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Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71 – 88

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Estimating Mediterranean Oligocene–Miocene sea-surface

temperatures: An approach based on coral taxonomic richness
Francesca R. Bosellini a,⁎, Christine Perrin b
a
Dipartimento di Scienze della Terra, Università di Modena e Reggio Emilia, largo S.Eufemia 19, 41100 Modena, Italy
b
Laboratoire des Mécanismes et Transferts en Géologie, Université Paul Sabatier, 14 avenue Edouard Belin, 31400 Toulouse & Muséum National
d'Histoire Naturelle, Département Histoire de la Terre, 8 rue Buffon, 75005 Paris, France
Received 2 July 2007; received in revised form 15 October 2007; accepted 22 October 2007

Abstract

During the Oligocene and Miocene, shallow-water carbonates of the Mediterranean region were rich in scleractinian corals
thriving within various depositional settings, including different reef types. Their diversity patterns, although related to a complex
interplay between a suite of environmental factors and palaeobiogeography, are considered to be strongly controlled by climate
variability and changes in sea-surface water temperature.
By using the quantitative relationship between present-day coral taxonomic richness and prevailing sea-water temperature,
underlined by the so-called “energy hypothesis”, we test zooxanthellate-coral generic richness values from a selection of 102
Oligocene–Miocene localities of the Mediterranean region as a proxy for relative palaeotemperatures.
For each Oligocene–Miocene stage, generic richness values per z-coral site are firstly examined, together with variations of the
Mediterranean z-coral generic pool. For better testing the method and assessing its potential application, patterns of generic richness
and inferred palaeotemperatures are then compared with global palaeoclimatic curves based on marine oxygen stable isotopes data or
other climate proxies, such as palaeoclimatic records from European continental floras and from fossil coral linear extension rate.
Results clearly show that fluctuations of coral richness-derived palaeotemperatures correspond relatively well with global
changes of sea-water temperature especially for the entire Oligocene, the Chattian–Aquitanian boundary and the Late Miocene.
The well known Mid-Miocene Climatic Optimum, however, is not recorded, suggesting that regional factors, acting together with
important palaeogeographical changes, exerted a strong control on the generic richness of Mediterranean z-coral communities.
A remarkable decline of taxonomic richness is recorded after the Burdigalian, together with a gradual decrease of palaeotemperatures
in the region. From the Middle Miocene onwards to the Messinian, however, an increase in the temperature range of z-coral localities is
clearly visible, indicating that z-coral communities were able to thrive and adapt to a wider temperature range, as the Mediterranean was
gradually migrating northwards, outside the tropical belt.
The “energy hypothesis”, if used at global or regional scale, can be considered a promising and reliable method for estimating
Cenozoic palaeotemperatures, from coral or other suitable fossil assemblages of shallow-water carbonates.
© 2007 Elsevier B.V. All rights reserved.

Keywords: Palaeotemperatures; Reef corals; Diversity; Oligocene; Miocene; Mediterranean

⁎ Corresponding author. Fax: +39 59 2055885.

E-mail addresses: frabos@unimore.it (F.R. Bosellini), christine.perrin@lmtg.obs-mip.fr (C. Perrin).

0031-0182/$ - see front matter © 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2007.10.028
72 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88
1. Introduction
Oligocene and Miocene shallow-water carbonates of
the Mediterranean region are well known to bear abundant
and diverse scleractinian corals that occurred isolated, or
formed small bioconstructions to large and luxuriant
framework reefs in a wide variety of structural and
depositional settings (Franseen et al., 1996; Perrin, 2002).
Their evolution and diversity patterns are certainly a
response to a complex palaeogeographic and climatic
history (including seasonality, Chevalier, 1977) with
stepwise transition towards cooler climates, biotic turn-
overs and the almost complete disappearance of zoox-
anthellate corals near the end of the Miocene (Chevalier,
1962; Esteban, 1996; Perrin, 2002).
Within this complex interplay of environmental con-
trols, those related to climate variability and changes in sea-
surface water temperature are undoubtedly of primary
importance. Fossil scleractinian corals are in fact widely
used for indicating past climatic conditions by simply
using an uniformitarian approach or by the use of database
that correlate patterns of coral taxonomic richness (Veron,
1995; Wilson and Rosen, 1998; Rosen, 1999; Bosellini and
Russo, 2000) with palaeoclimate. Assessment of ancient
reef ecosystems through comparative analysis of large
datasets also explains the latitudinal distribution of reef
biota, including corals, by linking it more or less directly to
palaeoclimate and, more precisely, to the attenuation of
sea-surface temperature with increasing latitude (Adams
et al., 1990; Kiessling et al., 1999; Kiessling, 2001).
In this picture, and within an increasing need of data for
better understanding the response of reef corals to climate
change, the Mediterranean region is a suitable and
promising “laboratory” for investigating spatio-temporal
changes in reef ecosystems and coral assemblages during
the Oligocene–Miocene climatic transition. Our detailed
REEFCORAL database provides actually information on
more than 2630 scleractinian-coral spatio-temporal occur-
rences from 285 localities, including the Mediterranean
Sea, Paratethys, Middle East and some European Atlantic
regions.
In this paper we use our database to test zooxanthel-
late-coral generic richness values from a selection of 102
Oligocene–Miocene localities from the Mediterranean
and adjacent areas as a proxy for relative palaeotem-
peratures. The approach is based on the quantitative
relationship that positively correlates taxonomic richness
and prevailing mean sea-surface temperature (SST)
underlined by the so-called “energy hypothesis” (Fraser
and Currie, 1996; Rosen, 1999).
After illustrating the method and dataset, and also
discussing biases and limitations, potentiality of this
approach is assessed by comparing patterns of generic
richness and inferred palaeotemperatures for each
Oligocene–Miocene stage with global palaeoclimatic
curves based on oxygen stable isotope data or other
available climate proxies. From this comparison, we test
our approach as a tool to gather palaeoclimatic in-
formation from shallow-water carbonates, additionally
or alternatively to the more conventional geochemical
technique of stable isotope palaeothermometry that
usually refers to the deep sea record.
2. Past climate from corals
Modern zooxanthellate scleractinian corals strictly
depend on a suite of limiting factors that control the
development of warm and shallow-water marine ecosys-
tems. They also show a distinct latitudinal boundary and
taxonomic richness that correlate well with sea-surface
temperatures (Wells, 1955; Stehli and Wells, 1971; Rosen,
1971, 1984; Veron and Minchin, 1992; Veron, 1995;
Chen, 1999), due to the general attenuation of taxonomic
richness with increasing latitude and decreasing tempera-
ture. By these assumptions, diversity patterns and
latitudinal distribution of z-corals in the geological record
have been traditionally used to infer past climates and
environmental conditions.
Latitudinal distribution of dominant reef biota, includ-
ing corals (Adams et al., 1990; Kiessling et al., 1999;
Kiessling, 2001), and patterns of coral taxonomic richness
(Veron, 1995; Wilson and Rosen, 1998) have been
analyzed through database in order to assess their link
with palaeoclimate. Major changes in diversity patterns
and turnovers, that punctuated the evolutionary history of
Cenozoic Caribbean reef corals, especially during the
Oligocene–Miocene and Pliocene–Pleistocene transi-
tions, have been documented through detailed speci-
men-based dataset at both species and generic level (for a
review see Budd, 2000). Climatic fluctuations related to
cooling episodes, acting with changes in oceanic circula-
tion, increased upwelling and associated turbidity, have
been inferred as the main controlling factors (Edinger and
Risk, 1994; Budd, 2000). Ultimately, biogeographic
factors linked to the emergence of the Central American
isthmus, also played a crucial role (Budd, 2000).
Concerning the Mediterranean region in particular,
Chevalier (1962) was the first to use the observed
reduction of coral richness for qualitatively supporting
cooler conditions through the Miocene. Rosen (1999) and
Bosellini and Russo (2000), respectively for the Miocene
and for the Italian Paleogene, estimated minimum sea-
water palaeotemperatures by using the positive correlation
between taxonomic richness of z-corals and prevailing
 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88
mean sea-water temperatures, the later being directly
related to incident solar energy, as underlined in the so-
called “energy hypothesis” (Fraser and Currie, 1996).
For the very recent past (Holocene), corals are un-
doubtedly very good high-resolution palaeoclimatic tra-
cers. Geochemical analyses of coral skeletons are in fact
used as proxy records for seasonal-to-centennial time-scale,
providing a better understanding of the recent temperature
history of the tropical oceans and of major climate
oscillations such as El Niño Southern Oscillation (ENSO)
(for a review see Grottoli, 2001). Reconstructions of sea-
water temperature, salinity, light levels, upwelling, nutrient
composition and other environmental parameters have
been obtained from coral records by measuring changes in
the skeletal isotope ratios (especially δ18O, δ13C), trace and
minor elements (Sr/Ca, Mg/Ca) and growth rate (Druffel,
1997; Grottoli, 2001).
Concerning coral growth rate, width, density and
chemical composition of each growth band are generally
thought to reflect the average environmental conditions
that prevailed during the time over which that portion of
the skeleton was calcified and, therefore, are looked as
high-resolution archives of past climates (Knutson et al.,
1972; Hudson et al., 1981; Felis and Pätzold, 2004).
These techniques, however, are usually limited to the
most recent geological past (in general not older than
Pleistocene), due to diagenetic processes that often
modify the biological aragonite of the coral skeleton.
Notwithstanding these limitations, ghost structures of
annual growth bands recognized in Late Miocene
massive Porites of Crete (Greece) have been recently
used as proxy records (Brachert et al., 2006a,b). These
relicts of growth bands allowed the application of the
quantitative relationship between SST and annual
extension rates of massive Porites observed from
various Indo-Pacific locations and known as “Porites
thermometer” (Lough and Barnes, 2000), and thus,
reconstruction of minimum SST (Brachert et al., 2006a).
Also, the stable isotope record (δ18O, δ13C) of Porites
skeleton still preserved in aragonite may document
seasonal and interannual climate variability (Brachert
et al., 2006b). These results are compared below (see
“The palaeoclimatic perspective of reef-coral taxonomic
richness”) with “our” palaeotemperatures inferred from
z-coral taxonomic richness.
3. Methods and dataset
3.1. The “energy hypothesis” method
Various studies have quantitatively shown that reef-
coral species richness increases with increasing sea-surface
73
temperatures (Wells, 1955; Stehli and Wells, 1971; Rosen,
1971, 1984; Scheer, 1984; Veron and Minchin, 1992;
Veron, 1995; Chen, 1999), thus leading to consider the
later factor as an important control on coral diversity.
This is consistent with the so-called “richness-energy
hypothesis” originally applied for understanding terres-
trial vegetation patterns, for which richness is a function
of temperature and evapo-transpiration rates, and thus
basically of energy supply (Wright, 1983; Wright et al.,
1993). The “energy hypothesis” has been subsequently
used by Fraser and Currie (1996) to explain modern
global patterns of diversity in z-corals. These authors
empirically related first order global patterns of z-coral
taxonomic richness to incident solar energy for which,
in marine environments, prevailing sea-water tempera-
ture can be considered as a good proxy (Fig. 1).
This rationale has been inverted by Rosen (1999) in
order to deduce palaeotemperatures from richness
values of a given coral fauna at a given place in the
fossil record. In particular, this author tested prelimina-
rily this approach by using richness data from 15
Miocene localities of the Mediterranean region.
The method (for a detailed explanation see Rosen,
1999) consists first of taking the square root of generic
z-coral richness from each locality. Secondly, using the
Fig. 1. The variation in the square root of modern coral generic
richness in 130 sites worldwide, expressed as a function of mean
annual ocean surface temperature (°C) at each site. The original plot of
Fraser and Currie (1996) has been modified by Rosen (1999), with the
integration of the envelope curve of maximum richness values (from
Rosen, 1999).
74 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88

Table 1
List of the 102 selected Oligocene–Miocene localities of the Mediterranean region
Locality Age Gen Gen T° References
(Z) (Z + NZ) (Z)
1 Mallorca, Balearic Islands (Spain) Lower Rupelian 14 15 19,20 Alvarez et al. (1989); Ramos-Guerrero et al.
(1989)
2 Marosticano, Vicentin (N Italy) Lower Rupelian 11 11 18,90 Pfister (1980a)
3 Gornji Grad (Slovenia) Lower-middle Rupelian 15 18 19,30 Nebelsick et al. (2000); Bosellini pers. data
4 Eastern Lessini, Vicentin (N Italy) Middle Rupelian 23 23 19,80 Frost (1981)
5 Marosticano, Vicentin (N Italy) Middle Rupelian 17 18 19,45 Frost (1981)
6 Eastern Lessini, Vicentin (N Italy) Middle Rupelian 16 16 19,40 Bosellini (1988)
7 Ovada, Piedmont (N Italy) Middle Rupelian 13 14 19,15 Fantini Sestini (1960)
8 Marosticano, Vicentin (N Italy) Middle Rupelian 20 20 19,60 Pfister (1980a)
9 Sassello, Liguria (N Italy) Upper Rupelian 18 18 19,50 Fravega et al. (1987)
10 Aqui, Piedmont (N Italy) Upper Rupelian 12 12 19,00 Pfister (1980b)
11 Cairo Montenotte, Liguria (N Italy) Upper Rupelian 22 22 19,75 Pfister (1985)
12 Mesohellenic Basin (Greece) Upper Rupelian 22 24 19,75 Schuster (2002c)
13 Abadeh (central Iran) Upper Rupelian 31 36 20,20 Schuster (2002a); Schuster and
Wielandt (1999)
14 Gaas, Aquitaine (France) Rupelian 11 13 18,90 Chevalier (1956)
15 Sirt Basin (Lybia) Rupelian 21 22 19,70 Hladil et al. (1991)
16 Salento Peninsula, Apulia (S Italy) Lower Chattian 26 27 19,95 Bosellini and Russo (1992); Bosellini and Perrin
(1994); Bosellini pers. data
17 Malte Chattian 21 21 19,70 Chaix and Saint-Martin (1994)
18 Landes, Aquitaine (France) Chattian 11 17 18,90 Chaix and Cahuzac (2001)
19 Dax, Aquitaine (France) Chattian 16 21 19,40 Chevalier (1962)
20 Mesohellenic Basin (Greece) Chattian 15 15 19,30 Schuster (2002b,c)
21 Peyrehorade, Aquitaine (France) Chattian 24 30 19,90 Cahuzac and Chaix (1994)
22 Sirt Basin (Lybia) Chattian 9 9 18,70 Hladil et al. (1991)
23 Aquitaine region (France) Chattian 45 73 21,70 Cahuzac and Chaix (1993,1996)
24 La Nerthe, Provence (France) Upper Chattian 29 32 20,10 Chevalier (1962)
25 Aquitaine region (France) Aquitanian 28 37 20,10 Cahuzac and Chaix (1993, 1996)
26 Gironde, Aquitaine (France) Aquitanian 15 18 19,30 Chevalier (1962)
27 Qom (central Iran) Aquitanian 2 2 16,50 Schuster (2002d); Schuster and
Wielandt (1999)
28 Dolianova, Sardinia (Italy) Aquitanian 6 6 18,00 Governato (1993, unpublished data);
Cherchi et al. (2000)
29 Makran Mountains (S Iran) Aquitanian 8 8 18,55 McCall et al. (1994)
30 Sirt Basin (Lybia) Aquitanian–Serravallian 16 17 19,40 Hladil et al. (1991)
31 Esfahan (central Iran) Aquitanian–Burdigalian 8 8 18,55 Schuster (2002d); Schuster and Wielandt (1999)
32 Paros Island (Greece) Aquitanian–Burdigalian 13 13 19,15 Collection MNHN
33 Cyprus Aquitanian–Burdigalian 15 15 19,30 Follows (1992); Follows et al. (1996)
34 Aquitaine region (France) Aquitanian–Burdigalian 16 18 19,40 Oosterban (1988)
35 NW Gulf of Suez (Egypt) Burdigalian 12 15 19,00 Schuster (2002b,e)
36 Qom (central Iran) Burdigalian 16 17 19,40 Schuster (2002d); Schuster and Wielandt (1999)
37 Zagros Mountains (SW Iran) Burdigalian 7 7 18,30 Schuster (2002d); Schuster and Wielandt (1999)
38 Makran Mountains (S Iran) Burdigalian 32 38 20,25 McCall et al. (1994)
39 Aquitaine region (France) Burdigalian 33 45 20,30 Cahuzac and Chaix (1993, 1996)
40 Gironde, Aquitaine (France) Burdigalian 22 24 19,75 Chevalier (1962)
41 Dax, Aquitaine (France) Burdigalian 24 27 19,90 Chevalier (1962)
42 Torino Hills, Piedmont (N Italy) Burdigalian–Langhian 35 76 20,40 Chevalier (1962); Zunino (2007, unpublished
data)
43 Valencia region (Spain) Langhian 5 5 17,80 Calvet et al. (1994)
44 Aquitaine region (France) Langhian 18 27 19,50 Cahuzac and Chaix (1993, 1996)
45 Manciet, Aquitaine (France) Langhian 4 6 17,50 Chevalier (1962)
46 Saubrigues, Aquitaine (France) Langhian 4 5 17,50 Chevalier (1962)
47 Styrian Basin (Austria) Lower Badenian (Langhian) 3 3 17,00 Friebe (1991)
48 Budapest area (Hungary) Lower Badenian (Langhian) 10 10 18,80 Oosterban (1990)
49 Murchas, Granada Basin (S Spain) Late Langhian 3 3 17,00 Braga et al. (1996)
50 Duplek (Slovenia) Badenian (Langhian–L.Serrav.) 4 4 17,50 Baron Szabo (1997)
51 Budapest area (Hungary) Badenian (Langhian–L.Serrav.) 4 4 17,50 Oosterban (1990)
 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88 75

Table 1 (continued )
Locality Age Gen Gen T° References
(Z) (Z + NZ) (Z)
52 Vienna Basin (Austria) Badenian (Langhian–L.Serrav.) 11 19 18,90 Piller and Kleemann (1991); Riegl and Piller
(2000)
53 Holy Cross Mountains (Poland) Badenian (Langhian–L.Serrav.) 7 18 18,30 Roniewicz and Stolarski (1991)
54 North Hurghada (Egypt) Langhian–L.Serravallian 14 17 19,20 Perrin et al. (1998)
55 South Quseir area (Egypt) Langhian–L.Serravallian 10 10 18,80 Perrin et al. (1998)
56 Hérault, Languedoc (France) Langhian–Serravallian 10 12 18,80 Chevalier (1962)
57 Loire Basin (France) Langhian–Serravallian 10 16 18,80 Chevalier (1962)
58 Djebel Chott, Dahra region Langhian–Serravallian 2 3 16,50 Belkebir et al. (1994)
(Algeria)
59 Aquitaine region (France) Serravallian 5 13 17,80 Cahuzac and Chaix (1993, 1996)
60 Midyan region (Saudi Arabia) Tortonian 3 3 17,00 Dullo et al. (1983)
61 Cyprus Tortonian 1 1 15,90 Follows (1992); Follows et al. (1996)
62 Heraklion Basin, Crete (Greece) Tortonian 5 7 17,80 Buchbinder (1996)
63 Granada Basin (S Spain) Tortonian 4 4 17,50 Braga et al. (1990)
64 Almanzora River Valley, Almeria Tortonian 1 1 15,90 Martin et al. (1989)
(SE Spain)
65 Kasaba area (Turkey) Tortonian 1 1 15,90 Hayward (1982); Hayward et al. (1996)
66 NW Crete (Greece) Tortonian 3 3 17,00 Baron Szabo (1995)
67 Modena area (N Italy) Tortonian 7 16 18,30 Montanaro (1929)
68 Modena area (N Italy) Tortonian 6 19 18,00 Chevalier (1962); Chevalier collection, MNHN
69 Tortona area, Piedmont (N Italy) Tortonian 3 17 17,00 Chevalier (1962)
70 Sangonera, Murcia (SE Spain) Upper Tortonian 2 2 16,50 Lopez Buendia (1992)
71 Caltanisetta region, Sicily (S Italy) Upper Tortonian 5 5 17,80 Catalano (1979)
72 Isparta region (Turkey) Tortonian–Messinian 5 5 17,80 Karabiyikoglu et al. (1999)
73 Mallorca, Balearic Islands (Spain) Tortonian–Messinian 3 3 17,00 Pomar (1991)
74 Menorca, Balearic Islands (Spain) Tortonian–Messinian 2 2 16,50 Obrador et al. (1992)
75 Gozo Island Tortonian–Messinian 1 1 15,90 Pedley (1979)
76 Sirt Basin (Lybia) Tortonian–Messinian 3 3 17,00 Hladil et al. (1991)
77 Calabria (S Italy) Tortonian–Messinian 4 8 17,50 Chevalier (1962)
78 Vigoleno, Piacenza (N Italy) Lower Messinian 2 2 16,60 Barrier et al. (1994)
79 Maiella Mountain (Central Italy) Lower Messinian 1 1 15,90 Danese (1999)
80 Cabo de Gata, Almeria (Spain) Lower Messinian 2 2 16,50 Esteban and Giner (1980)
81 Rosignano, Tuscany (Italy) Lower Messinian 5 5 17,80 Bossio et al. (1996); Chevalier (1962)
82 Vibo Valentia, Calabria (S Italy) Lower Messinian 3 3 17,00 Pedley and Grasso (1994); Romano et al. (2005)
83 Salento Peninsula, Apulia (S Italy) Lower Messinian 3 3 17,00 Bosellini et al. (2001, 2002)
84 Caltanisetta region, Sicily (S Italy) Lower Messinian 2 2 16,50 Grasso and Pedley (1989); Pedley (1996)
85 Melilla (N Morocco) Messinian 2 2 16,50 Saint-Martin and Cornée (1996)
86 San Miguel de Salinas Basin (SE Spain) Messinian 1 1 15,90 Reinhold (1995)
87 Alicante Basin (SE Spain) Messinian 1 1 15,90 Calvet et al. (1996)
88 Orania (Algeria) Messinian 2 2 16,50 Saint-Martin (1996)
89 Sicily (Italy) Messinian 2 2 16,50 Catalano (1979)
90 Malta and Gozo Islands Messinian 2 2 16,50 Bosence and Pedley (1982)
91 Malta Messinian 3 3 17,00 Chaix and Saint-Martin (1994)
92 Bou Meriem (Morocco) Messinian 3 3 17,00 Elhamzaoui and Lachkhem (1994)
93 Cabo de Gata, Almeria (SE Spain) Messinian 1 1 15,90 Esteban and Giner (1980)
94 Fes (Morocco) Messinian 4 4 17,50 Moissette and Saint-Martin (1995)
95 Nijar, Almeria (SE Spain) Messinian 1 1 15,90 Dabrio et al. (1981)
96 Lorca Basin, Murcia (SE Spain) Messinian 4 4 17,50 Rouchy et al. (1986)
97 Orania (N Algeria) Messinian 1 1 15,90 Rouchy et al. (1986)
98 Orania (N Algeria) Messinian 5 5 17,80 Saint-Martin (1996)
99 Melilla (N Morocco) Messinian 2 2 16,50 Saint-Martin and Cornée (1996)
100 Melilla (N Morocco) Messinian 1 1 15,90 Rouchy et al. (1986)
101 Santa Pola, Alicante (SE Spain) Messinian 1 1 15,90 Rouchy et al. (1986)
102 Sorbas Basin, Almeria (SE Spain) Upper Messinian 1 1 15,90 Riding et al. (1991); Braga et al. (2006)

envelope curve of maximum richness values that Rosen each locality. In this way, the application of this meth-
added to the original plot (see Fig. 1), it is possible od to the fossil record is facilitated and allows mini-
to read back the minimum possible temperature for mizing biases caused by the undoubtedly incomplete
76 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88
fossil record, and hence, not representative of original
richness values.
3.2. The coral dataset from Oligocene–Miocene
Mediterranean localities
In this paper, estimated palaeotemperatures using the
method described above, are derived from richness
values of z-corals from Oligocene and Miocene
localities of the Mediterranean region (Table 1).
Richness data have been extracted from our dataset,
and provided by the 102 selected localities ranging in
age from the lower Rupelian to the upper Messinian.
Although some stages appear less represented than
others, because of poor investigations or limited
stratigraphic resolution, the broad geographic distribu-
tion of the selected localities is considered to be
satisfactorily representative of the whole “Mediterra-
nean” region both in time and space (Table 1, Fig. 2).
As highlighted in the Introduction, Mediterranean
Oligocene–Miocene coral reef sites have been in general
well explored providing a huge literature dataset and
contributing to operate a relatively good control on
taxonomy and stratigraphy. It was thus possible, for almost
all selected localities, to refer to quite recent studies dealing
with the systematics and/or palaeoecology of their coral
fauna and associated with up-dated stratigraphy (Table 1).
Data concerning number of genera also integrate some
systematic revisions that we directly made at the generic
level within the database, and unpublished data from our
studies of coral collections, including published (i.e. the
classical original Collection of Chevalier 1962, at the
MNHN, Paris) and unpublished Museum collections,
together with our own collections.
As the “energy hypothesis” has been applied by
Fraser and Currie (1996) to “hermatypic” corals, richness
of z-corals is specifically indicated for each locality. We
include in the category of zooxanthellate corals (z-genera
in Table 1): 1) z-corals sensu Wilson and Rosen (1998,
Tab.2, p.170: genera that are still living and are known to
be zooxanthellate); and 2) corals designated as “z-like”
(i.e. extinct genera having morphology and skeletal
characters typical of living z-corals). In Table 1, the total
number of genera is also provided for comparison, and
includes non-zooxanthellate corals, but also those whose
symbiotic status is particularly uncertain.
4. Potential biases
When using the “energy hypothesis” method, potential
biases interfere directly at two main levels in the data
processing: 1) with the assessment of biodiversity patterns
and, 2) with the relationship between generic richness of
z-coral assemblages and sea-surface temperature.
4.1. Biases acting on the estimates of z-coral richness
through time
Potential biases acting on the estimates of z-coral
richness through time are those influencing the precision
of the stratigraphical range of any z-coral site, the
collection of data, the taxonomic identification, and the
attribution of the symbiotic status of any scleractinian
genus (Fig. 3).
It is well known that the age of coral-bearing facies,
including reefs, is often difficult to constrain due to the
lack of stratigraphical biomarkers within these facies,
especially for the Cenozoic, and to the difficult precise
correlations with contemporaneous layers containing
these biostratigraphical markers, such as basinal facies.
Other dating techniques, such as radiometric age and
grade-datations, although locally used in particular cases
(Magné et al., 1987), are often difficult to apply. Con-
sequently, the stratigraphical resolution of z-coral local-
ities remains highly variable through time and space and,
as far as the Mediterranean Oligocene–Miocene is
concerned, some stages such as Langhian and Serraval-
lian, and Tortonian and Messinian, are often difficult to
distinguish with confidence.
The quality and amount of data collected for any coral
locality (i.e. sampling) may also have a strong impact on
the estimates of taxonomic richness (see for discussion
Perrin et al., 1995). This is directly reflected both by the
number of coral specimens in a collection, and by the
level and amount of investigation in a given area. From
this point of view, it is important to keep in mind the
contrasted levels of investigation, and hence the different
amount of data available, between the eastern and
western Mediterranean. Some countries, such as Greece,
where Oligocene and Miocene carbonate platforms are
known to outcrop, obviously show a lack of data
concerning the distribution of z-corals (Fig. 2). In
addition, sampling itself may also be linked in some
way to coral preservation and diagenesis.
The identification of fossil taxa is always affected by
some potential biases. These are typically represented
by some degree of taxonomic subjectivity and the state
of preservation/diagenesis characterizing the studied
specimen. For scleractinian corals, the poor state of
systematics constitutes an additional limiting factor for
the precise estimates of coral assemblages. The systema-
tics of scleractinian corals is fundamentally based on
microstructural characters, which produce the three-
dimensional microarchitecture and micromorphology of
 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88 77

Fig. 2. The geographic distribution of the 102 Oligocene–Miocene localities of the Mediterranean region. Oligocene, Early Miocene, Middle
Miocene and Late Miocene localities are differentiated.
78 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88
septae. The microstructural/ultrastructural patterns,
which are the direct expression of the biomineralization
process, are less dependent on external abiotic para-
meters of the environment within which the scleractinian
individual grows, than the pure morphological char-
acters, which typically exhibit an important intraspecific
and even intracolonial plasticity. Microstructural char-
acterization of a fossil coral requires the skeleton to be
preserved completely or partially in aragonite for the
spatial arrangement of aragonite “fibres” and “calcifica-
tion centres” to be identified. However, aragonite
preservation is generally prevented in reef settings and
carbonate platforms, because the relatively high macro-
and microporosity of reef material favour circulation of
diagenetic fluids, and hence, leaching or recrystallisation
of the less chemically-stable aragonitic components.
This has two important consequences for estimates of
coral richness: 1) the taxonomic definition of some fossil
genera still lack the basic understanding of their
microstructural pattern, and 2) the potentially high-
diversity coral-bearing facies, such as reefs, commonly
are those offering the worst conditions for the good
preservation of coral skeletons. Additionally, it is well
known that coral taxa have different susceptibility to
taphonomy and diagenesis (Constantz, 1985; Dullo,
1986) depending on the microstructure and the micro-
architecture of their skeleton. These act together with the
growth morphology of the colony, to determine the
preservation potential of the different taxa in fossil reef
assemblages.
Moreover, as stressed by many authors, morphologi-
cal characters used for the distinction of species display a
strong variability, hence introducing a certain amount of
subjectivity in the definition and identification of both
Fig. 3. Estimating the z-coral richness through time: potential biases and their interactions.
living and fossil scleractinian species. Additionally, the
high potential of hybridization between species may
provide intermediate morphologies, thus further
accentuating difficulties in species identification both
in present-day and fossil coral communities (e.g.
Sheppard and Sheppard, 1991; Veron, 1995).
The attribution of a symbiotic status to a scleractinian
genus is directly based on the knowledge of the symbiotic
status of its living counterparts, either belonging to the
same genus or to the same family, if the latter shows a
unique status for all its living species. This implies the
basic assumption that the symbiotic requirement of a
particular genus has remained stable through time, at least
for the time-interval considered in the present study (i.e.
Oligocene-to-present). Hence, the attribution of a sym-
biotic status to any fossil specimen strongly relies on its
taxonomic identification and is, therefore, subject to the
same biases (i.e. taxonomic subjectivity and general state
of coral systematics).
4.2. Biases acting on the relationship between generic
richness of z-coral assemblages and sea-surface
temperature
These include biases acting at two very different
spatial scales: 1) those induced by local environmental
factors other than sea-surface temperature, and 2) those
related to biogeographical features.
The environmental factors, which can locally limit
z-coral taxonomic richness, are mainly turbidity affecting
light penetration in the water column, and local nutrient
inputs adjacent to river mouth or land runoffs, although
the distribution of habitats favourable to coral settlement
and growth also influence to some degree the general
 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88
pattern of coral diversity. One way to minimize these
problems is to analyse a significant number of z-coral
sites, and to consider the pattern obtained at regional or
global scale, rather than to deduce palaeotemperatures for
particular places.
Historical biogeography also exerts a strong control
over the diversity pattern, both at global and regional
scales. In particular, the lack of possible exchange
with z-coral faunas of the Indo-Pacific after the late
Burdigalian, due to the closure of marine seaways
through the Middle East (McCall et al., 1994; Perrin
et al., 1998; Rögl, 1998; Schuster and Wielandt,
1999), is one of the multiple causes of the decreasing
richness of the Mediterranean coral fauna during the
Miocene (Chevalier, 1962; Rosen, 1999; Perrin, 2002).
The importance of the biogeographical context has been
also underlined by Wilson and Rosen (1998). These
authors explain the low diversity of the Indo-West
Pacific Paleogene corals as mainly a result of a strong
tectonic control on the distribution of habitats suitable
for reef-coral fauna. The modern Indo-West Pacific
centre emerged, in fact, only in the Miocene after the
tectonic collision between the cratons of Australia and
mainland Asia (Hall, 1996), becoming richer from the
Neogene until the present time (Veron, 1995; Wilson
and Rosen, 1998; Rosen, 2002).
5. Taxonomic richness and palaeotemperatures:
the results
5.1. Coral diversity patterns
The regional generic pool of z-corals and its variations
during the Oligocene–Miocene time interval was esti-
Fig. 4. Variations of the generic pool of z-corals in the Mediterranean region through time. Estimates are based on 2637 spatio-temporal occurrences
of z-corals, and 285 localities having a well constrained stratigraphical range.
79
mated from 2637 z-coral spatio-temporal occurrences and
285 localities in the Mediterranean region (Fig. 4). This
was done after minor taxonomic revision and by
considering only the localities for which the stratigraphi-
cal range could be relatively well constrained. The total
number of coral genera for the Oligocene–Miocene time
slice is 103, 13 of which having an unknown symbiotic
status. 51 genera are common to Oligocene and Miocene,
while 37 are exclusively known from Oligocene localities,
and 15 were only mentioned in the Miocene. The highest
generic diversity for the whole region is recorded during
the Chattian, and is followed by a marked decrease in the
Aquitanian. The Burdigalian-to-Langhian interval is
characterized by relatively high values of generic
richness. Two strong decreases of the Mediterranean
generic pool are shown between the Langhian and the
Serravallian, and between the Serravallian and the
Tortonian, representing together a drop of more than
50%. It is also important to underline that the generic pool
of z-corals is not so different during the Tortonian and
Messinian time-slices (Fig. 4), although some rare upper
Messinian monogeneric Porites reefs, recorded from the
Sorbas Basin, in southern Spain (Riding et al., 1991;
Braga et al., 2006), underline a further decline of z-coral
generic richness during the Messinian.
The number of z-coral genera for each of the 102
selected localities (Table 1) is reported in Fig. 5A. The
stratigraphical range of each locality is represented in
million years, by the length of the horizontal bar. For each
stage, the generic richness between localities varies
around an average, except for a few localities character-
ized by very different values. Obviously, these strong
differences reflect some biases, either due to local
environmental conditions influencing coral richness at
 80
F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88
Fig. 5. A. Generic richness of z-corals expressed for the 102 Oligocene–Miocene coral localities of the Mediterranean region selected for this study. Numbers refer to the localities number listed in Table 1.
•: for lower Messinian and non-differentiated Messinian localities (no. 78-80-84-85-88-89-90-99) ¤: for lower Messinian and non-differentiated Messinian localities (no. 79-86-87-93-95-97-100-101). ⁎: for
the upper Messinian locality n°102. Bars represent the stratigraphical range of each locality. B. Palaeotemperatures inferred from generic richness at each locality. Time-scale of Gradstein et al. (2004).
 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88
these sites or to artefacts during data collection and
analysis. Therefore, these localities do not have to be
considered when analysing the general trend and fluctua-
tions of diversity through time. The highest anomalous
values in generic richness, which certainly are related to
excessive taxonomic splitting, are recorded at localities
no. 23 (Aquitaine region, France) in the Chattian, no. 25
(Aquitaine region, France) in the Aquitanian and no. 42
(Torino Hills, Piedmont, northern Italy) in the Burdiga-
lian–Langhian, and to a lesser degree by no. 39
(Aquitaine region, France) in the Burdigalian. When
compared to the total generic pool of the Mediterranean
region, the abnormally high generic richness at these sites
corresponds to more than 65% of the contemporaneous
regional pool of z-coral genera being represented at the
same place. The localities no. 13 (Abadeh, Iran) in the
Rupelian and no. 38 (Makran, Iran) in the Burdigalian,
both show relatively high generic richness, which
undoubtedly results from the Indo-Pacific affinity of
their coral fauna. The Abadeh Oligocene coral fauna is a
mixture of Mediterranean and Indo-Pacific elements
(Schuster and Wielandt, 1999), while the Makran
Burdigalian corals are entirely of Indo-Pacific affinity
(McCall et al., 1994). Both these localities represent
particular cases in the general pattern, due to the specific
significance of their coral faunas.
The generic richness per z-coral site slightly increases
between the Rupelian and the Chattian, and possibly
reaches its highest value for the strictly Mediterranean
coral faunas, during the later time interval. At the end of
the Chattian, an important drop is shown, while the
diversity of z-coral localities increases again markedly in
the Burdigalian, paralleling the fluctuations of the
Mediterranean z-coral generic pool. The Burdigalian–
Langhian boundary is marked by a decrease of z-coral
diversity per locality, following the generic richness of
the whole region. After the Langhian, the diversity record
successive decreases both in the number of z-coral genera
per site and in the overall generic coral diversity in the
Mediterranean region.
5.2. The inferred palaeotemperatures
Fig. 5B shows the palaeotemperatures inferred from
the number of z-coral genera at each locality by using
the “energy hypothesis” of Fraser and Currie (1996).
The fluctuations of these palaeotemperatures throughout
the Oligocene–Miocene follow the pattern described
above for the generic coral diversity. It should be note,
however, that the range of temperatures represented by
the coral localities is gradually increasing through time.
This widening of temperature range through time is
81
particularly well marked after the Burdigalian (Fig. 5B)
and cannot be related to a wider palaeolatitudinal range
of z-coral distribution in the area. It probably reflects the
gradual adaptation of the Mediterranean z-coral fauna to
a larger range of sea-surface temperatures as the regional
cooling was progressively occurring. The relatively low
temperatures estimated for the Late Miocene show a
value below the common threshold of 18°C annual
average for coral reef growth. As we are dealing,
however, with z-corals and not with coral reefs, these
values are consistent with present-day z-corals living for
example around Japan (Yabe and Sugiyama, 1932;
Hamada, 1977; Veron and Minchin, 1992).
6. The palaeoclimatic perspective of reef-coral
taxonomic richness
The complex relationship between richness pattern of
z-corals assemblages and sea-surface temperature has
been previously underlined, this link being strongly in-
fluenced by many interactions and feedbacks acting at
different spatio-temporal scales. Diversity patterns, in
fact, result both from ecological and historical causes, and
particular caution is thus necessary for not considering
them just in terms of stress response (Rosen, 1988; Jokiel
and Martinelli, 1992; Perrin et al., 1995). These important
limitations, which commonly affect the fossil record at
different levels of investigation, should be, and can be, in
part minimized before treating the data for palaeoenvir-
onmental and palaeoclimatic reconstructions.
6.1. Minimizing problems and biases
In order to overcome the different types of biases, those
having only a local influence and affecting some particular
localities can be minimized by considering a significant
number of z-coral sites, and by analyzing the pattern
obtained at regional or global scale. As regards biases
related to historical biogeography, that undoubtedly exerts a
strong control on the regional species pool and thus
influence to some degree the inferred temperature-diversity
pattern at regional scale, their effect can be estimated
precisely enough by comparing the results obtained with
independent data, such as global isotopic curves of
palaeotemperatures or other available climate proxies.
It is also important to underline that the “energy
hypothesis” approach basically relies on the assumption
that the relationship between generic richness of z-corals
at a given site and sea-water temperature did not change
considerably through time. Analogously, this prerequisite
has been considered as fundamental by the “nearest-
living-relative” method (NLR) and by the coexistence
82 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88
approach (CA), both commonly applied by palaeobota-
nists for quantitative reconstructions of Cenozoic con-
tinental palaeoclimates (Chaloner and Creber, 1990;
Mosbrugger and Utescher, 1997; Uhl et al., 2003;
Mosbrugger et al., 2005). In this case, biases undoubtedly
introduced by uniformitarianism can be minimized by
using our temperature-generic richness plot to discuss
relative palaeotemperatures through time rather than
absolute values (see also Rosen, 1999, p. 320).
6.2. Testing the method
Palaeotemperatures deduced for each coral locality by
applying the “energy hypothesis” allow a curve showing
the fluctuations of mean temperature to be reconstructed
through Oligocene–Miocene time (Fig. 6). Temporal
variations of diversity are evaluated by averaging the
generic richness of localities occurring in the same stage
(Fig. 6A). In addition, the richness values of the
northernmost localities of each stage were also considered
(Fig. 6B), because the z-coral communities developed at
the northern margin of their global distribution, and hence
close to the limits of their ecological requirements, are
expected to be more sensitive to climate fluctuations. In
case several z-coral sites occur at similar palaeolatitudes
for the same stage, the average of their generic richness
values was taken.
In order to test the reliability and potential application
of our empirical approach, our inferred palaeotempera-
tures are compared to similar results obtained from
independent methods, in particular palaeotemperature
curves derived from marine oxygen stable isotopes
(Dodd and Stanton, 1990; Abreu and Haddad, 1998;
Zachos et al., 2001) and from flora continental record
(Mosbrugger et al., 2005). Fig. 6 clearly shows that
minimum palaeotemperature values inferred from z-coral
richness match, in the general trends, the global
palaeoclimatic curves, demonstrating the consistence
and reliability of our results. Some discrepancies,
however, are shown for the Middle Miocene.
Palaeotemperatures inferred from z-coral richness
show a slight increase from the lower to the middle
Rupelian, followed by a slight decrease in the upper
Rupelian. This trend is also recorded by the isotopic data,
in particular the curves E and G of Fig. 6. In northern
Italy (Liguria, Piedmont and Vicentin Lessini Shelf),
several z-coral localities have a well constrained
stratigraphical range (Bosellini and Russo, 2000), and
for the Lessini Shelf in particular, a gradual increase of its
z-coral taxonomic richness has been documented, and
regarded as driven by change in sea-water temperature
throughout the Rupelian (Bosellini and Russo, 1988).
Our data also parallel the trend of the marine isotopic
curves and of the continental flora curve, which both
show a cooling at the Chattian–Aquitanian boundary.
The average generic richness records a drastic decline
from Chattian to Aquitanian, while this decrease appears
attenuated in the northernmost localities.
The highest diversity values at the northernmost z-
coral site are recorded in the Burdigalian, which also
corresponds to a relative high value of average richness.
These peaks of taxonomic richness are not directly
correlated with a marked increase in sea-water tempera-
ture at the global scale, although the composite curve of
Abreu and Haddad (1998; Fig. 6F) shows a general and
gradual increase in sea temperature from the early
Burdigalian to the Langhian. The two other isotopic
curves (Dodd and Stanton, 1990; Zachos et al., 2001;
Fig. 6E and G, respectively) are based on a compilation
of data at global scale, and both represent the variations
in deep-sea water temperature, for which no warming
trend in the Burdigalian is recorded. Therefore, the
taxonomic richness of Mediterranean z-coral commu-
nities in the Burdigalian is not directly related to the
global sea-surface temperature, but instead, it seems that
other factors arise.
Similarly, the cooling trend in our inferred palaeo-
temperatures, beginning in the Middle Miocene and
increasing in the Late Miocene, differs from the isotopic
temperatures curves, mainly because the Mid-Miocene
Climatic Optimum is not shown by a maximum in the
Mediterranean z-coral diversity. Although it has been
demonstrated that the Mid-Miocene Climatic Optimum
is recorded by shallow-water faunas and facies in the
Mediterranean (e.g. Bojar et al., 2004), it is not clearly
expressed in the generic diversity of z-coral commu-
nities of this age, especially when these are compared to
their Burdigalian counterparts. It is, however, worth to
underline that a similar pattern of increased decline in
taxonomic richness (at the species level) after the
isolation of the Mediterranean basin from the Indian
Ocean, has been reported for the tropical coralline algal
genus Sporolithon by Braga and Bassi (2007).
Two main causes can explain these discrepancies
between our inferred temperature curves and the global
isotopic curves. The first is the stratigraphical resolu-
tion, which is generally not very well constrained for
Mediterranean shallow-water carbonates both in the
Burdigalian and in the Langhian–Serravallian. The
second is linked to the particular palaeogeographical
changes occurring in this region during the Miocene.
Hence, the influence of regional factors at that time may
prevail over global climatic parameters. The occurrence
of a warm peak recorded by the European floras in the
 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88
Fig. 6. Patterns of z-coral diversity and inferred palaeotemperatures through the Oligocene–Miocene of the Mediterranean region, and comparison with global and European palaeotemperature curves
A. Changes in z-coral diversity, estimated by averaging the z-coral generic richness of localities of the same stage. B. Change of the z-coral generic richness based on the northernmost localities.
C. Change of minimum sea-surface palaeotemperature derived from z-coral generic richness. D. Continental temperature curve (cold month mean) for Central Europe (from Mosbrugger et al., 2005,
modified). E. Oxygen isotopic composition of multispecies assemblages of benthic foraminifera (from Savin et al., 1975; and Dodd and Stanton, 1990, modified). F. Composite smoothed δ18O curve
(from Abreu and Haddad, 1998, modified). G. Global deep-sea oxygen record based on data compiled from more than 40 DSDP and ODP sites (from Zachos et al., 2001, modified). Time-scale of
Gradstein et al. (2004).

83
84 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88
Burdigalian (Fig. 6D) and the similitude of this curve
with our inferred temperature curve between that time
and the Tortonian may corroborate this view. The
taxonomic richness of the Mediterranean scleractinian
fauna is known to have declined during the Miocene
(Chevalier, 1962). One of the main cause for this
decrease in coral diversity is considered to be the
isolation of the Mediterranean basin from the late Early
Miocene to the Middle Miocene. At that time, seaway
connections with the Indo-Pacific Ocean through the
Middle East were disrupted by emergent areas and
northward rotation of the Arabian Peninsula (Rögl,
1998; Harzhauser et al., 2002), although several reopen-
ings and closures of this seaway during the Middle
Miocene occurred (Rögl, 1998). In the Eastern Medi-
terranean and Middle East areas, no scleractinian fauna
resulting from a mixture of Mediterranean and Indo-
Pacific species are known to occur after the Early
Miocene (McCall et al., 1994; Schuster and Wielandt,
1999), and the Mediterranean pool of z-coral began to
decrease after the Langhian onwards. By contrast, after
the Middle Miocene, the variations of z-coral taxonomic
richness-inferred palaeotemperature follow the changes
of the global sea-water temperature and the regional
continental mean cold month temperature (Fig. 6). The
decline of temperature is marked in the Serravallian in
the isotopic curves and appears to have been more
gradual from the Serravallian to the Messinian in the
temperature curve inferred from the z-coral generic
diversity, attesting for a sort of time-lag in the response
of z-coral assemblages to the decreasing SST. The
minimum sea-surface temperatures inferred from the
generic richness of the Mediterranean localities are
18.30°C and 17.80°C, respectively for the Tortonian
and the Messinian. This is in relatively good agreement
with the palaeotemperatures derived from the linear
growth rate of Upper Miocene Porites from Greece,
suggesting winter sea-surface temperatures not exceed-
ing 20–21°C (Brachert et al., 2006b).
7. Conclusions
The Mediterranean region provides a large number of
coral reef localities of Oligocene and Miocene age, used in
this report to test zooxanthellate-coral generic richness
values as a proxy for relative sea-surface palaeotempera-
tures by applying the “energy hypothesis” method (Fraser
and Currie, 1996).
This study highlights first that the “energy hypoth-
esis”, applied to Oligocene–Miocene z-coral assem-
blages of the Mediterranean region, provides results in
good agreement with the main trends obtained from
independent methods, such as global isotopic data
compilations and regional palaeoclimatic records from
continental floras (Mosbrugger et al., 2005) and from
fossil coral linear extension rate (Brachert et al.,
2006b).
Fluctuations of the inferred palaeotemperatures
show, in fact, a relatively good correlation with global
changes of sea-water temperature, especially for the
entire Oligocene, the Chattian–Aquitanian boundary
and the Late Miocene. In contrast, the Mid-Miocene
Climatic Optimum is not recorded in the z-coral generic
richness, showing that other regional factors, in part
related to major palaeogeographical changes affecting
the Mediterranean at that time, may have prevailed over
global factors.
The marked decline in taxonomic richness after the
Burdigalian is well correlated with decreasing palaeo-
temperatures in the region. This is associated with an
increase in temperature range of z-coral localities, which
clearly reflects the gradual adaptation of z-coral com-
munities to a wider temperature range from the Middle
Miocene onwards to the Messinian, as the Mediterra-
nean area was gradually moving northwards, outside the
tropical belt.
Potentiality of this approach is enhanced if applied at
global or regional scales. Additionally, as isotopic data
usually derive from the deep sea record, it appears a
promising and reliable proxy for estimating palaeotem-
peratures from reef palaeoenvironments or, in general,
from shallow-water carbonates. As such, the potential
application of this method could be tested similarly also
for other regions and maybe for other types of shallow-
water organisms like, for example, larger foraminifera.
A question could also arise about how back in time this
kind of approach can be used. As also corroborated by
methods applied to flora continental record (Mosbrug-
ger et al., 2005), it is, however, unlikely to find time
intervals older than the Cenozoic providing fossil
assemblages for which it is possible to suppose a
relationship with sea-surface temperature similar to that
of their modern equivalents.
Acknowledgements
We are particularly grateful to Brian Rosen for
fruitful discussions with both of us on this topic and for
his advice at the early stage of this study. We would like
to thank also two anonymous reviewers for their helpful
comments and suggestions. J.C. Braga, and F. Dela
Pierre and M. Zunino are thanked for providing valuable
stratigraphical information about the Sorbas Basin and
the Torino Hill localities respectively.
 F.R. Bosellini, C. Perrin / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 71–88
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