ArkeIl et al. (1957, in Treatise), and we therefore included Erycitinae as synonyms of Hammatoceratinae. Fernfindez-Ldpez et al. (1988, p. 293) emphasized the direct relationship of Zurcheria with Erycites or Spinammatoceras, pointing out tha t the septal suture of Zurcheria is significantly different from that of Malladaites and Haplopleuroceras, and they figured several sutures of Zurcheria (Fern~ndez- Ldpez et al., op. cit., fig. 3 in text). However, if we compare the septal suture of Zurcheria with that of Malladaites and Haplopleuroceras we can see that such differences are minimal (see, for example, the

302

septa l su tures of M. pertinax and M. striatum, in L inares & Sandoval 1986, figs. 7a, 9a in text ; and Z. striata, Parazurcheria tenuissima and P. costula- ta, in Fern~ndez-L6pez et al. 1988, fig. 3). I f we bear in mind such cr i ter ia as s t ra t igraphica l range, ex te rna l morphology and septa l su ture develop- ment , then, we could see t h a t Spinammatoceras, Malladaites, Haplopleuroceras and, possibly, Zurcheria form p a r t of the s ame l ineage wi th in the Erycit idae.

The evolutive s tep f rom Malladaites to Haplopleu- roceras took p lace in the U p p e r A a l e n i a n (Concavum Zone and Subzone). I f we compare onto- genies, we obse rve t h a t adu l t spec imens of Haplopleuroceras are s imi l a r in size to adul t Malladaites and s imi lar in the i r morphology to juveni le Malladaites (as to the ven t ra l keel, single and b i tubercula te ribs, etc.), all of which leads us to suppose t h a t Haplopleuroceras is a paedomorphic descendan t of Malladaites (early innovat ion + neoteny, accord ing to t he concept p roposed by D o m m e r g u e s et al. 1986). Malladaites seems likewise to be a paedomorph of Spinammatoceras.

The genus Zurcheria, which (like Haplopleuroceras) h a s a less a c c e n t u a t e d d i m o r p h i s m t h a n do MaIladaites and Spinammatoceras, has no keel (except on the i nne rmos t whorls), the o rnamen ta - t ion of i ts i nne r whor ls is s imi la r to t h a t of HaplopIeuroceras, and its septa l su ture has a slightly more r e t r ac ted U3. Zurcheria m u s t therefore be a pe ramorph ic of Haplopleuroceras (delayed innova- t ion + acceleration).

On the other h a n d the hypothes is pu t forward by Fern~ndez-L6pez et al. (1988) is also reasonable. They claim a direct phylogenetic re la t ionship between Spinammatoceras, or r a t h e r Malladaites and Zurcheria. The only problem wi th this, a t leas t according to our field da ta f rom the Subbetic, is the v i r tua l absence of Malladaites and/or Zurcheria in the u p p e r m o s t C o n c a v u m Zone ( L i m i t a t u m Subzone). In this case, then, the evolutive step to Zurcheria would be m a r k e d by the g radua l disap- pea rance of the keel and the tubercles toward the

inner whorls, following a pe r amorph i c accelerat ion process.

Acknowledgement - We are very grateful to Prof. Dr. S. Elmi (Claude-Bernard University, Lyon), Prof. Dr. D. Contini (University of Besan~on) and Dr. L. Rulleau (Claude-Bernard University, Lyon) for their critical reading and kind suggestions. Likewise we would like to thank sincerely Prof. N. McLaren (University of Granada) for supervising the English translation. We thank also Dr. Busnardo for his critical reading and observa- tions. This study was funded as part of Project PB91-0733 of the DGICYT (Ministerio de Educaci6n y Ciencia) within Group EMMI (Junta de Andalucia).

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Figs. 1-4 - Haplopleuroceras inaequalicostatum GI~RARD. MOD.X.213, MOD.X.133, MOD.X.171, MOD.X.184, Upper Aalenian, Concavum Zone, Limitatum Subzone or Lower Bajocian, Discites Zone, Cortijo del Despefiadero section, Montillana area, province of Ja6n.

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A. LINARES & J. SANDOVAL Departamento de Estratigraf~a y Paleontolog~a

Facultad de Ciencias Universidad Granada 18002-Granada, Spain