TRIASSIC L I N G U L I D E B R A C H I O P O D S F R O M THE I B E R I A N R A N G E (SPAIN) ANA MJ~QUEZ-ALIAGA, CHRISTIANC. EMIG & JUAN M. BRITO MARQUEZ-ALIAGA A., EMIG C.C. & BRITO J.M. 1999. Triassic lingulide brachiopods from the Iberian Range (Spain). [Brachiopodes ]ingulides du Trias de la Cordill6re Ib6rique (Espagne)]. GEOBIOS, 32, 6: 815-821. Villeurbanne, le 31.12.1999. Manuscrit d6pos6 le 08.10.1998; accept6 d6finitivement le 15.01.1999. ABSTRACT - During the Middle Triassic marine transgression in Spain, several lingulide populations were fossilized in the Iberian Range (western part of Sephardic Province) and the recorded specimens have been described previously under several specific names. The paleontological aspects of the Jalance (Valencia province) section have been studied for the first time. Its exceptionally large population was fossilized in situ as flat-lying valves and can be interpreted as an autochthonous association related to a very shallow marine environment. The lingulide specimens belong to the genus Lingularia BIERNAT& EMIG, 1993, but the species name remains under debate, probably Lingularia smirnovae. Internal morphology and shell characters are described and compared with other Lingularia species. KEYWORDS: MIDDLE TRIASSIC, MUSCHELKALK, BRACHIOPODA, LINGULARIA, SPAIN. RI~SUMI~ - Au cours de la transgression marine du Trias moyen en Espagne, plusieurs populations de lingulid6s ont 6t6 fossilis6es dans la Cordill6re Ib6rique (zone ouest de la Province Sephardique) et les exemplaires d6crits ant6rieurement ont 6t6 attribu6s h plusieurs esp6ces. Les aspects pal6ontologiques du gisement de Jalance (province de Valence, Espagne) sont 6tudi6s pour la premi6re fois. La population de ce site a 6t6 fossilis6e in situ en valves s6par6es et interpr6t6e comme une association autochtone vivant darts un environnement marin en eau peu profonde. Tousles lingulid6s r6colt6s appartiennent sans contestation au genre Lingularia BIERNAT& EMIG,1993, mais l'esp6ce n'a pu 6tre d6termin6e avec certitude, probablement Lingularia smirnovae. L'6tude de la morphologie interne et des caract6res de la coquille a permis de faire une comparaison avec les autres esp6ces du genre Lingularia. MOTS-CLI~S: TRIAS MOYEN, MUSCHELKALK, BRACHIOPODA, LINGULARIA, ESPAGNE.
INTRODUCTION D u r i n g the Middle Triassic the first m a r i n e t r a n s gression began in the E a r l y Anisian, onlapping the l a n d - m a s s e s of t h e I b e r i a n P e n i n s u l a , N o r t h Africa and C e n t r a l Europe. In this way, fluvial and lacustrine red beds of the E a r l y Triassic (Bunts a n d s t e i n Facies) p a s s e d up into shallow-water m a r i n e c a r b o n a t e s ( M u s c h e l k a l k Facies). Such conditions prevailed d u r i n g the Middle Triassic. The last m a r i n e t r a n s g r e s s i o n took place at the end of the L a t e Triassic. The Iberian Peninsula (Figs 1, 2) was located in the western part of the Tethys Realm. The eastern border of the Iberian Peninsula was bounded by several sub-basins (Pyrenean, Catalonian, Ebro, Iberian and Betic), opening eastwards into the Tethys and s e p a r a t e d by high, f a u l t e d Palaeozoic blocks. Upland regions were drowned by the sea during the late Ladinian and a single basin was formed. The Middle - Late Triassic (Anisian-Norian) of the Iberian Peninsula has three carbonate intervals inter-
preted as prograding carbonate ramp sequences: the lower Muschelkalk (Anisian), the upper Muschelkalk (Ladinian), and the Im6n Dolomites Formation (Rhaetian). These t h r e e intervals were separated by two siliciclastic evaporitic periods interpreted as sabkha and saline deposits: the middle Muschelkalk facies (Late Anisian - Early Ladinian) and the Keuper facies (Carnian - Early Norian?). Those successions are of G e r m a n i c Triassic type with intercalations of continental and marine facies (Fig. 3). The Triassic fossil record is v e r y scarce and poorly preserved. Studies in Spain for over a c e n t u r y (1853-1958) were of limited geographical scope (Almera 1899; W u r m 1911; Sos 1933; D a r d e r 1945; B r i n k m a n n 1948). As the paleobiological concepts were not addressed in these studies, m a n y taxonomic problems remained. In the case of lingulide brachiopods, in the Catalonian Ranges, Almera (1899) cited Lingula cf. tenuissima BRON• in the Muschelkalk of Barcelona, which is probably the first citation of a Triassic lingulide occurrence in
816 shed a good description ofLingula polariformis aff. polaris LUNDGRENand Lingula sp. from the uppermost layers of the Muschelkalk below the Keuper of Monterde and of E1 Frasno (Zaragoza province).
Germanic Province, the Sephardic Province and the Tethys realm during the Middle Triassic (modified, after M&rquezAliaga et al. 1986). Schdmas des relations paleogdographiques de la Province Germanique, de la Province Sephardique et du domaine de la Tdthys au cours du Trias Moyen (modifid d'apr~s Mdrquez-Aliaga et al. 1986).
In the first comprehensive study of Triassic fossils of Spain, Schmidt (1935) established close relationships between Germanic and Spanish faunas, and several new taxa based on Spanish specimens were described. Lingula tenuissima var. zenkeri ALBERT1 has been recorded from the Ladinian of Villora (Cuenca province), in beds located in the uppermost part of the Muschelkalk near the Keuper facies (Fig. 3). Virgili (1958), working in the Catalonian Ranges, significantly increased knowledge of the Spanish Triassic; using Schmidt's schemes, this author pointed out that in Spain the Triassic sediments are of Germanic facies type but the fauna has many Alpine affinities (Alpine sensu Tethys Realm, Fig. 1). Lingula tenuissima BRONNhas been described from the uppermost part of the Muschelkalk of Pauls and Begues (Barcelona), and the presence of various mollusc species allowed a correlation of the Lingula beds with the Cordevolian (Upper Ladinian) (Virgili 1958).
Spain. From the Aragonian area of the Iberian Range, Wurm (1911, p. 123, Tabl. 7, Fig. 5) publi-
Local stratigraphic Triassic studies of the Iberian Range were made by Dupuy & Marin (1960), Hinkelbein (1969), Viallard (1973), M~rquez-Aliaga
FIGURE 1 - Sketch of the paleogeographical relationships of the
[ • Palaeozoic ] ~
Mesozoic
U pe, Cetaceous
[----] Tertiary ~
PrebeticZone
~
I I I I I I I I
~.,
&.-. d~d~
F1GURE 2 - Geographical distribution of the Triassic lingulide beds (black circles) in Spain. Rgpartition gdographique des niveaux d lingulidds (points noirs) en Espagne.
[~k~'~!!,'t/"-D~--lt/"
e~ BOYAR
JALANC]~- . . . . . . -( i.~.~
817 (1976), Sopefia (1979) and Ramos (1979), recording, in the uppermost part of the Muschelkalk, Lingula tenuissima BRONNin Albarracin (Teruel province), Henarejos and Villora (Cuenca province), Molina de Aragdn (Guadalajara province), and L. keuperea ZEN~R in Jalance, Chelva, Macastre (Valencia province) (Fig. 2). In two studies based mainly on bivalves from outcrops of the Iberian, Catalonian and the Betic Ranges, M/~rquez-Aliaga (1985), Budurov et al. (1993) and M~rquez-Aliaga & Martinez (1996) made comparisons with Triassic sections of Italy, England, Germany, and Bulgaria. These authors concluded that 73 % of the species in the Spanish fauna, including the lingulides, occur in both the Alpine Province (Tethys realm) and Germanic Province, while only 12 % of the Spanish species are exclusively of Alpine Province affinity. The other species (15 %) can be considered as autochthonous to the Sephardic Province, defined by Hirsch (1977) as the epicontinental southern border of the Tethys realm, extending from Arabia to the Iberian Peninsula (Fig. 1). Consequently, the correlation between the "autochtonous" species
Rhaetian - Norian I "~I Carniaa ........ kadinian
~Sandstone Limestone
,LJAi~ ~.~
and
~Gypsum ~ ~lDolomite ~ M a r l y Limestone . . . . . ~.. ~ u o o m i t e wire ~ M a r l and Clay u,..z,~m cro-brecc a
The lingulide specimens studied in this present paper are from Henarejos (Cuenca province) and Jalance (Valencia province), where large populations have been sampled. The aim is to resolve taxonomic and biostratigraphical problems.
The Middle Triassic sediments of t h e Catalonian, Southeast Iberian and External Betic Ranges have similar facies (Fig. 3: schematic section). From bottom to top, the three lithostratigraphic units are as follows: - a lower carbonate unit with bioclastic limestones, algal mats and marls, of variable thickness; - a middle, red siliciclastic-evaporitic layer of variable thickness, up to several hundred meters; and, - an upper carbonate unit consisting of dolomites and green-grey marlstones with bioclastic and oolitic limestones, algal buildups and shallowing-upward marlstone-limestones sequences, up to 100 to 140 m thick; ripple-marks, cross-bedding, mud-cracks, bioturbation, algal domes, breccias, tepees and iron-rich horizons are the main internal structures. There are abundant bivalves, gastropods, crinoids, and brachiopods, and sparse cephalopods and conodonts (Hirsch et al. 1987; M~rquez-Aliaga & Martinez 1996).
so~hi~~:i:{:{:i:iiii ::::::::::::
~Conglomerate ~Marl
Over the last decade (Fig. 2), "Lingula" specimens have been recorded from several locations: Henarejos (Cuenca province) by L6pez et al. (1987) and Mfirquez-Aliaga & Ldpez (1989); Minorca (Balearic province) by Llompart et al. (1987); Monterde (Zaragoza province) by Garc[a-Royo et al. (1989); Riba de Santiuste and Fuencaliente de Medina (Guadalajara province) by Mfirquez-Aliaga & Garcia (1991); Boyar (Cadiz province) by MartinAlgarra et al. (1995); Barranco del Contador by Arche et al. (1995); Calanda (Teruel province) by M&rquez-Aliaga et al. (1994); Jalance (Valencia province) by Prats et al. (1987). Consequently, the record of "Lingula" in the Iberian Range is abundant, always located in the uppermost part of the Muschelkalk facies, probably related to very shallow marine or brackish environments (Fig. 3).
STRATIGRAPHIC SETTING
Anisian
3
(sensu Mfirquez-Aliaga 1985) occurring in Spain, Jordan and Israel gives information on migration from east to west between Anisian to Ladinian times (Mfirquez-Aliaga & Hirsch 1988; Hirsch & M&rquez-Aliaga 1988; Budurov et al. 1993).
I
u Rhizocorallium ~ Lingulariasp. I
~ (bioturbation) ~ burrows ~Limea sp. (bioturbation)
FIGURE 3 - T r i a s s i c s c h e m a t i c s e c t i o n s h o w i n g two d e t a i l e d s t r a t i g r a p h i c sections of t h e u p p e r M u s c h e l k a l k a n d of t h e u p p e r M o y a M e m b e r . Section schdmatique du Trias avec deux
colonnes stratigraphiques ddtailldes du Muschelkalk supdrieur et du Membre Moya supdrieur.
From the outcrops in the Iberian Range studied by LSpez-GSmez & Arche (1992, 1993), the Dolomites Formation (upper Muschelkalk) has been described with five members named, from bottom to top (Fig. 3: detailed section of the upper Muschelkalk): Gorgocil, Henarejos, Huelamo, Valacloche, and Moya. This formation consists of a sequence of shallow-water carbonates, organized in a complete
818 transgressive-regressive cycle representing the origin, evolution and demise of a carbonate ramp, shallowing-upwards. The maximum flooding surface is located at the top of the Henarejos Member, and the regressive part of the unit ends in a coastal sabkha with clear subaerial exposure levels (Arche et al. 1995). The "Lingula" beds located in the Moya Member are related to very shallow marine environments (Fig. 3: detailed section of the upper Moya Member). SYSTEMATIC P A L A E O N T O L O G Y It is impossible to distinguish and identify lingulid genera and species on the basis of external shape and shell size. Thus, many previous authors attributed Paleozoic and Mesozoic lingulide specimens to the Recent genus Lingula (see Biernat & Emig 1993). Consequently, several hundred specimens collected in Jalance and Henarejos have been studied to obtain information to identify the genus and species of these specimens on the basis of morphological characteristics. Details of the internal morphology and shell characters, as defined by Emig (1982, 1983), have been observed only on 6 ventral and 4 dorsal valves.
convex. Umbonal regions have not been observed. The shell surface bears only subconcentric ribbing of variable prominence and spacing (Fig. 5). A comparison of shell ratios between our specimens and other Lingularia species is given on Table 1 and Figure 6. The W/L ratios of our specimens are close to those of the broadest Lingularia species: L. similis (Biernat & Emig 1993; Table 1, Fig. 6). On the other hand, our specimens appear to have a more general oval outline.
Ventral valve (Fig. 4): a pair of narrow, V-shaped grooves extend internally from the anterior adductor muscle pair to the posterior adductor, where the grooves join. As in Lingularia smirnovae, the body muscle arrangement is characterised by: - a small internal anterior oblique, located near the anterior oblique and above the internal posterior oblique; the heart-shaped posterior adductor is developed asymmetrically, i.e. more on the right part of the ventral side. The vLC, or ventral lophophoral cavity (i.e. the distance between the distal limit of the anterior adductor muscle scars in the ventral valve, DORSAL
The material studied herein is deposited in the "Museo de Geologfa, Universitat de Valencia" (MGUV), of the Departamento de Geologfa: JA samples from Ja]ance; HE from Henarejos.
~
D e s c r i p t i o n - The specimens show all the main characteristics of the genus Lingularia (Fig. 4). The shell is elongate oval in outline, lateral margins subparallel; anterior margin is broadly rounded, sometimes quite straight. Maximum width at the mid-length of the shell. The valves are weakly TABLE 1 - Range and m e a n value of the W/L ratios of the valves, w i d t h (W), length (L) in the Lingularia species. (*) data from B i e r n a t & E m i g (1993); n.i. = n u m b e r of individuals. Intervalle et moyenne du
rapport W / L des valves, de la longueur (L) et de la largeur (W) chez les esp~ces du genre Lingularia. (*) donndes d'apr~s Biernat & Emig (1993); n.i. = hombre d'individus.
S5%
vLC
McT
4r
i (___M_a~l~c_a_ rials
@ ......... 4;,?
The new supraordinal classification of the Brachiopoda according to Williams et al. (1996) is used here. Subphylum LINGULIFORMEAWilliams et al., 1996 Class LINGULATAGorjansky & Popov, 1985 Order LINGULIDAWaagen, 1885 Family LINGULIDAEMenke, 1828 Genus Lingularia BmRNAT & EMIG, 1993 Lingularia cf. smirnovae BIERNAT& EMIG, 1993
dLC
VENTRAL
....
\
,,
FIGURE 4 - A r r a n g e m e n t of the m u s c u l a t u r e w i t h perimial line and disposition of the two m a i n anterior m a n t l e canals in Lingularia cs smirnovae. The extension of the lophophoral cavity (LC) and the distance of the tip of the mantle canals to the anterior m a r ~ n (MET) are indicated by arrows. See also text for explanation. Arrangement de la musculature avec la ligne pdri-
myale et disposition des canaux antdrieurs du manteau chez Lingularia cf smirnovae. L'extension de la cavitd lophophorale (LC) et la distance entre la partie antdrieure des canaux du manteau et le bord antdrieur de la valve (McT) sont reprdsentdes par une double fl~che. Voir aussi les explications dans le texte.
Species
n.i.
W/L range
W/L mean
L range
L mean
W range
W mean
Lingularia cs smirnovae
43
0.62-0.75
0.67
5.7-15.4
10.5
4.0-10.2
7.2
Lingularia smirnovae (*)
17
0.47-0.55
0.51
11.4-23.3
15.9
6.1-12.0
8.1
Lingularia similis (*)
66
0.44-0.75
0.62
7.9-29,2
22,1
4.5-19.0
13.7
Lingularia siberica (*)
9
0.45-0.56
0,50
4.3-8.5
7.3
2.3-4.4
3.6
Genus Lingularia
139
0.44-0.75
0.61
4.3-29.2
16.5
2.3-19.0
10.1
819
FIGUUE 5 - Dorsal valves ofLingularia cf. smirnovae. Upper Muschelkalk from Jalance (JA), Valencia province and Henarejos (HE), Cuenca province. 1. JA-3470, x 5.2. JA-3469, x 5.3. HE-3467, x 5.4. JA-3472, x 5. All the specimens from the "Ana M~rquez" collection have been deposited at the Museo de Geolog~a, Universitat de Valencia (MGUV), of the Departamento de Geologla. Valves dorsales de Lingularia cf. smirnovae. Muschelkalk supdrieur de Jalance (JA), Valencia province et de Henarejos (HE), Cuenca province.. 1. JA-3470, x 5. 2. JA-3469, x 5.3. HE-3467, x 5. 4. JA.3472, x 5.Tous les exemplaires de la collection d'Ana Mdrquez ont dtg ddposds au Museo de Geologia, Universitat de Valencia (MGUV), du Departamento de Geologia.
and that of the anterior oblique scars in the dorsal valve, to the anterior margin of the valve) is about 47 % (Fig. 4), which is near the mean percentage in the genus Lingularia which the vLC range is 37-57 % and the mean 46 %. No measurement for L. smirnovae is given by Biernat & Emig (1993). The ventral mantle canals are similar in shape to that described in L. similis by Biernat & Emig (1993): with strong, S-shaped curves on both sides; nevertheless those ofL. smirnovae were not observed by Biernat & Emig (1993). The distance between the distal tip of the mantle canals and the anterior margin of the valve represents about 24 % of the whole valve length (Fig. 4), which is one of the lowest percentages observed in the genus Lingularia as a whole: range 24-37 % and mean 32 % (see Biernat & Emig 1993).
Dorsal valve (Fig. 4): a central narrow ridge extends generally over 1-2 millimetres posteriorly from the anterior oblique muscle scars. The complete muscle arrangement has not been observed but the heart-shaped posterior adductor is developed asymmetrically (more on the left of the dorsal side). The dLC or dorsal lophophoral cavity is about 35 %, which is within the range of L. smirnovae (33-36 %) but also near the mean value (33 %) for the genus Lingularia as a whole. Mantle canals has not been observed on the dorsal side. T a p h o n o m i c a s p e c t s - The abundance of lingulide fossils in Jalance appears to be exceptional in the Iberian Range (more t h a n 500 valves per square meter). The population was fossilized in situ as separate valves (see Emig 1986, for fossilization conditions). The shells are very thin showing the
growth lines perfectly (Fig. 5), sometimes fragmented. In many cases the shell is completely or partly lacking, but specimens remain as internal moulds. The fossil population can be interpreted as an autochthonous association preserved in a calcareous dolomite sediment with algal laminations and pseudomorphic anhydrite ghosts, related to a very shallow marine water environment. In Henarejos, the specimens are very scarce; the well-preserved shells are thicker than those of Jalance. The population is recorded as separate valves in an hardground with an abundant toollusc fauna. The autochthony of the population appears doubtful (M~rquez- Aliaga 1985; LSpezGSmez et al. 1987). O b s e r v a t i o n s - According to Biernat & Emig (1993), specimens (W/L = 0.58-0.75; mean 0.63) described as Lingula tenuissima by Broglio-Loriga (1968) from the Lower Triassic of the Dolomite Mountains (Italy) have strong similarities with Lingularia. The material seems to be well-presetved, but with internal morphology described and figured only poorly. From the Triassic of Northern Spain (Monterde: Fig. 2) Wurm (1911) described specimens with W/L = 0.47 and 0.62, and vLC = 42 % under the name Lingula polariformis aff. polaris LUNDGREN;these specimens have been related to Lingularia siberica by Biernat & Emig (1993). In Villora (Fig. 2), Schmidt (1935) identified various specimens as Lingula tenuissima var. zenkeri ALBERTI,but the specimens are too poorly described and figured to allow any relationships.
820
L. similis L. s p .
;~
L. smirnovae
: := i-'l
::::.I:,
.................
I0 ' '
'I'4
I::,10
--
di iil. ::ifi::li]!l::H[ .... 24 28
L
' ' ' 1'8'
'--
...................
.
.
.
.
.
.
.
.
.
.
.
2-
%
0,50
0,50
0,70 W/L
". . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
L. siOeric,~...........;m;rn.;;;; .................... ......
sp.
i~
~176
....
L.
i~
~
.L" smirnovae ,
.
.
8
12
16
20
24
.
.
.
.
.
.
28 m m
20 ~ Vm
2
6
N " 12
" " 10
"
mm
w
[] L. similis
y = - 1,67 = 0,70x R 2 = 0,83
A L. smimovae
y= y=
9 L. sp.
y= - 0,17 + 0,69x R 2 = 0,94
Lingularia
.~
0,20 + 0,50x R2 = 0,97
9 L. siberica Genus
mE
9 9e
0,61 + 0 , 4 2 x R 2 = 0 , 8 8
Z
i~ :
[]
i
y= - 0,25 + 0 , 6 3 x R 2 = 0,92
L 0
I0
20 m m
30
FIGURE 6 - H i s t o g r a m s of the valve l e n g t h (L) and width (W), of the W/L ratios and regression cmwes m e a s u r e d in the Lingularia species; our specimens are labelled L. sp. Histogrammes de la longueur (L) et de la largeur (W) des valves, du rapport V~/L et les courbes de rdgression pour les diffdrentes esp~ces du genre Lingularia; nos exemplaires apparaissent sous L. sp.
In conclusion, the studied specimens from Jalance and Henarejos belong to the genus Lingularia, whose occurrence extends from the Carboniferous to the Cretaceous. The specific affinity of the Iberian Range remains doubtful until more data can be obtained; nevertheless, our specimens appear rather close to Lingularia smirnovae, a species described by Biernat & Emig (1993) from the Cretaceous of Kazakhstan. A c k n o w l e d g e m e n t s - This research was supported by the Direcci6n General de Investigaci6n Ciencia y Tecnologia (DGICYT) - PB 95-0084; and partly by grant-in-aid from the Centre National de la Recherche Scientifique (France) and the Universitat de Valencia (Spain) for one of us (E.C.C.). We t h a n k two "anonymous" referrees, Prof. Michael Bassett and Dr. Lars Hotmer, who made helpful comments and corrected the English throughout the manuscript.
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della Universitat di Ferrara (9, Scienze geologica paleontologica), 4, 12: 189-202. -
821
BUDUROV K., CALVET F., GoY A., M~,RQUEZ-ALIAGA A., MARQUEZ L.,
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A. ~ Q U E Z - A L I A G A Instituto "Cavanilles", Departamento de Geologfa & de Biodiversidad y Biologia Evolutiva Universitat de Valencia C/Dr. Moliner, 50 E-46100 Burjasot, Valencia e-maih
[email protected]
C.C. EMIG
CNRS-UMR 6540, Centre d'Oc~anologie Station Marine d'Endoume Rue de la Batterie-des-Lions F-13007 Marseille e-mail:
[email protected]
J.M. BRITO Departamento de Geologfa, Universitat de Valencia C/Dr. Moliner E-46100 Burj asot, Valencia
