RUIZ BUSTOS 1999 - Biostratigraphy of The Continental Deposits in The Granada, Guadix and Baza Basins (Betic Cordillera)

Biostratigraphy of the Continental Deposits in the Granada, Guadix
and Baza Basins (Betic Cordillera)
Antonio Ruiz BUSTOS

Instituto Andaluz de Ciencias de la Tierra CSIC-UG
Facultad de Ciencias
18002 Granada
SUMMARY
In the Granada, Guadix and Baza Basins, a biostratigraphic synthesis has been performed using data on the
relative position of the deposits that fill the basins and on the fossil marnmals found in these basins.
The stratigraphic position of the deposits is explained by the Lacustrine Stages Model, in which stages
characterized primarily by erosion and fluvial sedimentation alternate with lacustrine stages characterized
predominantly by carbonate sedimentation. Each lacustrine stage, called by the same name as the age of the fauna
found there, is preceded by a detrital stage.
The criterion governing the phylogenies compiled in the Betic basins was the hierarchical arrangement of
the different traits in relation te the importance of their biological nature. The leading taxa of the chronology
were considered to be the subfamily Arradiculatine and the genera Stephanomys, Castillomys and Mimomys.The
palaeoecological conditions were deduced by the use of biocenograms, these being based on the relationship
which, according to the fauna and their feeding, allow inferences both the on habitat and on the climate capable
of bringing about the habitat.
The biostratigraphic scale is the synthesis of data coming from the relative position of the deposits and from
the study of the phylogenetic and palaeoecological characteristics of the fossil mammals. This scale consists of
the Upper Miocene divided into two mammalian levels: Turolian (MN11 and MN12) and Ventian (MN13). The
former is equivalent in part to the Tortonian, and the second to the Messinian. The Pliocene is split into two
other levels: Ruscinian (MN14 and MN15) and Villafranchian (MN16 and MN17). The Quaternary is divided
into 3 biozones of Pleistocene Mammals (MP), corresponding to the Lower Pleistocene (MP18),Nliddle (MP19)
and Upper (MP20).
RESUMEN
En las cuencas de Granada, Guadix y Baza se realiza una síntesis biostratrigráfica mediante los datos que
proceden de la posición relativa de los sedimentos que rellenan las cuencas y el estudio de los mamíferos fósiles
encontrados en estas cuencas.
La posición estratigráfica de los sedimentos se explica mediante el modelo de las Etapas Lacustres que indica
una alternancia en el tiempo entre etapas con predominio de los procesos erosivos y depósitos de sedimentos
fluviales, y etapas lacustres donde predomina el deposito de sedimentos carbonatados. Cada etapa lacustre se
denominan con el nombre de la edad de la fauna que contienen y está precedida por una etapa detrítica.
Las filogenias realizadas en las cuencas béticas tienen como criterio director: la jerarquización de los
caracteres en relación a la importancia de su naturaleza biológica y se consideran como taxones lideres de la
cronología la subfamilia Arradiculatine y los géneros Stephanomys, Castillomys y illímornys.
Las condiciones paleoecológicas son obtenidas mediante Biocenogramas estos se basan en la relación que a
través de la fauna y su alimentación nos conduce a inferir tanto el habitat como el clima capaz de originar el
habitat.
La escala biostratigráfica síntesis de los datos que proceden de la posición relativa de los sedimentos y
el estudio de las características filogeneticas y paleoecologicas de los mámiferos fósiles consiste en un Mio-
ceno superior dividido en dos Pisos de Mamíferos: Turoliense (MN11 y MN12) y Ventiense (MN13); el
primero equivale en parte al Tortoniense y el segundo al Messiniense. El Plioceno se separa en otros dos
pisos: Rusciniense (MN14 y MN15) y Villafranquiense (MN16 y MN17). El Cuaternario se divide en tres
Biozonas MP (Mamíferos Pleistocenos) correspondientes al Pleistoceno Inferior (MP18), Medio (MP19) y
Superior (MP20).
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I. INTRODUCTION
The Betic Cordillera, located in the southern and southeastern Spain, is an Alpine mountain chain that
acquired its present day framework during the Neogene. The continental deposits of the Betic Cordillera (Fig.
1) date from between the Upper Tortonian and the present and are composed mainly of lacustrine and fluvial
sediments deposited in the intramontane basins. These basins are located over the contact area between the
Internal and External Zones of the cordillera and became gradually individualized as the mountain chain
emerged. Situated in the centre of the cordillera, the Granada, Guadix and Baza Basins have the greatest
dimensions, and their deposits remain quite constant over time and they contain a rich mammalian fossil record.
These deposits being rarely represented in other Spanish basins.
The geological study of the central Betic basins began towards the middle of the last century, the most
notable writers (up to the present) being: C. Silvertop, R. von Drasche, L. Siegert, P. Fallot, E. Aguirre, J.A.
Vera, C. Sanz de Galdeano, C. Montenat, P. Ott d'Estevou, J.A. Peña, J. Gibert, J. Rodriguez Fernandez and
J. Fernandez. The stratigraphic record and palaeogeographical context the there basins has been summarized by
Sanz de Galdeano and Vera (1992).
In the present work the stratigraphic data from the continental deposits of the central Betic basins (Granada,
Guadix and Baza) in association with the dating and palaeoecology deduced from the fossil mammals according
to the biological evolution and feeding needs of each taxon were used in order to establish a biostratigraphic
system that would permit individual data to be evaluated in relation to the overall information.
2. STRATIGRAPHY OF THE GRANADA, GUADIX AND BAZA BASINS
The Lacustrine Stages Model is explained in Ruiz Bustos (1976, 1984, 1991a, 1992, 1993b, 1994b, 1995a)
and in Ruiz Bustos et al. (1992b). According to this model, the stratigraphic-synthesis column of the sedimentary
fill of the Granada, Guadix and Baza Basins (Fig. 2) consists of alternating stages; that is, predominantly erosive
processes and fluvial, detrital deposits, red in colour (originating in the flood plains), alternating with lacustrine
stages characterized primarily by the deposit of carbonate sediment, white in colour, with intercalations of dark
clayey material rich in organic matter. Each lacustrine stage was preceded by a detrital one. In the lacustrine
sediments, the fossil fauna is abundant and reveal their age, as opposed to the situation with the detrital
sediments. The successive lacustrine stages are here called by the narre of the age corresponding to their
respective fauna.
The Granada Basin, during the Miocene, presents the oldest lacustrine stage, called the Turolian Lake. This
stage has eutrophic features and the shore vegetation supported 6 different Proboscidea genera (Aguirre et al.
1974). In the Guadix Basin, the Turolian Lake is represented by the Bite of Salinas (Soria Mingorance and Ruiz
Bustos, 1992). This suggests temporally close beginnings for the continental deposit in the Granada and Guadix
basins. In the Baza Basin, the onset of the infilling must have been coetaneous with that of the other two basins,
but until now it has not been tested because the deposits from the Turolian Lake in the Baza Basin appear in
outcrops of limited surface area and without fauna.
The Turolian Lake was succeeded by the Ventian Lake. The latter was shallower than that of the former,
water salinity was higher and gypsum deposition was more frequent. In the Granada Basin, the Ventian Lake,
in its final stages, transformed into a palaeosoil and ended with the deposition of Paramos limestone. These
constituted the final lacustrine event which affected the entire Granada Basin (Ruiz Bustos ct al. 1992a, 1992b).
The Guadix and Baza Basins during the Pliocene exhibit a repetition, in general terms, of the sedimentological
characteristics of the preceding Miocene lacustrine sequence.The Ruscinian Lake (Lower Pliocene) was eutrophic
and marshy, and slightly folded. Next, the Villafranchian Lake reveals frequent deposits of gypsum, flint, dolomite
and ripples in its strata; the lake water was saltier, more turbulent and shallower than in the case of the Ruscinian
Lake. The two lakes were separated by a lithological package formed by red detrital sediments, which are
predominated by erosive phenomena and date approximately to the Lower Villafranchian. These characteristics
are described in the works of Viseras (1991), Guerra Merchan (1992) and Soria Mingorance (1993). In the
Granada Basin, lacustrine stages with Pliocene mammals are not found, and, during this time, red detrital
sediments are possibly deposited. This is interpreted as being duo to tectonic causes (Ruiz Bustos, 1992).
At the beginning of the Quaternary in the Granada and Baza Basins, there was a lacustrine stage called the
Lower Pleistocene Lake. The Middle Pleistocene marks the last lacustrine stage of great dimensions in relation
to the size of the basins, giving rise to the Middle Pleistocene Lake. This lake varied in size over time and was
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BIOSTRATIGRAPHY OF THE CONTINENTAL DEPOSITS IN THE GRANADA, GUADIX AND BAZA BASINS
not strictly coetaneous in the different basins, the bottom of this stage being the Cullar de Baza site (Baza Basin)
and the top represented by the Solana site (Guadix Basin). In general, the Quaternary lakes had greater environmental
diversity and were smaller than their forerunners in the Miocene and Pliocene.
3. BIOZONE MARK (BM)
In the Betic Cordillera, there is a high number of sites which are stratigraphically intercalated between the
first and last sites of each Biozone of Pleistocene and Neogene Mammals. This requires increasing the chronological
precision within the time span of each biozone by using regular and successive time intervals called Biozone
Marks (BM). Shown in Figure 2, these were first described by Ruiz Bustos (1992). The Biozone Marks represent
intervals of approximately 0.25±0.05 M.y., but tend to diminish in duration as the Biozone Marks approach the
present. These consist of two numbers separated by a hyphen, the first representing the biozone and the second
the position with the biozone. For example, Biozone Mark 13-1 refers to the bottom of the MN13 Biozone.
The Biozone Marks lose their meaning when separated from the Biozones to which they belong. This implies
that for proper use the Biozone Marks must not be confused with short-terco biozones or subzones, since the
principal characteristic of the Biozone Marks is that the time periods are successive within a biozone and
therefore between these the fauna show no differences of enough magnitude for the Biozone Marks to be
considered true Biozones. The morphological changes which occur between one Biozone Mark and the next
are mostly of intraspecific nature. This does not prohibit the appearance or disappearance of a taxon within a
biozone from being accurately identified with the corresponding Biozone Mark.
Sites represent isolated, local events within the sedimentary record, thus requiring the distinction between
the following concepts: a) Real Presence, with real time during which the species lived generation after generation
in the basin; b) Local Presence, indicated by each site; and c) Guide Presence, inferred from several instantes
of Local Presence. The duration which indicates the Guide Presence is equal to or greater than the Local
Presence marked by the sites and tends to equalize with the Real Presence. The Biozone Marks, due to their
short time period, allow inferences concerning the presence of taxa (exercising the necessary precautions) and
become the best representation possible of the fauna that live at each point in time in the Betic Basins. The rules
that limit the possibilities of inferring the presence of a taxon are as follows: a) When the taxon is a genus or
higher, and carries the terco continuity of presence cannot be inferred. b) Non-specified species do not
allow continuity of presence to be inferred. An exception crises when the absence is situated within a single
Biozone while there is majority presence of the taxon in the rest of the Biozone Marks of the Biozone. c) In
each species, while there is no Biozone or further separation, continuity of presence is inferred.
4. MAMMAL FAUNA AND THEIR EVOLUT IONARY PROCESSES
In the Betic basins, 1 have worked on the evolutionary processes undergone by mammal tooth morphology
using as a governing criterion the hierarchical arrangement of the traits in relation to the importance of their
biological nature.
The biological nature of the morphological traits, also called biological role or importance, expresses the
inheritabílity of the trait and the degree to which the ordinary vital activity of the individual requires the correct
functioning of the trait. The greater the biological nature of a trait, the greater is the capacity for modification,
subject to natural selection, and thus engender phylogenetic changes detectable by palaeontology. For example,
we might ask:Which trait has greater biological nature eye morphology of a deer which was not inherited and
carne about accidentally during the embryonic development of the individual, or a secondary sexual trait such
as a horn which was smaller than average in the population? Eye morphology can cause the individual die before
reproduction or simply have no affect on offspring; however, the horn affects both the life of the individual as
well as its offspring, and therefore the latter trait has more biological nature than che former.
Size, despite its biological nature determined by inheritance and the p eed to be optimal for daily activity,
strongly depends on the environment, in the broad sense. This can result in individuals with notably different
minirnum or maximum values. In the Betic Cordillera, as a rulo, body size registers a continuous increase over
time. Although at certain points of crisis this process can stop, but if the crisis is overcome without the
generalized extinction of the species within a genus, then the evolutionary trend towards increased size resumes.
This scenario appears in the size dispersion undergone by Stephanornys during the transition from Lower to
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Ruiz BUSTOS, A.
Upper Pliocene, reflected at the karstic site (Moreda) and the stratified lacustr ne site (CTH-3), as well as in
Apodemus gorafensis in which tooth size fluctuated at the end of the Ventian (Ruiz Bustos et al., 1992a).
The size of a protuberante or fold of tooth enamel is greater with relation to the rest of the crown the
greater its biological nature, for example, the distinction between initial and definitive folds in arvicolids. The
Enamel Units method (Ruiz Bustos, 1987, 1988, 1995b) is a tool to quantify the biological nature of the
morphological traits of the tooth crown.
The book entitled La Evolución Plástica (Ruiz Bustos, 1994a) explains in detail how to calibrate the biological
nature of traits. This enables inheritance laws to be applied to fossil populations in order to provide a biological
explanation for traits observed in the fossil record. The establishment of a hierarchy of morphological traits
according to their biological nature enables a calibration of the meaning of the morphologies, both accessory
and accidental, considering these essential for characterizing the species. In analyses using the Dental Equation
(Ruiz Bustos, 1987, pp. 8-13), different morphological features can be related and arranged into hierarchies
according to their greater or lesser biological nature, thereby defining the species that comprise a phylogenetic
sequence. This evaluation is the necessary condition to determine whether a given trait within a biocanton (Ruiz
Bustos, 1994a) can be used in taxonomy. An example of the use of this criterion is the study of the genus
Stephanomys by Ruiz Bustos (1986).
By the analysis of the biological nature of the traits, Ruiz Bustos (1976, 1977, 1978, y 1991b) explains the
phylogeny of the subfamily Arradiculatine and of the genera Stephanomys, Castillomys and Mimomys, as well as their
respective species and subspecies. Other systematics have been used with reference to Elephanticlae (Aguirre, 1968
and Mazo, 1989); Carnívora (Alcala and Morales, 1989 and Martinez Navarro,1991); Rhinocerotidae (Cerdeño,
1989); Bovídae (Azanza and Morales,1989); Cervidae (Azanza and Morales, 1989 and Ruiz Bustos et al., 1990)
and Lagomorpha (Lopez Martinez, 1989). These phylogenies offer the following data:
1) A list of the taxa described in the Betic basins from the beginning of the sedimentary infilling until the
present (see Appendix II).
2) The number of sites at which a certain taxon is present (see Appendix I). This enables the calculation
of the Presence (P) and Abundance (A) Indexes of a taxon in each Biozone Mark. The definition of these
concepts is: P=(nx100)/Y; A=(Txn)/Y, in which n represents the number of the sites where the taxon
was present in a Biozone Mark; Y is the number of sites in a Biozone Mark; T is the number of different
mammalian taxa in a Biozone Mark. For example, in the BM 12-2, there were 3 sites, where a total
of 16 different taxa were collected. Hipparion sp. cf. concudense and Rhinocerotidae gen. indet. appear at
two sites at the sanee time and no taxon was found at all three sites. Translated into percentages of
Presence, this indicates that no taxon reaches 100% Presence, and only two taxa reach 66.66% while the
remaining 14 taxa register 33.33%. The calculation of the Abundance Index for the taxon Hipparion sp.
cf. concudense is A=(16x2)/3= 10.66.
3) Dating. Values for taxon presence in each Biozone Mark, and therefore in each biozone, give a detailed
biostratigraphy by groups of taxa. Ascertaining the age of a given fauna requires identifying the Biozone
Mark which according to cladistic analysis (where the importante of each taxon is evaluated) is closest
to the fauna being dated.
4) Appearance and extinction. The point in time of the first and last record of each taxon in the basin
allows the calculation of the percentage of taxa that appear for the first time and how many appear for
the last time in each Biozone Mark in relation to the total number of taxa; that is, these which at some
point have been or are present in the Betic sedimentary basins. From these evolutionary processes, it is
possible to identify which taxa are immigrants and which are the products of evolution in the region.
5. PALAEOECOLOGY FOSTERING MAMMALIAN FAUNA
The palaeological conditions of a region can be deduced from biocenograms (Ruiz Bustos, 1976, 1990b,
1993a 1995c and 1995d). These consist of double-entry tables where the most general and basic plant habitats
of a region are plotted on the abscissa, and the mammals living in these habitats at a given site (MN and MP
Biozone or Biozone Mark) are plotted on the ordinate. The relationship between the two axes is established
according to the following criteria: value (0), mammals do not live in the biotope; value (1), the animal lives
in the habitat, but with difficulty; and value (2) the animal lives under optimal conditions in the habitat. The
sum of the values in the column corresponding to each habitat, and expressed in percentages with respect to
the other habitats, constitute the Habitat Index (HI).
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BIOSTRATIGRAPHY OF THE CONTINENTAL DEPOSITS IN THE GRANADA, &JADE\ AND BAZA BASINS
Basic plant habitats of a region are considered those which refe.r only to the size and number of the
individuals inhabiting a plant mass. re p.-ardless of the floristic composition. Elemental tupes of basic plant habitats
include: Deserts, Forests, Prairies and Forest Meadows. For a specific region. to the extent that we know other
characteristics of the plant masse.s_ ve can specifv and amplifv these basic habitats. each of which constitutes a
basic biotope for the mammalian fauna.
The characteristics of the present-dav vegetation and the. fossil data in the Iberian southeast, as in the rest
of the western Mediterranean area (Suc. 1984 and Barron et al.. 1994). indicate a similar structure and organization
of the basic plant habitats from the Vendan ro the present. This does not ir/1ply that the floristic composition
has never undergone transformations by replacement of one plant species by another, the appearance of new
species or changes in the dimensions of the plant biotopes as a result of shifts in regional climate.
In the sphere of continental Europe, and in the present work for the Betic Cordillera, basic plant habitats
and related climatic conditions are considered to be: A= snowline and tundra (cold and dry); B= short-grass area
and steppe (cold and dry); C= coniferous forest and taiga (cold and wet); D= Deciduous forest (hot and wet);
E= sclerophyllous forest (hot and dry); F= shrublands and dry grassland (hot and dry); G= cold continental
waters (cold and wet); H= warm continental waters (hot and wet) and I= Desert (hot and dry).
The Tendencies of mammals in the Betic Cordillera in relation to the Basic Biotopes A, B, C, D, E, F, G,
H and I are expressed by a succession of numbers with a constant order (000112200). In this succession, the
first number corresponds to the Basic Biotope A, the second to B and the third to C, etc... The Tendency of
each taxon, enumerated in alphabetical order, is expressed in Appendix II.
Any type of basic plant habitat can be included in one of four climatic categories: cold and dry, b) cold
and wet, c) hot and dry and e) hot and wet. From this relationship, four climatic parameters can be calculated:
Cold (C'), resulting froni the sum of the Habitat Indices (In), in which the habitats are classified as cold, And
the calculation is repeated successively for the other parameters, Hot (H'), Dry (D') and Wet (W'). These
parameters are expressed by two binomials referring to temperature: C'+H'=100 and in relation to wetness:
D'+W'=100.
The interpretation of the data obtained by biocenograms requires the use of a system of measurement based
on the diversity and ecological trend of the rnammals with their habitat; the unit of this system, called Mammal
Diversity Unit (MDU), is defined as the climatic conditions of a hypothetical habitat where only one mammal
species lives under minimum conditions for viability. The relationship fauna-vegetation-climate is quantified by
biocenograms and expressed in MDU.
From the data of Guide Presence of each taxon and its Trend towards the basic plant biotopes, it is possible
to construct biocenograms for each Biozone Mark. These provide their respective Habitat Indices. The totality
of the biocenograms obtained constitute the Biocenotic Spectrum of the basin. This indicates how the biomass
of the vegetation in the basin evolved over time:The evolution of the climatic conditions in the Betic Cordillera
and its biotopes are shown in the Synthesis Table (Table 1).
6. CONCLUSION
The Guide Table for the Betic Cordillera (Hg. 3) is constructed with the stratigraphic sequence of the
continental fill involved in the model of the Lacustrine Stages in the Betic basins and the dates for the rnammals
(explained in Appendix I and II). Because of the printing difficulties poned by the large dimensions of this table,
distribution is advisable in the forro of a computerized version of the Guide Table for the Betic Cordillera in
Quattro Pro. This diskette is available from the author upon request.
The summarized explanation of the data in the Guide Table constitute the Synthesis Table (Table 1). This
reflects the following conclusions:
1) The climatic conditions of the Betic basins evolved from a hot, dry climate when the infilling of the
basins began, with values of the climatic parameters expressed in Mammal Diversity Units (MDU) of roughly
Hot>80, Cold<20, Dry>60 and Wet<40, to present-day values of about Hot_50, Cold_50, Dry_50 and Wet_50.
This climatic evolution, by successive fluctuations, gradually shifted far froni the initial hot and dry values.
2) The climatic break of the greatest magnitude in the entire sedimentary sequence occurred around 1.6
M.y. This coincides with the limit for the Quaternary proposed by Aguirre and Pasini (1985) and distinguishes
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a Mio-Pliocene phase (Phase A) which began approximately 8 My and lasted until around 1.6 M.y. and a
Pleistocene phase (Phase B) which lasted from 1.6 M.y. to the present.
These conclusions go against the opinion of Bonadonna and Alberdi (1987a and b) and Alberdi (1995), who
date the beginning of the Quaternary towards 2.5 M.y, citing, as arguments, the appearance of the genus Equus
and climatic fluctuation. In relation to the first argument, the comparisons within the data set indicate that all
of the Pliocene species in the region which were present on the arrival of the genus Equus survived beyond 2.5
M.y.; significantly, this fauna went extinct towards 1.6 M.y., bringing about a complete faunistic renewal, lead
by the appearance of rootless arvicolids. With regard to the second argument, as indicated by the climatic values
in Mammal Diversity Units (Table 1), the shift in climate during the appearance of the genus Equus, around 2.5
M.y., does not disrupt the climatic continuity of the Pliocene, but only foreshadows the characteristics of the
climatic break that occurred towards 1.6 M.y. The consequences of the evaluation by Bonadonna and Alberdi
become evident when they characterize their perceived change at 2.5 M.y. as marking the beginning of the
mammalian level "Middle Villafranchian"; they cite no species to characterize this level in relation to the "Upper
Villafranchian" and they overlap the Upper Villafranchian with the Quaternary. This sketch is used by Bonadonna
and Alberdi on a global scale, incorporating the work of Zanchetta et al. (1995), correlating the Betic Cordillera
with some geological formations of the southern hemisphere (Argentina) on the basis of the rough temporal
coincidences of certain events among completely different fauna (such as South American and European) without
attempting to use methodologies capable of converting different and distant fauna to the same scale of climatic
parameters, as can be achieved, for example, by using the biocenogram.
3) Phase A (Mio-Pliocene) has 4 stages: Al, A2, A3 and A4. These in broad terms indicate an alternation
of climatic conditions, although in detail each stage has its peculiarities. The warm and dry conditions of Stage
Al reappear in Stage A3, and the coldest conditions of Stage A2 are repeated in Stage A4.
The Stage Al deposits accumulated in the eutrophic lakes differ lithologically with respect Stage A2 deposits
from lakes rich in salts, and the faunal types also differ in each type of lacustrine deposit. Both of these
differences indicate the difficulty in associating the fauna of Stage Al, which correspond to the Biozones MN11
and MN12, with the fauna from Stage A2, which correspond to the Biozone MN13. This problem is indicated
in the stratigraphic division of the continental Neogene proposed by Aguirre (1975), Aguirre et al. (1976),Weer
(1976), Alberdi et al. (1977) and Morales (1984).
The biostratigraphic scale proposed by Mein (1975, 1976 and 1990) does not sufficiently account for the
data from the Betic Cordillera, and the biostratigraphy presented by Calvo et al. (1993) is a detailed application
of Mein's scale to the Spanish basins and thus does not distinguish, either, between the successive lacustrine
phases of the Betic Cordillera. Therefore, Mein and Calvo did not separate the Ventian from the Turolian but
rather kept the division of Crusafont (1965) despite that today more data are available.
The equivalence between the Ventian (MN13) and Messinian proposed by Aguirre (1975) on the basis of
mammalian fauna was placed in doubt by Agusti et al. (1981) with the site called Casa del Acero, located in the
Betic basin of Fortuna. According to these authors, the site dates from the Turolian MN12 and lies aboye a level
of reefs attributed to the Messinian by Santisteban (1981), although this author, in the same work of 1981 (p.
401), indicates that the fauna of the site are redeposited. In addition, the possibility remains, according to the
regional geology and the age of the reefs in other points of the cordillera, that the age of the reefs are older.
The site of Casa del Acero should be reviewed again.
In Stage A2, the crisis a2.1, with a value for the parameter Co1d=22 and a2.3, with a value of Cold=21,
are highly similar and thus reflect a cyclic process. This agrees with the data pointing to a cyclic process in the
salinity crisis during the Messinian, but casts doubt on the possibility that high temperatures were the cause.
The Pliocene in the Betic Cordillera shows two stages: Stage A3, equivalent in its fauna to the Biozones
MN14 and MN15, and Stage A4, characterized by the arrival of the genus Equus MN17. The two stages are
separated by detrital deposits lacking fauna equivalent to those of the site of Villarroya (Villalta, 1952).
4) Phase B (Pleistocene) shows, by its three stages Bl, B2 and B3, a cyclic process in which the beginning
of each stage implies an increase in the Cold parameter, which decreases slightly over this stage until the
beginning of the next.
Stages Bl, B2 and B3 represent long interglacial periods each interrupted by short glacial periods, in a
fleeting pulsation of cold and dry. The sequence formed by these periods is termed "Mediterranean" and can
be correlated with the sequence of the Alps, taking into account that the Mediterranean Gunz-Mindel, the
Mediterranean Mindel-Riss and Mediterranean Riss-Würm lasted longer than their homologues in the Alps. In
addition, the glacial stages: Mediterranean Gunz, Mediterranean Riss and Mediterranean Würm were far briefer
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than in the higher latitudes of the continent. The b3.1 crisis reflects the intens ty of the Mediterranean Würm
in the region.
5) The biostratigraphic scale proposed by Ruiz Bustos (1990a. 1991c, 1992 and 1993b), shown in Figure
2, accounts for data published since 1974 and synthesized in the present work. This scale takes into account the
Lacustrine Stages Model for the stratigraphy of the Betic basins and shows the Upper Miocene divided into two
mammalian levels: the Turolian (MN11 and MN12) and Ventian (MN13). The former is equivalent in part to
the Tortonian and the later to the Messinian. The Pliocene is split into another two levels: Ruscinian (MN14
and MN15) andVillafranchian (MN16 and MN17). The Quaternarv is divided into 3 Pleistocene Mammal (PM)
Biozones: Lower Pleistocene (PM18), Middle (PM19) and Upper (PM20).
In accord with these observations, the Synthesis Table (Table 1) reveals an internal logic which converts the
manimalian fauna, on the basis of their rnorphology, into a tool to interpret the history of successive events.
REFERENCES
AGUAYO DE HOYOS P., ORTIZ RISCO E, SANCHEZ QUIRANTES L. y RUIZ BUSTOS A. (En prensa).
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APPENDIX I: CATALOGUE OF SITES IN THE GRANADA, GUADIX AND BAZA BASINS
The sequence of the sites enumerated represents the synthesis of two sources of information: on the one
hand, the dating given by the mammalian fauna and, on the other, the chronology implicit in the superposition
of the deposits in the regional stratigraphic sequence. The nomenclature of the mammalian levels is that adopted
by Ruiz Bustos (1990a).
SITES OF THE UPPER TUROLIAN
Canteras de Jun (Aguirre et al., 1974; Dabrio et al., 1978; Padial, 1986; Padial y Ruiz Bustos, 1989; Ruiz
Bustos et al., 1992b). Biozone Mark: 12-2. Fauna: Cervidae indet., Eliomys sp., Gomphotherium sp., Hipparion
sp. cf. concudense, Insectivora indet., Machairodus aphanistus, Metailurus sp. aff. major, Occitanomys adroveri,
Prolagus crusafonti, Rinocerotidae gen. indet., Ruscinomys schaubi, Valerymys juniensis, Zygolophodon sp.,
Molino de Alfacar (Aguirre et al., 1974).Biozone Mark: 12-2. Fauna: Pentalophodon sivalense, Deinotherium
giganteum, Chooerolophodon pentelici.
Monachil (Dabrio and Ruiz Bustos, 1979).Biozone Mark: 12-2. Fauna: Hipparion sp. cf. concudense,
Rinocerotidae gen. indet..
Barranco del Beiro (Aguirre et al., 1974; Padial, 1986).Biozone Mark: 12-3. Fauna: Proboscidea indet.,
Occitanomys adroveri, Stephanomys cf. stadii, Valerymys turoliensis, Ruscinomys schaubi, Prolagus crusafonti.
Los Arcos (Aguirre et al., 1974; Padial, 1986). Biozone Mark: 12-3. Fauna: Valerymys turoliensis, Prolagus
crusafonti, Insectivora indet..
Salinas (Soria Mingorance y Ruiz Bustos, 1992). Biozone Mark: 12-6. Fauna: Occitanomys adroveri,
Ruscinomys schaubi. Aves indet..
Pulianas-1 (Padial, 1986). Biozone Mark: 12-7. Fauna: Occitanomys adroveri, Stephanomys ramblensis
ramblensis, Apodemus gudrunae, Ruscinomys schaubi, Prolagus cf. michauxi.
Pulianas-2 (Padial, 1986). Biozone Mark: 12-7. Fauna: Stephanomys ramblensis ramblensis, Apodemus
gudrunae, Ruscinomys schaubi, Prolagus cf. michauxi, Insectivora indet., Reptilia indet.
Pulianas-3 (Padial, 1986). Biozone Mark: 12-8. Fauna: Apodemus gudrunae, Ruscinomys schaubi, Dipoides
problematicus. Prolagus cf. michauxi, Insectivora indet.
SITES OF THE VENTIAN.
Pino Mojon (Alonso Diago, 1989; Sese, 1989). Biozone Mark: 12/13. Fauna: Occitanomys adroveri,
Stephanomys ramblensis ramblensis, Apodemus gudrunae, Ruscinomys schaubi, Myocricetodon sp., Eliomys
truci, Prolagus cf. michauxi, Insectivora indet..
Calerico-F1 (Aguirre et al., 1974, Bone et al., 1978; Paial, 1986). Biozone Mark: 13-1. Fauna: Occitanomys
adroveri, Stephanomys ramblensis ramblensis, Apodemus gorafensis, Ruscinomys schaubi, Cricetus cf. kormosi,
Dipoides problematicus, Prolagus cf. michauxi.
Mina-5 (Bone et al., 1978; Paial, 1986). Biozone Mark: 13-2. Fauna: Apodemus gorafensis.
Mina-M12 (Bone et al., 1978; Paial, 1986). Biozone Mark: 13-2. Fauna: Stephanomys ramblensis ramblensis,
Paraethomys cf. miocaenicus, Dipoides problematicus, Castor sp.
Dehesa-D4 ( Aguirre et al., 1974, Bone et al., 1978; Paial, 1986; Alberdi and Bone, 1978). Biozone Mark:
13-3. Fauna: Anancus arvernensis, Hipparion gromovae granatense, Microstonyx sp., Rhinocerotidae indet.,
Bovidae indet.I, Parabos sp., Palaeoryx sp., Cervidae indet., Hippopotamus crusafonti, Ictitherium sp., Occitanomys
adroveri, Stephanomys ramblensis ramblensis, Paraethomys cf. miocaenicus, Dipoides problematicus, Testudo cf.
bolivari.
Dehesa-D5 (Bone et al., 1978; Paial, 1986; Alberdi and Bone, 1978). Biozone Mark: 13-3. Fauna: Occitanomys
adroveri, Stephanomys ramblensis ramblensis, Apodemus gorafensis, Paraethomys cf. miocaenicus, Cricetide
microtide cf. Pannonicola, Prolagus cf. michauxi.
Botardo-B (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 13-3. Fauna: Occitanomys adroveri.
Curva (Ruiz Bustos et al., 1992b). Biozone Mark: 13-4. Fauna: Stephanomys ramblensis postremus, Apodemus
gorafensis, Paraethomys cf. meini, Cricetus cf. kormosi, Ruscinomys cf. lasallei, Eliomys truci, Dibolia dekkersi,
Prolagus cf. michauxi.
Aguila-1 (Padial, 1986). Biozone Mark: 13-4. Fauna: Stephanomys ramblensis postremus,Apodemus gorafensis,
Paraethomys cf. meini, Cricetus cf. kormosi, Ruscinomys cf. lasallei.
Aguila-2 (Padial, 1986). Biozone Mark: 13-4. Fauna: Occitanomys adroveri, Stephanomys ramblensis
postremus, Paraethomys cf. meini, Ruscinomys cf. lasallei, Chalicomys jaegeri.
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Nivar (Ruiz Bustos et al., 1992b). Biozone Mark: 13-5. Fauna: Stephanomys ramblensis postremus, Apodemus
gorafensis, Paraethomys cf. meini, Cricetus cf. kormosi, Ruscinomys cf. lasallei, Eliomys truci, Dibolia dekkersi,
Prolagus cf. michauxi.
Purcal (Ruiz Bustos et al., 1992b). Biozone Mark: 13-5. Fauna: Stephanomys ramblensis postremus,
Apodemus gorafensis, Paraethomys cf. meini, Cricetus cf. kormosi, Ruscinomys cf. lasallei, Eliomys truci,
Atlantoxerus cf. adroveri, Muscardinus sp., Dibolia dekkersi, Prolagus cf. michauxi.
Calicasas (Dabrio et al., 1978; Padial, 1986; Ruiz Bustos et al., 1992b). Biozone Mark: 13-5. Fauna:
Stephanomys ramblensis postremus, Apodemus gorafensis, Paraethomys cf. meini, Ruscinomys cf. lasallei, Castor
sp., Prolagus cf. michauxi.
Colorado-1 (Guerra Merchan et al., 1991). Biozone Mark: 13-5. Fauna: Stephanomys ramblensis postremus,
Apodemus gorafensis, Paraethomys cf. meini, Eliomys truci, Prolagus cf. michauxi.
Cacin-1 (Ruiz Bustos et al 1992a). Biozone Mark: 13-5. Fauna: Stephanomys ramblensis postremus,
Apodemus gorafensis, Dibolia dekkersi, Prolagus cf. michauxi, Sorex sp., Lacertidae indet.
Cerro limones (Estevez et al., 1982). Biozone Mark: 13/14. Fauna: Hipparion gromovae granatense.
Abla (Cuevas et al., 1984). Biozone Mark: 13/14. Fauna: Hipparion gromovae granatense.
Botardo-C (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 13/14. Fauna: Apodemus gorafensis,
Apodemus gudrunae, Stephanomys donnezani cordii, Paraethomys cf. meini, Occitanomys cf. adroveri, Castillomys
crusafonti gracilis, Cricetus cf. kormosi, Protatera sp., Eliomys intermedius, Galerix sp., Dibolia dekkersi, Sorex
sp.
Botardo-2 (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 13/14. Fauna: Apodemus gudurnae,
Paraethomys cf. meini, Sthepanomys donnezani cordii, Castillomys crusafonti gracilis.
Botardo-3 (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 13/14. Fauna: Occitanomys cf.
adroveri, Pareathomys cf. meini, Sthephanomys donnezani cordii, Apodemus gudrunae, Galerix sp..
SITES OF THE LOWER. RUSCINIAN
Cuzo-1 (Guerra Merchan y Ruiz Bustos, 1991). Biozone Mark: 14-1. Fauna: Apodemus gudrunae, Paraethomys
cf. meini, Occitanomys cf. brailloni, Cricetus barrieri, Atlantoxerus cf. adroveri, Prolagus cf. michauxi, Episoriculus
gibberodon, Galerix sp.
Bacochas (Alonso Diago, 1989; Sese, 1989). Biozone Mark: 14-1. Fauna: Occitanomys cf. brailloni,
Stephanomys donnezani cordii, Apodemus gudrunae, Paraethomys cf. meini, Myocricetodon sp., Cricetus barrieri,
Eliomys Intermedius, Atlantoxerus cf. adroveri, Prolagus cf. michauxi, Leporidae indet., Soricidae indet., Erinaceidae
indet., Chiroptera indet.
Cuzo-2 (Guerra Merchan y Ruiz Bustos, 1991). Biozone Mark: 14-1. Fauna: Apodemus gudrunae.
Cuzo-3 (Guerra Merchan y Ruiz Bustos, 1991). Biozone Mark: 14-1. Fauna: Apodemus gudrunae, Apodemus
gorafensis, Paraethomys cf. meini, Occitanomys cf. brailloni, Cricetus barrieri, Eliomys truci, Prolagus cf.
michauxi, Galerix sp.
Gorafe-A (Ruiz Bustos et al., 1984). Biozone Mark: 14-2. Fauna: Stephanomys donnezani cordii, Occitanomys
cf. brailloni, Apodemus gorafensis, Paraethomys cf. meini, Ruscinomys lasallei, Trilophomys vandeweerdi, Cricetus
barrieri, Prolagus cf. michauxi.
Gorafe-1 (Bruijn, 1974). Biozone Mark: 14-2. Fauna: Stephanomys donnezani cordii, Paraethomys cf.
meini, Protatera sp., Trilophomys pyrenaeicus, Castillomys crusafonti gracilis.
Colorado-2 (Guerra Merchan et al., 1991). Biozone Mark: 14-2. Fauna: Occitanomys cf. brailloni, Apodemus
gorafensis, Paraethomys cf. meini, Ruscinomys lasallei, Cricetus barrieri, Prolagus cf. michauxi, Erinaceidae
indet., Diplocynodon sp., Emydidae indet., Ciprinidae indet.
Aljibe-2 (Guerra Merchan et al., 1991). Biozone Mark: 14-3. Fauna: Paraethomys cf. meini, Paraethomys
jaegeri, Cricetus barrieri, Eliomys intermedius, Prolagus cf. michauxi.
Aljibe-3 (Guerra Merchan et al., 1991). Biozone Mark: 14-3. Fauna: Paraethomys cf. meini, Paraethomys
jaegeri, Occitanomys cf. brailloni, Castillomys crusafonti gracilis, Stephanomys donnezani cordii, Micromys sp.,
Cricetus barrieri, Prolagus cf. michauxi.
Yeguas (Soria Mingorance y Ruiz Bustos, 1991). Biozone Mark: 14-4. Fauna: Paraethomys cf. meini,
Occitanomys cf. brailloni, Castillomys crusafonti gracilis, Trilophomys vandeweerdi, Eliomys intermedius, Prolagus
cf. michauxi. Crocidura sp., Erinaceidae indet.
Rambla del Conejo (Alonso Diago, 1989; Sese, 1989). Biozone Mark: 14-4. Fauna: Stephanomys donnezani
cordii, Trilophomys vandeweerdi, Eliomys intermedius, Prolagus cf. michauxi, Erinaceidae indet.
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RUIZ BUSTOS, A.
Gorafe-4 (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 14-4. Fauna: Apodemus gorafensis,
Occitanomys cf. brailloni, Paraethomys cf. meini, Castillomys crusafonti gracilis, Cricetus barrieri, Protatera sp.,
Eliomys intermedius, Trilophomys vanderverdi, Dibolia dekkersi.
Stephanomys donenzani adroveri, Paraethomys jaegeri, Occitanomys cf. brailloni, Castilomys crusafonti gracilis,
Cricetus cf. barrieri, Ruscinomys europeus, Trilophomys vanderverdi, Eliomys intermedius, Dolomys sp., Sorex
sp..
Gorafe-2 (Bruijn, 1974). Biozone Mark: 14/15. Fauna: Stephanomys donnezani adroveri, Paraetomys
jaegeri, Trilophomys vandeweerdi, Eliomys intermedius, Dolomys sp.
SITES OF THE UPPER RUSCINIAN
Huescar-3 (Mazo et al., 1985). Biozone Mark: 15-1. Fauna: Stephanomys donnezani donnezani, Castillomys
crusafonti gracilis, Paraethomys cf. meini, Cricetus barrieri, Blancomys neglectus, Mimomys (Cseria) sthelini,
Prolagus cf. michauxi, Trischizolagus aff. maritsae, Anancus arvernensis.
(Alonso Diago, 1989; Sese, 1989). Biozone Mark: 15-1. Fauna: Stephanomys donnezani donnezani, Castillomys
crusafonti crusafonti, Blancomys neglectus, Eliomys intermedius, Mimomys (Cseria) sthelini, Trischizolagus aff.
maritsae, Erinaceidae indet.
Santa (Guerra Merchan y Ruiz Bustos, 1991). Biozone Mark: 15-1. Fauna: Occitanomys cf. brailloni,
Stephanomys donnezani donnezani, Apodemus dominans, Paraethomys cf. meini, Mimomys (Cseria) sthelini,
Micromys sp., Prolagus cf. michauxi.
Cañada del Castaño-1 (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 15-1. Fauna: Paraethomys
jargeri, Occitanomys brailloni, Castillomys crusafonti crusafonti, Rhagapodemus sp., Stephanomys donnezani
donnezani, Apodemus dominans, Eliomys intermedius, Muscardinus sp., Sorex sp.
Stephanomys donnezani adroveri, Paraethomys jaegeri, Occitanomys brailloni, Castillomys crusafonti crusafonti,
Cricetus barrieri, Ruscinomys europeus, Eliomys intermedius, Mimomys (Cseria) sthelini, Sorex sp., Trilophomys
vanderverdi.
Nuca-1 (Ruiz Bustos 1991a). Biozone Mark: 15-2. Fauna: Occitanomys cf. brailloni, Stephanomys donnezani
donnezani, Castillomys crusafonti crusafonti, Apodemus dominans, Cricetus aff. barrieri,Trilophomys vandeweerdi,
Trilophomys sp., Eliomys intermedius, Mimomys (Cseria) sthelini, Micromys sp., Rhagapodemus sp., Prolagus
calpensis., Talpidae indet.
Cómodo (Guerra Merchan y Ruiz Bustos, 1991). Biozone Mark: 15-2. Fauna: Occitanomys cf. brailloni,
Stephanomys donnezani donnezani, Castillomys crusafonti crusafonti, Apodemus dominans, Mimomys (Cseria)
sthelini, Prolagus calpensis.
Barranco de cariuelas-4,5 (Alonso Diago, 1989; Sese, 1989). Biozone Mark: 15-2. Fauna: Stephanomys
donnezani donnezani, Stephanomys donnezani adroveri, Castillomys crusafonti crusafonti, Apodemus dominans,
Ruscinomys europaeus, Trilophomys vandeweerdi, Eliomys intermedius, Mimomys (Cseria) sthelini, Prolagus
calpensis, Leporidae indet.
Cortijo del Muro (Alonso Diago, 1989; Sese, 1989). Biozone Mark: 15-2. Fauna: Stephanomys donnezani
donnezani, Castillomys crusafonti crusafonti, Trilophomys vandeweerdi, Eliomys intermedius, Prolagus calpensis,
Leporidae indet.
Galera-1 (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 15-2. Fauna: Mimomys (Cseria)
sthelini, Stephanomys donnezani donnezani, Occitanomys cf. brailloni,
Cañada del Castaño-2 (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 15-2. Fauna: Occtanomys
brailloni, Castillomys crusafonti crusafonti, Stehpanomys donenzani, Apodemus dominans, Eliomys intermedius,
Mimomys (Kislangia) cappettai.
Fonelas-1 (Alonso Diago, 1989). Biozone Mark: 15-3. Fauna: Occitanomys cf. brailloni, Castillomys
crusafonti crusafonti, Apodemus dominans, Mimomys (Kislangia) cappettai.
SITES OF THE LEWER VILLAFRANCHIAN
Moreda (Ruiz Bustos, 1986). Biozone Mark: 15/16. Fauna: Occitanomys cf. brailloni, Stephanomys amplius,
Castillomys crusafonti crusafonti, Apodemus dominans, Apodemus jeanteti, Blancomys neglectus, Trilophomys
vandeweerdi, Eliomys intermedius, Eliomys sp., Muscardinus sp., Glis sp., Atlantoxerus cf. Adroveri, Mimomys
166
(Cseria) sthelini, Mimomys (Kislangia) cappettai, Micromys sp., Rhagapodemus sp., Beremendia sp., Episoriculus
gibberodon, Dibolia sp., Soricidae indet., Prolagus calpensis, Cercopithecidae indet., Gazella sp..
TCH-3 (Agustí et al.,1990). Biozone Mark: 15/16. Fauna: Stephanomys amplius, Castillomys crusafonti
crusafonti, Apodemus dominans, Eliomys intermedius, Mimomys (Cseria) sthelini, Mimomys (Kislangia) cappettai.
SITES OF THE UPPER VILLAFRANCHIAN
Huelago (Alonso Diago, 1989; Sese, 1989). Biozone Mark: 17-1. Fauna: Stephanomys laynensis, Castillomys
crusafonti meini, Apodemus dominans, Castor sp., Mimomys (Cseria) pliocaenicus, Mymomys (Villanyia) reidi,
Mimomys (Kislangia) rex, Talpidae indet., Soricidae indet., Prolagus calpensis, Oryctolagus sp., Mammuthus
meridionalis, Equus stenonis livenzovensis, Stephanorhinus cf. etruscus, Cervidae indet., Croizetoceros ramosus,
Eucladoceros cf. senezensis, Gazella borbonica, Gazellospira torticornis, Hesperidoceras merlae, Leptobos cf.
elatus.
Barranco de cañuelas-1,2,3 (Alonso Diago, 1989; Sese, 1989). Biozone Mark: 17-1. Fauna: Stephanomys
laynensis, Mymomys (Villanyia) reidi, Equus stenonis livenzovensis.
Nuca-2 (Ruiz Bustos 1991a). Biozone Mark: 17-1. Fauna: Mimomys (Villanyia) reidi, Mimomys (Kislangia)
rex, Drepanosorex praearaneus, Prolagus calpensis, Talpidae indet..
Toyo (Soria Mingorance y Ruiz Bustos, 1991). Biozone Mark: 17-2. Fauna: Stephanomys laynensis, Castillomys
crusafonti meini, Apodemus dominans, Micromys sp., Mimomys (Cseria) pliocaenicus, Prolagus calpensis, Oryctolagus
sp., Drepanosorex praearaneus, Sorex sp., Talpidae indet..
Tapia (Alonso Diago, 1989; Sese, 1989). Biozone Mark: 17-2. Fauna: Apodemus dominans, Mimomys
(Cseria) pliocaenicus, Mymomys (Villanyia) reidi, Mimomys (Kislangia) rex, Prolagus calpensis, Equus stenonis
ssp.
Agua-3 (Ruiz Bustos 1991a). Biozone Mark: 17-2. Fauna: Mimomys (Kislangia) rex, Drepanosorex praearaneus,
Prolagus calpensis.
Nuca-3 (Ruiz Bustos 1991a). Biozone Mark: 17-2. Fauna: Mimomys (Villanyia) reidi, Castillomys crusafonti
meini, Talpa cf. europea.
Galera-2 (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 17-2. Fauna: Apodemus dominans,
Castillomys crusafonti meini, Crocidura kornfeldi, Eliomys intermedius, Galemys kormosi, Mimomys (Cseria)
medasensis, Mimomys (Kislangia) rex, Sorex prearaneus, Stephanomys laynensis.
Cortes Baza-3 (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 17-2. Fauna: Apodemus dominans,
Castillomys crusafonti meini, Mimomys (Villanyia) reidi.
Alquería (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 17-2. Fauna: Apodemus dominans,
Castillomys crusafonti meini, Eliomys intermedius, Mimomys (Villanyia) reidi, Galemys kormosi, Sorex sp.
Alfarerias (Ruiz Bustos, 1986). Biozone Mark: 17-3. Fauna: Stephanomys balcellsi, Prolagus calpensis.
Fuentenueva-1 (Alberdi y Ruiz Bustos 1985; Moya Sola et al., 1987). Biozone Mark: 17-3. Fauna:
Castillomys plinensis, Apodemus dominans, Mymomys (Villanyia) reidi, Gazella borbonica, Equus stenonis cf.
vireti.
Cortes de Baza-1 (Peña et al., 1977). Biozone Mark: 17-3. Fauna: Castillomys plinensis, Mimomys
(Cseria) medasensis.
Orce-1 (Gibert et al. 1994). Biozone Mark: 17-3. Fauna: Mimomys (Villanyia) reidi.
Areba (Guerra Merchan y Ruiz Bustos, 1991). Biozone Mark: 17-4. Fauna: Mymomys (Villanyia) savini,
Oryctolagus sp.
Orce-2 (Gibert et al. 1994). Biozone Mark: 17-4. Fauna:
Apodemus cf. jeanteti, Apodemus dominans, Castillomys crusafonti meini, Drepanosorex prearaneus, Eliomys
intermedius, Equus stenonis cf. vireti, Galemys kormosi, Gazellospira torticornis, Leptobos etruscus, Mimomys
(Ceceria) medasensis, Mimomys (Villanya) reidi,
Cementerio Orce-B (Gibert et al. 1994). Biozone Mark: 17-4. Fauna: Apodemus dominans, Castillomys
crusafonti meini, Eliomys intermedius, Equus stenonis ssp., Leptobos etruscus, Mamuthus meridionalis, Mimomys
(Cseria) medasensis, Mimomys (Villanyia) reidi.
Cortijo Don Alfonso (Gibert et al. 1994). Biozone Mark: 17-4. Fauna: Castillomys crusafonti meini,
Equus stenonis, Mimomys (Villanyia) reidi, Mimomys (Cseria) medasensis.
Cortijo Don Diego (Gibert et al. 1994). Biozone Mark: 17-4. Fauna: Mamuthus meridionalis, Mimomys
(Cseria) medasensis, Soergelia minor.
Barranco del Paso (Gibert et al. 1994). Biozone Mark: 17-4. Fauna: Bubalus sp., Equus stenonis,
Mamuthus meridionalis, Mimomys (Cseria) medasensis, Soergelia minor, Stephanorhirnus etruscus.
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RUIZ BUSTOS, A.
Orce-D (Gibert et al. 1994). Biozone Mark: 17-4. Fauna: Apodemus dominans, Equus stenonis vireti,
Gazella borbonica.
Orce-C (Gibert et al. 1994). Biozone Mark: 17-4. Fauna:
Castillomys crusafonti meini, Cervus sp., Eliomys intermedius, Gazella sp.
Barranco de los Conejos-1 (Gibert et al. 1994). Biozone Mark: 17-4. Fauna: Mimomys (Cseria) medasensis.
SITES OF THE LOWER PLEISTOCENE
Plines-1 (Ruiz Bustos 1991b). Biozone Mark: 18-2. Fauna: Castillomys plinensis, Apodemus aff. sylvaticus,
Arvicola deucalion, Talpa cf. europea, Sorex sp., Crocidura sp. Hippopotamus antiquus, Aves indet., Lacertidae
indet., Barbus aff. boragei, Leuciscus pyrenaicus.
Cerro Parejo (Ruiz Bustos et al., 1990). Biozone Mark: 18-2. Fauna: Arvicola deucalion, Hippopotamus
antiquus.
Orce-3 (Gibert et al. 1994). Biozone Mark: 18-2. Fauna: Apodemus sylvaticus, Arvicola deucalion, Castillomys
plinensis, Crocidura kornfeldi, Eliomys intermedius, Episoriculus gibberodon, Galemys pyrenaicus, Mimomys
(Villanyia) savini, Sorex sp.,
Venta Micena (Gibert et al. 1983, Martinez Navarro, 1991, Ruiz Bustos, 1993c). Biozone Mark: 18-3.
Fauna: Castillomys plinensis, Apodemus aff. mystacinus, Eliomys intermedius, Arvicola deucalion, Euphaiomys
azarai, Hystrix major, Prolagus calpensis, Oryctolagus cf. lacosti, Galemys pyrenaicus, Episoriculus gibberodon,
Equus stenonis granatensis, Mammuthus meridionalis, Stephanorhinus etruscus brachicephalus, Cervus acoronatus,
Cervus elaphoides, Praemegaceros verticornis, Bison sp., Praeovibos sp., Soergelia minor, Capra sp., Hippopotamus
antiquus, Ursus etruscus, Canis etruscus, Canis falconeri,Vulpes praeglacialis, Homotherium latidens, Megantereon
sp., Lynx sp., Pachycrocuta brevirostris, Meles sp., Horno sp.
Lachar (Aguirre et al, 1974; Ruiz Bustos 1975). Biozone Mark: 18-3. Fauna: Equus stenonis granatensis,
Mammuthus meridionalis, Stephanorhinus etruscus brachicephalus, Praemegaceros verticornis, Praedama savini,
Leptobos sp. cf. etruscus.
Fuensanta (Aguirre et al, 1974; Ruiz Bustos 1975). Biozone Mark: 18-3. Fauna: Equus stenonis granatensis,
Mammuthus meridionalis.
Fuentenueva 2-3 (Gibert et al. 1994). Biozone Mark: 18-3. Fauna: Equus stenonis granatensis.
Barranco de Leon 1 (Gibert et al. 1994). Biozone Mark: 18-3. Fauna: Euphaiomys azarai.
Barranco de Leon 2-3 (Gibert et al. 1994). Biozone Mark: 18-3. Fauna: Apodemus aff. mystacinus,
Apodemus sylvaticus, Bubalus sp., Canidae indet., Capra sp., Castillomys plinensis, Cervus sp., Eliomys intermedius,
Equus stenonis granatensis, Euphaiomys azarai, Hippopotamus sp., Homotherium latidens, Mamuthus meridionalis,
Prolagus calpensis, Soergelia minor, Bison sp.,
Barranco de los Conejos-2 (Gibert et al. 1994). Biozone Mark: 18-3. Fauna: Arvicola deucalion,
Mimomys (Villanyia) savini, Praeovibos sp.
Cañada de Murcia (Alonso Diago 1989; Sese, 1989). Biozone Mark: 18-4. Fauna: Arvicola deucalion,
Euphaiomys azarai.
Cortijo de las Nieves 18-4 (Ruiz Bustos et al., 1990). Biozone Mark: 18-4. Fauna: Praedama savini.
Loma Quemada 1-2 (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 18-4. Fauna: Apodemus
sylvaticus, Apodemus aff. mystacinus, Arvicola deucalion, Castillomys plinensis, Euphaiomys azarai, Galemys
pyrenaicus, Micromys sp., Mimomys (Villanyia) savini, Sorex sp.
Puerto Lobo 1-4 (Martin Suarez, 1988; Gibert et al. 1994). Biozone Mark: 18-4. Fauna: Arvicola
deucalion, Euphaiomys azarai, Mimomys (Villanyia) savini.
Orce-6 (Gibert et al. 1994). Biozone Mark: 18-4. Fauna: Arvicola deucalion, Euphaiomys azarai, Bovidae
indet.
Orce-7 (Gibert et al. 1994). Biozone Mark: 18-4. Fauna:
Apodemus sylvaticus, Apodemus aff. mystacinus, Arvicola deucalion, Castillomys plinensis, Euphaiomys
azarai, Galemys pyrenaicus, Sorex sp.
Orce-4 (Gibert et al. 1994). Biozone Mark: 18-4. Fauna: Arvicola deucalion, Euphaiomys azarai, Mimomys
(Villanyia) savini.
Orce-5 (Gibert et al. 1994). Biozone Mark: 18-4. Fauna:Castillomys plinensis, Euphaiomys azarai.
Orce-P (Gibert et al. 1994). Biozone Mark: 18-4. Fauna: Euphaiomys azarai.
Atalaya (Ruiz Bustos y Pérez López 1992). Biozone Mark: 18-5. Fauna: Castillomys plinensis, Apodemus
aff. sylvaticus, Eliomys quercinus quercinus, Euphaiomys azarai, Arvicola deucalion, Lepus sp., Crocidura sp,
Capra sp., Vulpes vulpes, Felis sp.. Reptilia indet., Aves indet.
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Huescar-1 (Mazo et al., 1985). Biozone Mark: 18-6. Fauna: Palaeoloxodon antiquus, Hippopotamus
antiquus, Castillomys plinensis, Apodemus aff. mystacinus, Eliomys quercinus quercinus, Mimomys (Villanyia)
savini, Microtus brecciensis brecciensis, Terricola duodecimcostatus primus. Terricola huescarensis huescarensis.
Oryctolagus sp., Leporidae indet., Lepus cf. granatensis, Equus stenonis ssp., Stephanorhinus etruscus brachicephalus.
Cervus elaphoides, Praemegaceros verticornis, Leptobos sp., Ursus sp.. Panthera sp., Enhydrictis cf. ardea.
Caniles (Guerra Merchan y Ruiz Bustos 1992). Biozone Mark: 18-6. Fauna: Mimomvs (Villanyia) savini.
SITES OF THE MIDDLE PLEISTOCENE
Cullar de Baza-1 (Ruiz Bustos, 1976). Biozone Mark: 19-2. Fauna: Apodemus sylvaticus, Allocricetus
bursae, Eliomys quercinus quercinus, Arvicola mosbachensis, Microtus brecciensis mediterraneus,Terricola huescarensis
cullarensis, Lepus cf. granatensis, Sorex sp., Neomys sp., Crocidura sp., Equus süssenbornensis, Equus altidens,
Stephanorhinus etruscus brachicephalus, Mammuthus trogontherii, Cervus acoronatus, Praemegaceros verticornis,
Bison sp., Capra sp., Sus scrofa, Canis etruscus, Vulpes sp., Vulpes praeglacialis, Crocuta sp., Testudo sp., Lacerta
sp., Teleosteos.
Cuarterones (Ruiz Bustos et al., 1990). Biozone Mark: 19-4. Fauna: Arvicola cf. mosbachensis.
Majolicas-D (Ruiz Bustos et al., 1990, Cerdeño, 1990). Biozone Mark: 19-4. Fauna: Magacerini indet.,
Cervus cf. acoronatus, Capreolus sp., Stephanorhinus hemitoechus.
Solana Z.(Martin Penela, 1987, Cerdeño, 1990). Biozone Mark: 19-5. Fauna: Apodemus cf. flavicollis,
Allocricetus bursae, Eliomys quercinus quercinus, Eliomys quercinus lusitanicus, Arvicola cantiana, Microtus
brecciensis mediterraneus, Oryctolagus cf. cuniculus, Lepus capensis, Sorex sp., Crocidura sp., Macaca sylvanus,
Mammuthus trogontherii, Palaeoloxodon antiquus, Equus caballus, Stephanorhinus hemitoechus, Cervus elaphus,
Capreolus capreolus, Dama sp., Bos primigenius, Bison priscus, Hippopotamus sp. cf. amphibius, Sus scrofa,
Canis lupus, Panthera leo spelaea, Lynx cf. pardina, Felis sylvestris.
SITES OF THE UPPER PLEISTOCENE
Daimuz (Aguirre et al., 1974). Biozone Mark: 20-1. Fauna: Mammuthus intermedius.
Cueva del Agua (Lopez Martinez y Ruiz Bustos, 1977, Sevilla, 1988). Biozone Mark: 20-1. Fauna:
Apodemus sylvaticus, Allocricetus bursae, Eliomys quercinus quercinus, Eliomys quercinus lusitanicus, Mymomys
(Villanyia) savini, Phomys sp., Clethrionomys sp., Arvicola cf. sapidus, Microtus brecciensis mediterraneus,
Terricola duodecimcostatus duodecimcostatus, Oryctolagus cuniculus, Lepus capensis, Talpa europea, Sorex sp.,
Neomys sp., Crocidura sp., Rhinolophus euryale, Myotis myotis, Myotis nattereri, Plecotus auritus, Barbastella
bastastellus, Nyctalus lasiopterus, Nyctalus leisleri, Pipistrellus pipistrellus, Miniopterus schereibersi, Equus caballus,
Cervus elaphus, Capreolus capreolus, Ursus sp., Lynx sp.. Lacertidos, Colubrinae, Anguido.
Majolicas-J (Ruiz Bustos et al., 1990). Biozone Mark: 20-1. Fauna: Microtus brecciensis mediterraneus,
Terricola duodecimcostatus duodecimcostatus.
Cueva Horá (Fuentes y Mejide 1975). Biozone Mark: 20-2. Fauna: Apodemus sylvaticus, Eliomys quercinus
lusitanicus, Allocricetus bursae, Microtus arvalis arvalis, Microtus brecciensis cabrerae, Terricola duodecimcostatus
duodecimcostatus, Arvicola cf. sapidus, Pliomys sp., Oryctolagus cuniculus, Lepus capensis, Equus caballus,
Equus hydruntinus, Stephanorhinus hemitoechus, Cervus elaphus, Bos sp., Capra pyrenaica, Canis lupus, Vulpes
vulpes, Lynx pardina, Felis sylvestris.
Serra-1 (Delgado Castilla et al., 1993). Biozone Mark: 20-2. Fauna: Microtus brecciensis cabrerae, Terricola
duodecimcostatus duodecimcostatus, Arvicola cf. sapidus.
Artesa (Ruiz Bustos y Pérez López 1992). Biozone Mark:20-2. Fauna: Terricola duodecimcostatus
duodecimcostatus, Equus caballus, Cervus elaphus, Capra sp., Canis lupus.
Cruz-1 (Pérez López y Ruiz Bustos, 1991). Biozone Mark: 20-2. Fauna: Apodemus aff. flavicollis, Allocricetus
bursae.
Yedras (Ruiz Bustos, 1978, Sevilla, 1988, Cerdeño, 1990). Biozone Mark: 20-2. Fauna: Apodemus flavicollis,
Allocricetus bursae, Eliomys quercinus quercinus, Microtus brecciensis cabrerae, Microtus arvalis arvalis, Microtus
oeconomus ratticeps,Terricola duodecimcostatus duodecimcostatus, Oryctolagus cuniculus, Lepus capensis, Erinaceus
europaeus, Crocidura suaveolens, Myotis myotis, Myotis nattereri, Equus caballus, Stephanorhinus hemitoechus,
Cervus elaphus, Capreolus capreolus, Dama sp., Bison sp., Vulpes vulpes.
Carihuela (Ruiz Bustos y Garcia Sanchez, 1977; Vega Toscano et al., 1989; Sevilla, 1988; Cerdeño, 1990).
Biozone Mark: 20-3, Apodemus cf. flavicollis, Eliomys quercinus lusitanicus, Allocricetus bursae, Microtus arvalis
169
Ruiz BUSTOS, A.
arvalis, Microtus nivalis, Microtus brecciensis cabrerae, Terricola duodecimcostatus duodecimcostatus, Arvicola cf.
sapidus, Pliomys sp., Oryctolagus cuniculus, Lepus capensis, Erinaceus europeus, Crocidura russula, Sorex cf.
araeus, Sorex cf. minutus, Neomys sp., Equus caballus, Equus hydruntinus, Stephanorhinus hemitoechus, Cervus
elaphus, Bos sp., Capra pyrenaica, Ursus sp., Panthera sp., Canis lupus,Vulpes vulpes. Rhinolophus ferrumequinum,
Rhinolophus hipposideros, Rhinolophus euryale, Myotis myotis, Myotis bechsteini, Myotis nattereri, Myotis
emarginatus, Plecotus auritus, Barbastella bastastellus, Miniopterus schereibersi.
Zafarraya (Barroso et al., 1984). Biozone Mark: 20-4. Fauna: Apodemus cf. flavicollis, Eliomys quercinus
lusitanicus, Microtus arvalis arvalis, Microtus brecciensis cabrerae, Terricola duodecimcostatus duodecimcostatus,
Oryctolagus cuniculus, Lepus capensis, Equus caballus, Cervus elaphus, Bos sp., Capra pyrenaica, Sus scrofa,
Ursus sp., Canis lupus.
Pastora (Aguayo de Hoyos et al., in press). Biozone Mark: 20-4. Fauna: Apodemus cf. flavicollis, Eliomys
quercinus quercinus, Microtus arvalis arvalis, Microtus brecciensis cabrerae, Terricola duodecimcostatus
duodecimcostatus, Arvicola sapidus, Crocidura russula, Oryctolagus cuniculus, Cervus elaphus.
HOLOCENE.
Holocene Fauna. Biozone Mark: 0-1. Fauna: Sciurus vulgaris, Eliomys quercinus quercinus, Eliomys
quercinus lusitanicus, Apodemus sylvaticus, Terricola duodecimcostatus duodecimcostatus, Arvicola sapidus, Microtus
nivalis, M. brecciensis cabrerae, Castor fiber, Lepus granatensis, Lepus capensis, Oryctolagus cuniculus, Erinaceus
europaeus, Erinaceus algirus, Talpa caeca, Crocidura russula, Suncus etruscus, Neomys anomalus, Equus caballus,
Equus hydruntinus, Meles meles, Martes foina, Putorius putorius, Mustela ibérica, Lutra lutra, Ursus arctos,
Canis lupus,Vulpes vulpes, Genetta genetta, Herpestes ichneumon, Felis sylvestris, Lynx pardina, Capra pyrenaica,
Capreolus capreolus, Dama dama, Cervus elaphus, Sus scrofa.
PRESENT
Living Fauna Biozona Mark: 0-0. Fauna: Erinaceus algirus, Talpa caeca, Crocidura russula, Suncus etruscus,
Neomys anomalus, Meles meles, Martes foina, Putorius putorius, Mustela iberica, Lutra lutra, Canis lupus,Vulpes
vulpes, Genetta genetta, Herpestes ichneumon, Felis sylvestris, Lynx pardina, Sciurus vulgaris, Eliomys quercinus
quercinus, Eliomys quercinus lusitanicus, Apodemus sylvaticus, Terricola duodecimcostatus duodecimcostatus,
Arvicola sapidus, Microtus nivalis, M. brecciensis cabrerae, Lepus granatensis, Lepus capensis, Oryctolagus cuniculus,
Capra pyrenaica, Capreolus capreolus, Dama dama, Cervus elaphus, Sus scrofa.
APPENDIX II: TENDENCIES ATTRIBUTED TO THE MAMMALS PRESENT IN THE GRANADA, GUADIX AND BAZA BASINS
The tendencies of each taxon, listed in alphabetical order, are:
Allocricetus bursae = 020001000; Anancus arvernensis = 000222010; Apodemus gudrunae = 001221000;

Apodemus gorafensis= 001221000; Apodemus dominans= 001122000; Apodemus jeanteti= 001221000; Apodemus
aff. mystacinus = 001221000; Apodemus sylvaticus = 001122000; Apodemus flavicollis= 002210000; Arvicola
deucalion= 000000220; Arvicola cantiana = 000000220; Arvicola mosbachensis= 000000220; Arvicola sapidus=
000000220; Atlantoxerus cf. adroveri= 000122000; Beremendia sp. = 000112010; Bison sp.= 021110000; Bison
priscus= 021110000; Blancomys neglectus = 000012000; Bos sp.= 010012000; Bos primigenius = 010012000;
Bovidae indet. = 000001000; Bubalus sp = 000000020; Canis etruscus = 121112000; Canis falconeri= 121112000;
Canis lupus = 121112000; Capra sp.= 121111000; Capra pyrenaica= 121111000; Capreolus sp.= 012220000;
Capreolus capreolus = 012220000; Castillomys crusafonti gracilis= 000112000; Castillomys crusafonti crusafonti=
000112000; Castillomys crusafonti meini = 000112000; Castillomys plinensis = 000112000; Castor sp. = 001110210;
Castor fiber 001110210; Cercopithecidae indet.= 000221000; Cervidae indet.= 002220000; Cervus acoronatus=
002220000; Cervus elaphoides= 002220000; Cervus elaphus = 002220000; Cervus sp. = 002220000; Chalicomys
jaegeri = 001110220; Chooerolophodon pentelici = 000222010; Clethrionomys sp.= 002220000; Cricetide microtide
cf. Pannonicola = 000012000; Cricetus cf. kormosi = 010002000; Cricetus barrieri= 010002000; Crocidura
kornfeldi = 000012000; Crocidura russula = 001122000; Crocidura sp. = 001122000; Crocidura suaveolens = 001122000;
Crocuta sp. = 000012010; Croizetoceros ramosus = 002220000; Dama sp. = 002221000; Dama dama= 002221000;
Deinotherium giganteum = 000222010; Dibolia dekkersi = 000111020; Dibolia sp.= 000111020; Dipoides
170
B1OSTRATIGRAPHY OF THE CONTINENTAL DEPOSITS IN TI 1E GRANADA, GUADIX AND BAZA BASINS
problematicus= 001110220; Dolomys sp.= 000012000; Drepanosorex praearaneus= 021100100; Eliomys sp.=
002221000; Eliomys truci = 002221000; Eliomys Intermedius = 002221000; Eliomys quercinus quercinus= 002221000;
Eliomys quercinus lusitanicus= 002221000; Enhydrictis cf. ardea = 002220000; Episoriculus gibberodon= 000110020;
Equus stenonis livenzovensis = 010012000; Equus stenonis ssp.= 010012000; Equus stenonis cf. vireti= 010012000;
Equus stenonis granatensis = 010012000; Equus süssenbornensis = 020011000; Equus altidens = 010012000; Equus
caballus = 020011000; Equus hydruntinus= 000012000; Erinaceidae indet.= 001211000; Erinaceus europaeus=
001211000; Erinaceus algirus = 001121000; Eucladoceros cf. senezensis = 002220000; Euphaiomys azarai= 020001000;
Felis sp. = 002220000; Felis sylvestris = 002220000; Galemys kormosi = 000111020; Galemys pyrenaicus= 000111020;
Galemys sp. = 000111020; Galerix sp. = 001211000; Gazella sp. = 000012000; Gazella borbonica= 000012000;
Gazellospira torticornis = 000012000; Genetta genetta= 002221000; Gis sp. = 002220000; Gomphotherium sp.=
000222010; Herpestes ichneumon = 000022000; Hesperidoceras merlae = 000012000; Hipparion sp. cf. concudense=
000012000; Hipparion gromovae granatense = 000012000; Hippopotamus crusafonti = 000000020; Hippopotamus
antiquus = 000000020; Hippopotamus sp. cf. amphibius= 000000020; Homotherium latidens= 000012000; Hystrix
majar= 000012000; Ictitherium sp.= 000122000; Leporidae indet. = 000122000; Leptobos sp.= 000012000;
Leptobos cf. elatus = 000012000; Leptobos etruscus = 000012000; Lepus sp. = 011122000; Lepus cf. granatensis=
011122000; Lepus granatensis = 011122000; Lepus capensis= 011122001; Lynx sp. = 002221000; Lutra lutra=
000000220; Lynx pardina = 002222000; Macaca sylvanus = 00022100; Machairodus aphanistus= 001221000;
Mammuthus merídionalis = 000122010; Mammuthus trogontherii = 000122010; Mammuthus intermedius=
020110100; Megacerini indet. 002220000; Megantereon sp.= 001221000; Martes foina = 012220100; Metailurus
sp. aff. major= 001221000; Micromys sp. = 000112000; Microstonyx sp.= 002221010; Microtus brecciensis
brecciensis = 021110000; Microtus brecciensis mediterraneus = 021110000; Microtus brecciensis cabrerae = 021110000;
Microtus arvalis arvalis = 021100000; Microtus oeconomus ratticeps= 221000000; Microtus nivalis= 120000000;
Mimomys (Cseria) sthelini = 000012000; Mimomys (Kislangia) cappettai= 000012000; Mimomys (Cseria)
pliocaenicus = 000012000; Mimomys (Kislangia) rex = 000012000; Mimomys (Cseria) medasensis = 000012000;
Mymomys (Villanyia) savini = 000012010; Mimomys (Villanyia) reidi= 000012010; Meles sp. = 002220000; Meles
meles = 002220100; Muscardinus sp.= 002220000; Mustela iberica = 012221000; Myocricetodon sp.= 000012000;
Neomys sp. = 011100200; Neomys anomalus = 011100200; Occitanomys adroveri= 000112000; Occitanomys cf.
brailloni = 000112000; Oryctolagus sp. = 001122000; Oryctolagus cf. lacosti= 001122000; Oryctolagus cuniculus=
001122000; Pachycrocuta brevirostris= 000012010; Palaeoloxodon antiquus = 000122010; Palaeoryx sp. = 000012001;
Panthera leo spelaea = 000122000; Panthera sp. = 002220000; Parabos sp. = 000012000; Paraethomys cf. miocaenicus=
000122000; Paraethomys cf. meini= 000122000; Paraethomys jaegeri= 000122000; Pentalophodon sivalense=
000222010; Terricola duodecimcostatus primus = 010122110; Terricola duodecimcostatus duodecimcostatus=-
010122110; Terricola huescarensis huescarensis = 010122110; Terricola huescarensis cullarensis = 010122110; Pliomys
sp.= 020000000; Praedarna savini = 002220000; Praemegaceros verticornis= 002220000; Praeovibos sp.= 120000000;
Prohoscidea indet.= 000222010; Prolagus crusafonti= 000022000; Prolagus cf. michauxi = 000022000; Prolagus
calpensis = 000022000; Protatera sp. = 000012000; Putorius putorius = 002221000; Rhagapodemus sp. = 000122000;
Rinocerotidae gen. indet. = 000122000; Ruscinomys europeus = 010012000; Ruscinomys lasallei= 000012000;
Ruscinomys schaubi = 000012000; Sciurus vulgaris = 002210000; Soergelia minor= 012220000; Sorex cf. araeus=
021100100; Sorex cf. minutus = 020000100; Sorex prearaneus = 021100100; Sorex sp. = 021100100; Stephanomys
cf. stadii = 000112000; Stephanomys ramblensis ramblensis= 000112000; Stephanomys ramblensis postremus=
000112000; Stephanomys donnezani cordii= 000112000; Stephanomys donnezani adroveri = 000112000; Stephanomys
donnezani donnezani = 000112000; Stephanomys laynensis = 000112000; Stephanomys amplius = 000112000;
Stephanomys balcellsi= 000112000; Stephanorhinus cf. etruscus = 000122000; Stephanorhinus etruscus brachicephalus=
000122000; Stephanorhinus hemitoechus = 000122000; Suncus etruscus = 001112000; Sus scrofa= 012221110;
Talpa caeca = 021110000; Talpa europea = 021110000; Trilophomys vandeweerdi = 000012000; Trilophomys
pyrenaeicus = 000012000; Trischizolagus aff. maritsae = 000122000; Ursus arctos = 012220100; Ursus etruscus=
002220100; Ursus sp.= 002220100; Valerymys juniensis= 000211000; Valerymys turoliensis= 000211000; Vulpes
sp. = 002220000; Vulpes praeglacialis = 122220000; Vulpes vulpes = 012221000; Zygolophodon sp.= 000222010.
171
Ruiz BUSTOS, A.
Figure 1: Location map of the Betic Cordillera.
System of the Guide Table of a basin

Taxa MN-00
Present
Tendencies in the SI-1 SI-2 SI-3 SI-4 ...Biozone Marks
basin
ABCDEFGH 5 3 7 1 ...n2 of Sites
Presence:
00000001 species X 1* 7. * first time
00222000 species Z 2 x 1. . last time
11000020 species Y 1* 1 1. x inferred
1, 2, 7, times
Figure 3: Systgem of the organization of the data obtained in a basin from the mamalian fauna in order to
construct the Guide Table of the basin. For example, Biozone Mark SI-3 contains 7 sites. In all of these, species
X is present, indicated by the number of times "7". Species Z, however, is present at none of the sites, but its
presence can be inferred. This is indicated with the sign "x". Finally, species Y is present in only one of the 7
sites, as indicated by the times number "1".
172
BIOSTRATIGRAPHY OF THE CONTINENTAL DEPOSITS N THE GRANADA, GUADIX AND BAZA BAS[NS
A C P S 8 m Mammal s Betic
g h o e t eBlostratigraphy C o r d i 1 l e r a
e r 1 r a d L1thology
s o aigi
n r e e t Stages Biozon. Biozon oo alluv.sedim.
Ma. s Isse Marks x lake sedim.
(CO2) t rr. = erosion.surf.
H. Versil. 0-0 00
0.01 ' 0-1 00
20-4 00
Tyrrh. upp. MP20 20-3 oo
C1n P P 20-2 00
L L 20-1 00
E E 19-5 x
I I 19-4 x tltddle
S Sici1. S med. MP19 19-3 x Ple1stocene
T T 19-2 x lake
0 0 19-1 oo
C C 18-6 x
C1r.1n E E 18-5 x juer
N N 18-4 x Pleistocene
Calabr. E 1ou. MP18 18-3 x lake
18-2 x
18-1 00
1.6 17/18 00
C2n U 17-4 x
C2r.1n I 17-3 x VIllafrench.
L upp. MN17 17-2 x lake
L 17-1 x
Piacen. A 16/17 00
F 16-4 00
C2An.1n P R lou. MN16 16-3 oo
L A 16-2 00
C2An.2n I N. 16-1 00
3.3 O 15/16 x
C2An.3n C 15-3 x
E R upp. MN15 15-2 x
N U 15-1 x
E and. S 14/15 x Ruscinian
C3n.ln C 14-4 x lake
C3n.2n I lou. MN14 14-3 x
C3n.3n N. 14-3 x
C3n.4n 14-1 x
5.3 13/14 00
U 13-5 x
Messin. E MN13 13-4 x Ventian
C3An.1n N 13-3 x lake
T 13-2 x
C3An.2n M IAN 13-1 x
6.6 I 12/13 x
0 12-8 x
C3Bn C T 12-7 =
C3Br.1n E U 12-6 x
C3Br.2n N Torton. R 12-5 x Turolian
E O upp. MN12 12-4 x lake
C4n.ln L 12-3 x
1 12-2 x
C4n.2n A 12-1
8.0 N 12/11
Figure 2: Chronology, biostratigraphy and regional lithological sequence of the continental sedirnents in the Betic
Cordillera (opposite page).
173
Ruiz BUSTOS, A.
TABLE 1:
Table Synthesis of the biostratigraphy and palaeoccology of the Betic basins of Granada,
Guadix and Baza. In the columns of the table, the following explanations are provided for
each Biozone Mark: the values of the Habitat Indices; the climatic conditions expressed in
Mammal Diversity Units (MDU) and Phases (A and B), Stages (Al, A2, 13, 14, Bl, B2 and
B3) and Crisis (a2.1, a2.2 and b3.1), which involve the data of the Synthesis Table.
Biozone Climatic Condit.

Marks Hatitat Indices in MDU Events
ABCDEFGHI C' H' D' W'
0-0 1 9 21 21 25 15 5 3 0 36 64 50
50
0-1 1 10 20 21 24 15 6 3 0 37 63 50 modern
50
20-4 2 12 19 21 24 17 3 2 0 36 64 46
54
20-3 2 15 18 20 22 15 5 3 0 40 60 45 *b3,1
55
20-2 3 15 18 19 23 17 2 2 0 38 62 42 B3
58
20-1 1 13 19 21 24 15 5 3 0 38 62 48
52
19-5 1 10 17 20 26 19 3 4 0 31 69 44
56
19-4 0 9 17 21 27 15 4 7 0 30 70 49
51 B
19-3 B2
19-2 2 15 15 18 23 18 5 5 0 37 63 58 42
19-1
18-6 0 5 16 21 30 21 3 4 0 23 77 57 43
18-5 1 7 19 22 27 17 3 5 0 29 71 52 48
18-4 0 4 19 24 28 15 4 7 0 26 74 46 54 B1
18-3 3 9 16 20 26 19 2 6 0 30 70 57 43
18-2 0 11 11 14 22 22 8 11 0 31 69 56 44
18-1
17/18
17-4 0 9 9 14 32 36 0 0 o 18 82 77 23
17-3 0 7 7 12 31 43 0 0 o 14 86 81 19
17-2 0 11 9 14 24 36 4 2 o 24 76 71 29
17-1 0 5 12 16 28 31 5 3 o 22 78 65 35
16/17 A4
16-4
16-3
16-2
16-1
15/16 o 1 10 21 31 31 1 5 0 12 88 63 37
15-3 o 0 7 15 33 46 0 0 0 7 93 78 22
15-2 o 3 6 15 29 44 2 2 0 10 90 76 24
15-1 o 2 5 19 29 44 0 2 6 94 74 26 A
14/15 o 2 4 16 31 47 0 0 6 94 80 20 A3
14-4 o 2 7 19 30 42 0 0 9 91 74 26
14-3 o 3 5 17 30 46 0 0 7 93 78 22
14-2 o 2 7 18 29 42 0 0 2 9 91 75 25
14-1 o 3 11 23 29 30 1 3 1 14 86 63 37
13/14 o 0 9 18 36 36 0 0 0 9 91 73 27
13-5 o 5 11 19 27 27 5 5 0 21 79 60 40 *a2,2
13-4 o 2 9 18 26 30 7 9 0 18 82 58 42
13-3 o 1 9 18 28 30 4 7 2 15 85 61 39 A2
13-2 o 2 11 20 26 26 9 7 0 22 78 54 46 *a2,1
13-1 o 3 11 18 26 32 5 5 0 18 82 61 39
12/13 o 2 11 19 26 28 6 6 2 19 81 57 43
12-8 o 3 8 20 25 28 8 8 0 19 81 56 44
12-7 o 3 7 21 28 35 3 3 0 14 86 66 34
12-6 o 0 0 14 29 57 0 0 0 100 86 14
12-5 Al
12-4
12-3 0 3 3 24 26 35 3 6 0 9 91 65 35
12-2 0 1 8 26 30 27 1 7 0 10 90 58 42
12-1
12/11
174

RUIZ BUSTOS 1999 - Biostratigraphy of The Continental Deposits in The Granada, Guadix and Baza Basins (Betic Cordillera)

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RUIZ BUSTOS 1999 - Biostratigraphy of The Continental Deposits in The Granada, Guadix and Baza Basins (Betic Cordillera)

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Biostratigraphy of the Continental Deposits in the Granada, Guadix

and Baza Basins (Betic Cordillera)

Antonio Ruiz BUSTOS

2. STRATIGRAPHY OF THE GRANADA, GUADIX AND BAZA BASINS

3. BIOZONE MARK (BM)

4. MAMMAL FAUNA AND THEIR EVOLUT IONARY PROCESSES

5. PALAEOECOLOGY FOSTERING MAMMALIAN FAUNA

APPENDIX I: CATALOGUE OF SITES IN THE GRANADA, GUADIX AND BAZA BASINS

SITES OF THE UPPER TUROLIAN

SITES OF THE LOWER. RUSCINIAN

SITES OF THE UPPER RUSCINIAN

SITES OF THE LEWER VILLAFRANCHIAN

SITES OF THE UPPER VILLAFRANCHIAN

SITES OF THE LOWER PLEISTOCENE

SITES OF THE MIDDLE PLEISTOCENE

SITES OF THE UPPER PLEISTOCENE

The tendencies of each taxon, listed in alphabetical order, are:

Allocricetus bursae = 020001000; Anancus arvernensis = 000222010; Apodemus gudrunae = 001221000;

Figure 1: Location map of the Betic Cordillera.

System of the Guide Table of a basin

Biozone Climatic Condit.

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