Rodent Palaeoecology of the Continental Upper Miocene of Crevillente (Alicante, SE Spain)

Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 349–356 www.elsevier.nl/locate/palaeo

Rodent palaeoecology of the Continental Upper Miocene of Crevillente (Alicante, SE Spain)

Elvira Martı´n-Sua´rez a,*, Matthijs Freudenthal a,b, Jorge Civis c a Departamento de Estratigrafı´a y Paleontologia, Universidad de Granada, 18071 Granada, Spain b Nationaal Natuurhistorisch Museum, Postbus 9517, 2300 RA Leiden, The Netherlands c Departamento de Geologia, Universidad de Salamanca, Salamanca, Spain Received 15 November 1999; accepted for publication 19 July 2000

Abstract

The Upper Miocene succession of Crevillente presents an alternation of continental and marine beds. The marine fossils have been studied in order to obtain a reliable correlation with other marine sections. On the basis of these data, the entire section can be dated as Upper Tortonian/Messinian. The alternation of continental and marine beds ﬁts the known eustatic sea-level and climatic curves. These correlations are used to assign ecological preferences to the components of the micromammal faunas at species level; it becomes clear that working at the genus level does not provide valid results. With the assignments, it is possible to demonstrate consistent climatic diﬀerences between the coastal plain of Crevillente and the area of Teruel on the Meseta of Central Spain. © 2001 Elsevier Science B.V. All rights reserved.

Keywords: Messinian; Miocene; palaeoecology; Rodentia; Spain

1. Introduction of the succession in order to achieve a dating, and we have tied it in with the eustatic sea-level curve. The area studied is located north of the village This gives us a correlation between the intercalated of Crevillente (province of Alicante), as indicated micromammal localities and the eustatic curve, in Fig. 1. The sedimentary succession under study which permits a climatological interpretation of lies unconformably on the Mesozoic substratum the mammalian faunas. and consists of alternating continental and marine beds (Freudenthal et al., 1991;Martıń-Sua´rez and Freudenthal, 1998). Towards the north, the deposits are slightly inclined, and towards the south, 2. Age of the continental deposits they are folded with inclinations of up to 60° SSE. The marine and brackish deposits contain fora- The relative dating of the continental deposits minifera and ostracods, and the continental depos- has been carried out by various authors, most its have yielded a number of fossil mammal recently by Martıń-Sua´rez and Freudenthal, (1998). localities. We have studied the marine fossil content The mammal-bearing localities are mainly located in the sections A–A∞ and B–B∞ of Fig. 1. The * Corresponding author. Fax: +34-58-243203. locality Crevillente 2 (CR2), the oldest mammal E-mail address: [email protected] (E. Martıń-Sua´rez) locality of the succession, is the reference locality

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3. Age of the marine deposits

The study of the marine deposits presents some problems, since most of the samples contain reworked foraminifera, mainly from the Cretaceous and the Paleogene. Two samples have given useful results. In the lowermost part of the succession, 60 m below CR2, the sample CRF-13, has turned out to be poor in planktonic foraminifera, among which have been distinguished Globigerina bulloides, Globigerinoides obliquus, Globorotalia pseu- dobesa, Globorotalia gr. menardii (sinistral ), Neogloboquadrina acostaensis (sinistral ), Orbulina Fig. 1. Map of the Crevillente area. universa, Globoquadrina dehiscens. The presence of N. acostaensis (sinistral ) and G. menardii (sinis- of the Neogene mammal unit MN 11 (Mein, tral ), though scarce, permits a dating as Late 1990). The locality CR15 may be correlated Tortonian. G. menardii has not yet been replaced biostratigraphically with Los Mansuetos, strato- by the dextral form, which means that the deposits type of the Turolian, and reference locality of unit do not belong to the latest Tortonian. MN12. The locality CR6, the youngest of the In the highest part of the succession, between the succession may be correlated with El Arquillo 3, two reef episodes (Fig. 2), the sample CR14-F3 has which is the reference locality of unit MN 13 yielded benthonic foraminifera, which indicate a very (Mein, 1990). Thus, the continental deposits in shallow environment (Ammonia becardii, Elphidium the area cover most of the Turolian. For a biostrat- macellum, Asterigerinata planorbis, Lobatula lobatula, igraphical zonation, see Martı´n-Sua´rez and Nonion boueanum, Cribononion sp.; other, poorly Freudenthal (1998). represented, forms are Elphidium aculeatum,

Fig. 2. Correlation of the mammal localities of Crevillente with those of Teruel, and with the eustatic sea-level curve of Haq (1991) and Brachert et al. (1996). The vertical scale of the eustatic curve is distorted, because it is adapted to an even distribution of the mammal localities. For locality codes, see the caption of Fig. 3. : regression; ÷: transgression; yy: reefs; nn: gypsum. E. Martı´n-Sua´rez et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 349–356 351

Uvigerina sp. and Reussella spinulosa). Planktonic 1989) and with the Granada Basin (Braga et al., foraminifera are scarce: Globoquadrina cf. dehiscens, 1990). They correspond to the eustatic maximum Globigerinoides obliquus, Globigerina bulloides, Globi- of cycle TB 3.2 of Haq (1991). gerina sp., Globorotalia cf. mayeri and Globorotalia The next set of strata, containing the mammal gr. miotumida.TheGloborotalia miotumida group is localities CR2 to CR18, represents the transition understood as defined by Sierro et al. (1993), and to dominantly continental sedimentation, after the the forms present correspond to morphotype G. eustatic maximum of TB 3.2. conomiozea. From these data, its level can be attrib- The next span of the succession contains uted to the Messinian, to the replacement of fundamentally continental deposits, with several Globorotalia menardii by the group Globorotalia mio- mammal localities (CR8, CR7, CR17). The micro- tumida (G. menardii has not been found). fossils of these levels comprise some fragments of stems and gyrogonites of Charophyta. The associa- Thus, the marine beds correlate to the final % Tortonian and the Messinian. Fig. 2 gives the tion of ostracods tends to be monospecific: 90 belongs to Cyprideis torosa with practically smooth relative datings of the succession, both for conti- shells, suggesting a high calcium content in the nental and marine levels, as well as the correlations Mg/Ca ionic equilibrium. This indicates cold con- between these datings. ditions that appear to correspond to the regressive episode of cycle TB 3.3, which, in other basins, is represented by the deposition of bryomol calcirud- 4. Correlations with the eustatic curve ites (Brachert et al., 1996). The eustatic maximum of this cycle is repre- In the Upper Neogene of the SE part of the sented by two reefs that correspond to the Early Iberian Peninsula, six episodes of reef growth have Messinian reef episode in other Neogene basins of been identified, alternating with deposition of calci- SE Spain, like Sorbas, Cabo de Gata and Nı´jar. rudites (Brachert et al., 1996). The reef deposits, Their build-up took place during a transgression, indicating relatively high temperature, can be corre- which was correlated by Brachert et al. (1996) lated to sea-level highstands in the eustatic sea-level with the eustatic maximum of cycle TB 3.3 of the curve of Haq (1991), and the bryomol carbonates, global sea-level chart of Haq et al. (1988, 1991). that indicate lower temperature, correspond to sea- Cycle TB 3.4 begins with a regression that level lowstands. These sea-level variations in the coincides with the deposition of the gypsum in the Late Neogene are linked to glacial phenomena, Sorbas Basin (Martıń and Braga, 1994) and the low-stands and high-stands correlating with cold calcarenites of Cabo de Gata (van de Poel et al., and warm phases, respectively, depending on the 1984). In Crevillente, it results in an unconformity amount of water stored in glaciers. on top of the reefs. The next transgression leads Three of the reef phases are dated as earliest to the development of more reefs in the Sorbas Late Tortonian, Early Messinian, and latest Basin (Martıń and Braga, 1994) and Cabo de Messinian, respectively, and correspond to the Gata (Martıń et al., 1993; Brachert et al., 1996). highstands TB 3.2, TB 3.3, and TB 3.4 of Haq In Crevillente, the sedimentation during this trans- (Brachert et al., 1996). gression is first continental (CR6) and later The deposits of Crevillente were formed in more becomes marine. The succession is unconformably shallow conditions than the sections studied by overlain by a thick succession of supposedly Brachert et al. (1996), and therefore, the cyclicity Pliocene conglomerates. is expressed as an alternation of marine and continental deposits. In the oldest, marine, deposits of Crevillente, 5. Paleoecology we have found Neogloboquadrina acostaensis and 5.1. Ecological preferences per taxon Globorotalia menardii (sinistral ), which permit a correlation with the Sorbas Basin (Martıń and The basis for the assignment of preferences to Braga, 1994), with Almanzora (Martıń et al., the mammal species found is the correlation of 352 E. Martıń-Sua´rez et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 349–356 our Crevillente succession with the eustatic sea- is probably a vicariant descendant of A. barbarae; level curve and the corresponding temperature their co-existence in CR8 means that the speciation curve (Fig. 2). Species found in localities corre- must have taken place in another area and that A. lated with eustatic maxima are assigned a meini in CR8 is an immigrant. Thus, A. meini and warm/humid preference, and species from localities its ancestor A. barbarae may have had different in eustatic minima are assigned a cold/dry ecological preferences (Vrba, 1980). Therefore, preference. and because of its presence in the cool episode of Apart from the basis provided by these climatic CR8/CR7, A. meini is interpreted as an indicator fluctuations, we use several assumptions to attri- of cool climate. bute ecological preferences: The fauna of CR7 contains only two species: 1. Taxons within the same anagenetic evolutionary Occitanomys adroveri and Apodemus meini. This lineage have equal preferences. According to poor diversity is an additional indicator of rather Vrba (1980), vicariant speciation involves a adverse climatic conditions. Castromys is a descen- change in physical environment. In such a case, dant of Apodemus meini that appears in CR17 and ancestor and descendant may have opposite is assigned a preference for cool conditions too. ecological preferences. In the Crevillente sec- Occitanomys sondaari and its descendant tion, this assumption gives no contradictory Occitanomys adroveri are less frequent in CR2 and results and appears to be correct. CR15 than the Apodemus barbarae lineage, and 2. Taxons with opposite frequency peaks have Occitanomys reaches its maximum at the cool peak opposite ecological preferences. of CR17. The peaks of Occitanomys and Apodemus 3. Immigrants replacing taxons with highly similar are clearly opposed. We attribute cool preference dental pattern have opposite preferences. This to this group. assumption implies that the dental pattern in In CR22, a new representative of the genus O. itself is not a clue to ecological preference, as alcalai appears; it co-occurs with O. adroveri, and assumed, for example, by van Dam and Weltje after the level of CR31 Occitanomys adroveri disap- (1999). We assume that taxons with a similar pears. In agreement with assumption 3, O. alcalai dental pattern may be competitors, and chang- is supposed to have warm preference: it replaces— ing ecological conditions may force an existing gradually—a related, previously existing lineage, taxon to yield to a better adapted competitor. and this replacement coincides with a temper- The murid species Apodemus lugdunensis and ature change. its descendant Apodemus barbarae are dominant Paraethomys meini appears in our area in the in the warm lower part of the succession of warm phase of CR6. Its preference is interpreted Crevillente (CR2, CR15), and this lineage disap- as warm. pears in the cool phase of CR8 to CR17. van Dam Huerzelerimys is present in the warm lower part and Weltje (1999) attribute a neutral humidity of the succession and then disappears. There is no score and an intermediate temperature preference obvious replacement, so the preference of this to their group Muridae II (which includes taxon cannot be evaluated reliably. Apodemus) on the basis of actualism. We give it a Stephanomys ramblensis is present in low num- warm preference because of its presence in the bers from CR22 onwards. Thus, it appears when warm phase of CR2–CR15. climatic conditions improve, and one would be Apodemus gudrunae may be a descendant of the inclined to assign it warm preference. However, previous lineage that returns to the area in CR6, no equivalent taxon is present in earlier levels, and when climatic conditions have strongly improved. this assignment would not be reliable. Anyway, its We consider it therefore an indicator for warm attribution to Muridae I, as supposed by van Dam climate. and Weltje, is at least doubtful. Apodemus meini appears in the cool phase of In the warm phase of CR2–CR15, CR8, where it co-exists with A. barbarae, and it is Neocricetodon occidentalis and Apocricetus plinii the only Apodemus in the cool maximum CR7. It are found, whereas Cricetulodon lucentensis is E. Martıń-Sua´rez et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 349–356 353 present in the cool phase of CR8–CR17, a perfect Mein et al. (1990) and unpublished data made parallel with the distribution seen in Apodemus. available by Dr Mein (Lyon). The preferences for these Cricetinae are assigned In both areas, the oldest Turolian faunas (CR2, accordingly. In the supposedly coldest episode, CR4B, PM, TOA5) are dominated by the same CR7, no Cricetinae have been found. This may be two species: Occitanomys sondaari and Apodemus due to a small sample size, but it is also possible lugdunensis. In Crevillente, Apodemus is more that C. lucentensis is an indicator of coolness, but frequent than Occitanomys; in Teruel, it is the not too extreme. reverse. This may mean that the Teruel area After CR17, the temperature rises in the phase was cooler/drier than Crevillente. Furthermore, represented by CR22, CR14, and CR31. Cricetinae Huerzelerimys and Neocricetodon are frequent in are frequent, classified as Neocricetodon seseae by Crevillente and scarce in Teruel. The higher fre- Freudenthal et al. (1998), who presented the alter- quency of Neocricetodon supports the supposition native of two species being present: N. occidentalis that the Crevillente area was warmer (and more and A. plinii. humid) than Teruel. The very high percentage of A. alberti in the In the second group of faunas (CR15, MDV2, warm maximum of CR6 makes its preference clear. CC3, LM, MDV5), the dominant Muridae are In the Late Miocene, the Cricetinae is the only replaced by their descendants, Occitanomys adro- taxon that belongs to the Cricetidae II group of veri and Apodemus barbarae, respectively. Again, van Dam and Weltje (1999) that is supposed to in Crevillente, Apodemus is more frequent, while score neutral on humidity and intermediate on Occitanomys dominates in Teruel. Percentages for temperature. This indifferent score may be due to Huerzelerimys vary but are in the same order of the fact that, as defined, the group is too hetero- magnitude in both areas. Neocricetodon is frequent geneous. It is quite possible, that Megacricetodon in Crevillente and almost absent in Teruel. This has a preference for dry conditions, and the again means that Teruel was cooler/drier than Cricetinae indicate more humid climate. Our data Crevillente. Pseudoruscinomys is present in various show that not all Cricetinae are indicators of Teruel localities and abundant in Los Mansuetos, humid conditions. whereas it is absent in Crevillente. This may mean Hispanomys is the only member of van Dam that this taxon had cool preference and supports and Weltje’s Cricetidae I group in our material. It the interpretation by van Dam and Weltje (1999). is present in more or less constant percentages In Crevillente, the next group of localities is from CR2 to CR14, with the exception of CR7, CR8, CR7, CR17. Unfortunately, equivalent where it is absent. It is apparently not, or only faunas are unknown in Teruel. This episode repre- slightly, affected by the degree of climatic changes sents an important hiatus in the Teruel succession that we observe in our series. It is also possible that was not detected by van Dam and Weltje that treatment at genus level obscures differences (1999). In Crevillente, in this phase, Apodemus is at the species level. Several lineages are present, less frequent than Occitanomys, and the Apodemus and the relationships between them are not clear. material contains a small minority of Apodemus If these lineages have different ecological prefer- barbarae and a majority of the newcomer ences, our material is not good enough to discern Apodemus meini. In CR8, Neocricetodon is replaced these. by Cricetulodon, which is absent in CR7 and present again in CR17. CR17 is characterized by 5.2. Comparison of Crevillente and Teruel the highest frequency of Occitanomys of the entire Crevillente succession and the lowest frequency The best record of faunas of similar age as (or even absence) of Cricetinae. Castromys is a Crevillente is without any doubt found in the area new element that appears in Teruel somewhat of Teruel, and we will make a comparison with it, later. This phase is the maximum peak of coolness. based on data by van de Weerd (1976), van de CR22, CR14 and CR31 form the next group of Weerd and Daams (1978), Alcala´ et al. (1991), localities in Crevillente, possibly equivalent to 354 E. Martıń-Sua´rez et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 349–356

GLO6 in the Teruel area. In Crevillente, percent- in Teruel, and Apocricetus is scarce. The low- ages of Occitanomys remain high, accompanied by temperature indicator, Castromys, has disappeared rather important numbers of Castromys and in Crevillente and is present in Teruel. In Teruel, Neocricetodon. Climatically, this represents the Occitanomys alcalai is an important element of transition of the cool peak of CR8–CR17 towards the fauna. a higher temperature, reﬂected in the transgression The warmth indicator, Apodemus gudrunae,is and reef deposits, above CR17. scarce in Crevillente and more frequent in Teruel. In Teruel (GLO6 and VDC3), we witness the Apparently, the conditions in Teruel were cooler entrance of Stephanomys, which soon becomes the (and/or drier) than in Crevillente, as shown by the dominant species. In Crevillente (CR6, younger co-occurrence of warm and cool indicators. than VDC3), Stephanomys is very scarce, and The scarceness of Stephanomys in Crevillente, the fauna is dominated by Paraethomys and at the same time that it is abundant in Teruel, Apocricetus, both interpreted as indicators of warm might be used as an argument to assign cool conditions. On the contrary, Paraethomys is absent preference to this taxon. However, it is an

Fig. 3. Frequency curves of taxons for Crevillente (above) and Teruel (below). Horizontal scales represent percentages of rodent fauna. Peaks from left to right represent taxons with supposedly dry preference, peaks from right to left represent those with supposedly wet preference. 1: Cricetulodon lucentensis. 2: Other Cricetinae (Neocricetodon occidentalis, N. plinii, N. seseae and Apocricetus alberti). 3: Occitanomys sondaari+Occitanomys adroveri.4:Apodemus lugdunensis+Apodemus barbarae.5:Apodemus gudrunae.6:Apodemus meini+Castromys littoralis+Castromys inflatus.7:Paraethomys meini.8:Pseudoruscinomys.9:O. alcalai. 10: Huerzelerimys. 11: Stephanomys. 12: Hispanomys. 10–12: Catholic taxons and those without clear preference. Locality codes: CC: Concud; CEL: Celadas; GLO: La Gloria; LM: Los Mansuetos; MDV: Masada del Valle; PM: Puente Minero; TOA: Tortajada A; VDC: Valdecebro; VILL: Villastar. E. Martıń-Sua´rez et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 349–356 355 important element of the fauna of Purcal 4 blance to the curve that may be constructed from (Granada Basin,Martıń-Sua´rez et al., 1998), our Fig. 4, but there is a problem. van Dam and which is of about the same age, and that fauna Weltje (1999) come to the conclusion (their page contains mainly warmth indicators. Furthermore, 289) that aridity and temperature are fully and Stephanomys is present in Teruel in very constant positively correlated. However, following Brachert percentages, independent of the fluctuations of et al. (1996), we assume a positive correlation percentages of warm and cool indicators. Thus, between humidity and temperature. Of course, we maintain our doubt about this taxon, but Teruel lies on the Central Spanish meseta, and meanwhile think that it is more probably an indica- Crevillente lies in the coastal plain, so climates tor for high temperature and humidity than for may be different, but the mentioned difference is coolness and aridity. difficult to explain. In Fig. 3, we give the frequencies in percentages of the rodent fauna, using the scale from left to right for taxons with supposed cool preference, 6. Conclusions and from right to left for those with warm preference. The catholic species and those with an The Upper Miocene sediments in the area of unknown preference are represented separately. In Crevillente were deposited in a delta with alternat- Fig. 4, the curves of the taxons with known prefer- ing marine and continental conditions. The conti- ence have been added, and a similar graph has nental deposits are rich in fossil micromammals, been drawn for Teruel. Not only is Teruel con- and can be correlated with the stratotype of the stantly cooler/drier than Crevillente, but the peak Turolian; the marine beds can be correlated to the of dry climate corresponding to Los Mansuetos Upper Tortonian and the Messinian. The bound- may mean that the coolest phase begins earlier in ary between these two stages coincides in Teruel than in Crevillente (supposing, of course, Crevillente with continental deposits, and must be that our biostratigraphical correlations are not located shortly above CR17. The two reef deposits diachronic). in the upper part of the succession can be corre- At first sight, the humidity/temperature curve lated with the Early Messinian reef phase 5 of by van Dam and Weltje (1999) shows some resem- Brachert et al. (1996).

Fig. 4. Accumulated frequency curves for Crevillente (left) and Teruel (right). Wet–dry direction and abbreviations as in Fig. 3. The vertical position of the localities represents the biostratigraphical correlation between the two areas. Horizontal scales represent percentages of rodent fauna. 356 E. Martı´n-Sua´rez et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 349–356

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