Palaeobiogeography of the Early Cretaceous corals

Palaeogeography, Palaeoclimatology, Palaeoecology, 92 (1992): 233-247

23~

Elsevier Science Publishers B.V., Amsterdam

Palaeobiogeography of the Early Cretaceous corals Louise Beauvais lnstitut de Pah;ontologie, MNHN, URA 12 du CNRS, 8 rue B~4[Jbn, 75005 Paris, France

(Received November 28, 1989; revised version accepted September 20, 1990)

ABSTRACT Beauvais, L., 1992. Palaeobiogeography of the Early Cretaceous corals. Palaeogeogr., Palaeoclimatol., PalaeoccoI., 92:233 247. A compilation of the geographical distribution of about 1000 species of corals from the Early Cretaceous (Valanginian to Cenomanian) is presented. After the reefal regression during the latest Jurassic, no corals appear in the Berriasian, and only a few species are known from the Valanginian and Hauterivian. In many regions it is impossible to distinguish pre-Barremian stages. Many of the pre-Barremian corals reported are Hauterivian in age. The pre-Barremian displays an explosion of the Scleractinia: more than 300 species extended through Europe, Africa, and North and South America. During the Aptian and AIbian, coral distribution was about the same as during the pre-Barremian but during the Albian the number of species decreased and the proportion of ahermatypic forms increased. Because of pronounced endemism during the Albian, faunal similarity no longer existed between the Americas and Europe. During the Cenomanian the Madreporaria nearly disappeared from North America but were still well represented in Europe. Half of the known Cenomanian coral species are ahermatypic, and they appear to be distributed on the two sides of the hermatypic buildup along the Tethyan Realm.

Introduction

Despite the insufficient information available about the Early Cretaceous corals, it seems useful to examine their worldwide biogeographic distribution. This adds to the information provided in a series of papers by Beauvais (1973, 1974, 1979, 1981 b, 1982, 1986 and 1989) on the palaeobiogeography of the Mesozoic Scleractinia. The present work is a first approach to assessing the biogeography of Early Cretaceous Scleractinia. The study is almost exclusively based on data derived from the literature. Because of the state of knowledge of Mesozoic Scleractinia, it is as yet provisional. More than 1350 species of Madreporaria were described by D'Orbigny (1848, 1850, 1862), De Fromentel (1857, 1862 1887), Koby (1897), Trautschold (1886), Thomas and Peron (1893), Vaughan (1899), De Angelis d'Ossat (1905), Gerth (1928), Alloiteau (1941, 1948, 1958), Alloiteau and Hup6 (1947), and others, and in more recent studies by Bendukidze (1961), Zlatarski (1967a, 1967b, 1968a, b), Turn~ek and Buser (1974), Sikharulidze (1979a, b), Morycowa (1964, 1976), and 0031-01~2/92/$05.00

'(i 1992

others. Unpublished determinations by M. Beauvais for the Urgonian of southeastern France (collections of J.-P. Masse and A. and H. Arnaud's) have also been used for this study. The biostratigraphical potential of Madreporaria is as yet unknown. A preliminary analysis to provide a stratigraphical base for the midCretaceous has been attempted (Beauvais 1981a), but since then no more work has been tmdertaken. The difficulty is due to the slow evolution of the group, in which the duration of species extends over more than one stage. For instance, it is often impossible to distinguish the pre-Barremian stages. In Japan, Eguchi (1951) described more than 200 hermatypic and endemic species as "Lower Cretaceous corals". More recently Wu and Liao (1981) cited from Xizang (China) eight coral species from the "Early Cretaceous", which have ranges from the Tithonian to the Cenomanian. These authors note: "it should be pointed out that Early Cretaceous fauna was still kept in a [.ate Jurassic aspect". Actually, in many areas, it is difficult to distinguish the Early Cretaceous from the latest Jurassic corals, and numerous species existed from

Elsevier Science Publishers B.V.

234

the Kimmeridgian up to the pre-Barremian (Diplocoenia stellata Etallon, Microphyllia undans Etallon, Maeandraraea gresslyi Etallon, etc.). Likewise, some Early Cretaceous species are found in the Late Cretaceous, e.g., Latusastraea provincialis d'Orbigny and Actinacis remesi Felix. Morever, authors like Wijsman-Best (1974), Pfister (1977), Beauvais (1977, 1982) and Lathuili&e (1988) remarked that numerous Mesozoic coral species are not true species, but represent varieties related to the environmental conditions. Thus the distribution of such species might reflect the distribution of such particular environments. For the conclusions of the present paper to be reliable, a systematic revision of every Early Cretaceous species must be completed, with simultaneous sedimentological studies and, in general, through the use of multiple criteria which can help to resolve potential errors of interpretation. This paper deals with the geographical distribution of species and their associations and with the environmental distribution of the hermatypic (colonial) and ahermatypic (solitary) forms. Where hermatypic corals are abundant, this is 'normally an indication of a reef environment. Where ahermatypic forms predominate, conditions for Scleractinian life were generally difficult (Stanley and Cairns, 1988).

L. BEAUVAIS TABLE 1 Comparison of Valanginian coral assemblages N

Switzerland Crimea East Africa Mexico

14 4 2 5

Herm.

Aherm.

Endem.

NT SET

(%)

(%)

(%)

(%) (%)

64 50 100 50

36 50 0 50

36 100 0 100

50 14 100

N = n u m b e r of species; Herm.=percentage of hermatypic species; Aherm. = percentage of ahermatypic species; Endem. = percentage of endemic species; NT=percentage of species shared with the northern Tethys (Great Britain, France, Switzerland, Spain, Poland, Romania, Crimea, Causasus); SWT= percentage of species shared with the south-western Tethys (Italy, Greece, Yugoslavia, North Africa, Sinai, Israel, Lebanon); SET = percentage of species shared with the south-eastern Tethys (Arabia, East of Africa, Madagascar, India); N A = percentage of species shared with North America (Texas, Mexico); SA = percentage of species shared with South America (Peru, Chile, Venezuela, Argentina). TABLE 2 Comparison of Hauterivian coral assemblages N

Eastern France Crimea Georgia Mexico

13 39 1 I0

Herm.

Aherm.

Endem.

NT NA

(%)

(%)

(%)

(%) (%)

100 97 100 70

0 3 0 30

85 60 100 40

8 26

8 13

50

10

Pre-Barremian coral distribution (Figs. I and 2, Tables 1-3)

For abbreviations, see Table 1.

In general, it is difficult to separate the three stages Berriasian, Valanginian and Hauterivian by means of corals. In earlier publications, the Valanginian often included the Berriasian. After a retreat of the coral facies at the end of the Late Jurassic, no corals occur in the Berriasian s.str. During the Valanginian s.str., they seem to have still a very limited distribution: fourteen madreporarian species have been described from Switzerland (Koby, 1897), four from Yugoslavia (Turn~ek and Buser, 1974), four from the Crimea (Trautschold, 1886; Kuzmicheva, 1967), five from Mexico (Burckhardt, 1930; Wells, 1946) and two from Kenya (Dietrich, 1926). They are long-lived species which in other areas persisted until the Aptian (Dimorphastraea tenuistriata de Fromen-

tel), Albian (Pleurosmilia stutzi Koby) or even Cenomanian (Actinastraea pseudominima Koby). At every locality, nearly half of the Valanginian species are ahermatypic, which indicates that conditions for coral reef development were limited. The Mexican Valanginian species are endemic and, consequently seem to define an isolated province. The Crimea also appears to represent a distinct basin. A comparison of the distribution of the Scleractinia with that of the belemnites for the Valanginian shows that the belemnites have approximately the same geographic distribution as in the Tithonian: Cylindroteuthididae continued isolated within the Boreal Realm, while Hibolites except at one locality in northern Scandinavia,

PALAEOBR)GEOGRAPHY OF I HE EARLYCRETACEOUS('ORALS

• 0

235

Vatanginian herrnaiypic corals Vatanginian ahermatypic corats

Fig. I. Distribution of Scleractinia in the Valanginian s e n s u stricto. The palaeogeographic map is by Smith and Briden (1977, Early Cretaceous reconstruction).

which probably indicates a warm current flowing north, occurs in the Tethyan Realm (The Crimea, The Caucasus, Somalia, Madagascar, India, China, Indonesia, northern Australia). Thus the scarcity of corals during the BerriasianValanginian does not seem to be due to climatic reasons; the limiting factors for corals may have been sedimentological events, such as high mud precipitation, resulting in lower light penetration, chemical composition of seawater, etc. On the other hand, the pre-Barremian displays a rapid rise in the number of localities with Madreporaria and in the number of madreporarian species. Moreover, 87% of the pre-Barremian and

Hauterivian species are hermatypic. These facts indicate that conditions for reef existence were more favourable than during the Valanginian. In France, Alloiteau (unpublished results) attributed an Hauterivian age to the faunas collected in the departments of Yonne and Haute-Marne, which were dated as "Neocomian" by De Fromentel (1857), but this fact cannot be generalized for every pre-Barremian coral-bearing formation. PreBarremian corals are found in France (HauteMarne, Aube, Yonne, Hautes-Alpes, Vaucluse, Var, Aude: De Fromentel, 1857), Switzerland (Koby, 1897), the Polish Carpathian mountains (Morycowa, 1964, 1971), Georgia (SSR) (Benduk-

236

L. BEAUVAIS

[rimea

[a~tUcla.sus

Vene

East Africa [hile

\

0 Ne0c0miancoral distribution • Hauterivian ss sfr, c0ra[ distribution ~_.%--~Percenfage of shared species Fig. 2. Distribution of the Scleractinia in the pre-Barremian. The palaeogeographic map is by Smith and Briden (1977, Early Cretaceous reconstruction).

idze, 1961), Yugoslavia (Burckhardt, 1930), Hungary (Kolosvary, 1 9 5 4 , 1959), Romania (Simonescu, 1962), Mexico (Sonora, Durango, Tehucan: Felix, 1891; Gerth, 1928; Wells, 1946), Peru (Fritsche, 1924), Texas (Wells, 1932), Argentina and Venezuela (Fritsche, 1924; Gerth, 1928), Tanganyika (near Tendaguru: Gregory, 1927) and Algeria (Alger and Constantine provinces: Coquand, 1879-1880). The distribution of the species indicates the existence of a Tethyan Realm, including France, Switzerland, Poland, Romania, the Crimea and an American Realm, including Texas, Mexico, Venezu-

ela and Chile. In both these realms the endemism of species defines the provinces: In the Tethyan Realm two provinces may be distinguished: (a) a North Tethyan Province (France, Switzerland, Poland, Romania) containing Actinastraea magnifiea de Fromentel, Actinastraea tenuis Koby, Clausastraea alloiteaui Morycowa, Eugyra digitata Koby, Ellipsocoenia regularis d'Orbigny, Hydnophora crassa de Fromentel, Latusastraea exigua de Fromentel, Pentacoenia microtema d'Orbigny, Stylina regularis de Fromentel, Stylina micropora Koby and Stylina pachystellina Kobby. A Crimean

237

P A L A E O B I O G E O G R e ~ P H Y O F T H E EARLY C R E T A C E O U S C O R A L S

TABLE 3 Comparison of pre-Barremian coral assemblages N Northeastern France Switzerland Southeastern France Poland Romania Crimea Algeria East Africa Venezuela Chile Mexico Texas

Herm. (%)

Aherm. (%)

Endem. (%)

NT (%)

SWT (%)

SET ~%)

I

0.7

142 21 22

85 57 72

16 33 27

70 57 73

13 42 26

25 15

100 93

0 6

43 70

43 30

11 10 10 5 4 32 27

91 70 80 100 100 100 93

9 30 20 0 0 0 7

83 73 55 60 25 79 89

18 33 40 75 18 4

SA (%~ 2 1

s

8

NA i%) 3 9

s ll 3 7

For abbreviations, see Table 1. Subprovince contains 83% of endemic species, which are poorly related to species of the N o r t h Tethyan Province; (b) a Southwest Tethyan Province (Algeria) comprising 73% of endemic species, 18% of species shared with other localities of the Tethyan Realm, and 8% of species c o m m o n with N o r t h America. In the American Realm three provinces have been recognized: (a) a Texan-Mexican Province with nearly 80% of endemic species, the other being c o m m o n with the Tethyan Realm: (b) an Argentine-Chilean province strongly related to N o r t h Tethys; (c) a Venezuelan Province with 60% of endemism and 40% of species shared with the Tethyan Realm. No connections seem to occur between these three provinces. Rare species occur on both sides of the Atlantic: Actinastraea minima de Fromentel has been found in eastern France, Switzerland, Poland and Chile; Actinastraea globosa de Fromentel has been described from eastern France, Mexico and Argentina: Co,ptocoenia neocomiensis de Fromentel occurs in France, Italy, R o m a n i a and Poland, and has also been found in Texas; Siderastraea cuyleri Wells occurs in Texas and Poland; Thamnasteria tobleri K o b y has been reported from Switzerland and Venezuela; Eugyra cotteaui de Fromentel, a

long-ranging species (Hauterivian to Cenomanian), has been found in eastern and southern France, Switzerland, Yugoslavia, Poland, The Crimea, Algeria and Mexico; Stylosmila alpina K o b y has been found in Switzerland, Georgia (USSR) and Venezuela: Siderofimgia zitteli occurs both in the Crimea and Mexico, and Siderofungia irregularis in the Crimea, Mexico and Texas. In comparison with the belemnite distribution (Stevens, 1973), Hibolites had a limited distribution during the Valanginian, became more abundant and then spread more widely, invading all of western Europe, North America and Patagonia. It is interesting to note that a s yet no corals have been described from around the Austral continent.

Urgonian-Aptian coral distribution (Fig. 3, Tables 4, 5). During the Urgonian to Aptian, the coral distribution area increased: Madreporaria reached southern England and Japan. The Urgonian coral facies (with 90% of hermatypic forms) is well developed in southeastern France (collections of Alloiteau, M. Beauvais and Masse), southern Switzerland (Koby, 1897), the Isle of Capri (De Angelis d'Ossat, 1905), eastern Serbia and Slovenia (Yugoslavia) (Turn~ek and Mihajlovic, 1981), the Polish Tatra Mountains (Morycowa, 1966), Georgia and

238

L. BEAUVAIS

o/

D D

D

0

0

~q PDOp/~Ce

~,AIM 0 Ppo,, ,d.

/ j

*

Urgonian hermafypic total distribution Aptian hermatypic coral distribution [] Aptian ahermatypic coral distribution ~_%..~Percentage of shared species Parahibolites realm (after Stevens 19731 - ~ Oinosaurus distribution (after [harig 1973] •

Fig. 3. Distribution of the Scleractiniain the Urgonian-Aptian. The palaeogeographicmap is by Smith and Briden (1977, mid Cretaceousreconstruction). the Crimea (SSR) (Solomko, 1887; Bendukidze, 1961; Sikharulidze, 1979b); Kenya (Gregory, 1930) and Venezuela and Chile (Fritsche, 1924). During the Aptian, the area of madreporarian occurrence extended widely: rich hermatypic coral faunas occurred in the Aptian of Great Britain (20 species: Duncan, 1879; Thomas, 1963), Spain (Catalonia and Aragon: Coquand, 1865; De Angelis d'Ossat, 1905; Bataller, 1937, 1949; Alloiteau and Hup6, 1947), eastern Carpathian Mountains (Romania and Hungary: Kolosvary, 1954, 1959; Morycowa,

1971), Venezuela (Gregory, 1927; Wells, 1944), Mexico (Wells, 1946), Trinidad (Wells, 1948), southern France (collections of Altoiteau and Beauvais), Switzerland (Perte-du-Rh6ne) (Pictet and Renevier, 1858), Bulgaria (Zlatarski, 1967a, b, 1968a, b, 1970) and Algeria (Coquand, 18791880). Only ahermatypic genera have been reported from the Aptian. In Japan a rich hermatypic fauna consisting of about 50 endemic species (Eguchi, 1951) is referred to the Aptian-Albian. The Urgonian-Aptian species display important

.3

PAI.AEOBIOGEOGRAPHYOF THE EARLYCRETACEOUSCORALS TABLE 4

Comparison of the Urgonian coral assemblages N

Switzerland Southeastern France Italy Poland Yugoslavia Crimea East Africa Venezuela Chile

Herm.

Aherm.

End.

NT

SWT

SET

NA

SA

(%)

(%)

(%)

(%1

(%)

(%)

(%)

(%)

98 100 91 83 84 86 95 86 i00

2 0 9 16 16 14 5 14 0

62 0 41 54 49 34 53 63 20

21 74 59 29 48 34 38 25 60

9 22

herm.

aherm,

end.

(%)

(%)

(%)

NT (%)

SWT (%)

20 11 6 3

75 100 0 0

25 0 100

30

100

40 23 12 46 18 13 12

97 17 75 56 83 100 92

0 3 83 25 43 17 0 8

95 32 83 0 33 39 87 3l 75 88 91 81

50 12 22 18 69 7 22 7 4

6

I 4

17 I1 8 12 20

3 22

For abbreviations, see Table 1.

TABLE 5

Comparison of the Aptian coral assemblages N

Great Britain Is6re-Ain Switzerland Southeastern France Poland Romania Bulgaria Yugoslavia Spain

Venezuela Mexico Trinidad

100

43 17

SET (%)

NA (%)

SA (%)

17

100 21 14 13 62 13

45 44

4

7 2

4 12

9 18

For abbreviations, see Table i.

endemism with a geographic distribution nearly corresponding to the pre-Barremian provinces outlined above. The general Urgonian endemic rate is 47%; this rate increases to 59% in the Aptian. In Great Britain 95% of the species are endemic, in Switzerland, 83%, in Bulgaria, 87%, in Spain, 75%, in Venezuela, 68%, in Mexico, 91%, and in Trinidad, 81%. Similarly, high endemism existed also in the Callovian (Beauvais, 1979). It seems to imply that species evolved in situ in each area as a result of the formation of different basins by geographical barriers which halted the dispersal of the planulae. These barriers may have been islands. river mouths, currents, volcanic eruptions rifts..

etc. In central Europe, the distribution of the dinosaurs (Charig, 1973) seems to reveal the presence of islands which might have isolated the different coral areas. Chemical changes (Renard, 1986) or a long-term warming of the oceans indicated by oxygen isotope analyses (Fischer and Arthur, 1977; Scholle and Arthur, 1980; Arthur et al., 1985) during this period may also have been responsible for the Barremian Aptian crisis. The Tethyan Realm included France, Switzerland, Poland, Spain, Italy, Yugoslavia, Romania, the Crimea, and the Caucasus. The northern part of this realm had closer relations with the southern part during the Aptian than during the pre-

240

L. BEAUVAIS

Barremian and Urgonian. In the same way the southern part of the Tethyan Realm was more closely related to Africa and Madagascar during the Aptian than during the pre-Barremiam. During the pre-Barremian, only Poland and the Crimea showed relationships with the east coast of Africa, while during the Aptian almost every basin (Great Britain, Is6re-Ain, Romania, and Spain) shared species with Madagascar. The American Realm remained unchanged from the pre-Barremian, with a strong endemism in the different basins and very few communications with Europe (8% of species in common between America, Spain and Poland) and few communications between the different basins (17 to 19% of shared species). Some more resistant species were probable able to adapt to certain unfavourable ecologic conditions and could therefore pass from one basin to another: Acanthocoenia neoeomiensis Volz has been found in the Aptian of Switzerland and Yugoslavia; Platyeyathus orbignyi de Fromentel and Parasmilia aptiensis Piet have been found in the Aptian of southern France, Spain (Aragon) and Algeria; Eugyra cotteaui de Fromentel occurred in the preBarremian through Aptian of Spain, the southern Alps, Yugoslavia and Japan. Only one coral species has been found to be common to both Europe and America, Stephanocoenia aragonensis Alloiteau, which has been described from Spain and Mexico. The species Thamnasteria decipiens

(Michelin), known from the Aptian of Mexico, migrated later (Cenomanian) into Czechoslovakia and Greece, probably as a result of the closure of the Tethys in the west towards the end of the Early Cretaceous. The relationship between Apulia and Magrheb exhibited by some fossil groups [rudists (Masse, 1985) and ostracods (Damotte, 1985)], is poorly demonstrated by the corals; as yet only one species is known to be common to the Aptian of Spain and the pre-Barremian of Tunisia. Among the belemnites the Cylindroteuthididae disappeared during the Aptian, while simultaneously Parahibolites spread widely throughout the world. This seems to indicate an important general warming of the earth, which started during the Hauterivian-Barremian as shown by the invasion of Hibolites into the Boreal Realm. This global warming is well evidenced by oxygene isotopes (Scholle and Arthur, 1980; Scott, 1988). Albian coral distribution (Fig. 4, Table 6) During the Albian, the distribution of the Scleractinia was similar to that of the Aptian, thus confirming endemism in every studied area. However, the number of rocks with colonial forms decreased while rocks with solitary forms increased. Albian coral rocks spread over Great Britain (Devonshire, Folkestone, Haldon and Yorkshire:

TABLE 6 Comparison o f t h e A l b i a n c o r a l a s s e m b l a g e s N

Great Britain Northeastern France Is+re-Ain Switzerland Southwestern France Georgia Tunisia Madagascar Venezuela Mexico

27 1 4 6 8 4 I 15 21 2

For abbreviations, see Table 1.

Herm.

Aherm.

End.

NT

SWT

NA

(%)

(%)

(%)

(%)

(%)

(%)

55 0 0 17 87 0 0 40 81 I00

44 100 100 83 12 I00 100 60 19 0

84 0 25 67 100 75 I00 93 81 50

16 100 75 34 25 6 5 50

14

241

PALAEOBIO(~EO(iRAI~HY OF THE [i'~RLY CRETACEOUS CORALS

\ Texas

LM 4-

Subprovint:e

1

/

• Hermatypic coral distribution in the A[bian 0 AhermatypiE coral distribution in the A[bian 4-- % ~ Percentage of shared species ~ Parah/bo/ifes realm (after Stevens 1973)

Fig. 4. Distribution of the Scleractinia in the Albian. The palaeogeographic map is by Smith and Briden ( 1977, mid Cretaceous reconstruction).

Jukes-Browne, 1875: Tomes, !885; Thpmas, 1963) with a great number of qherrnat.ypi~ geDera (44%). In France [Paderrl (Audei All,oi~eau, 1948), department of Ain, Pays-de-Bray (Beauvais, 1982)], rocks without buildups contain a majority of solitary corals. In Switzerland (Koby, 1897) and Georgia (Silkharulidze, 1972, 1979a) the coral fauna is entirely ahermatypic. In Mexico (Burckhardt, 1930), Texas (Wells, 1932) and Venezuela (Wells, 1944), the Albian coral reefs are overlying the Aptian reefs. In Madagascar (Alloiteau, 1958), among the fifteen known species, nine are solitary.

This abundance of ahermatypic corals cannot be explained by a cooling of the seas, since, after the early Aptian, boreal belemnites disappeared and were abse~at during the Albian (Stevens, 1973). It appears that general environmental conditions characteristic of this age such as abundant argillaceous sedimentation or oxygen deficiency in the sea waters (Chamley, 1984), hindered the growth of hermatypic Scleractinia. Sedimentological and geophysical studies are necessary to explain the phenomenon. The high rate of endemism noted in Great-Britain, Central America, Madagascar and

242

L. BEAUVAIS

Japan can probably also be explained by a comparison between the distribution of the corals as related to that of the sedimentological facies. The Tethyan Realm was well defined in the preBarremian and Aptian, maintained during the Albian with no distinction possible between North and South Tethys except with regard to Madagascar; this Tethyan Realm included Great-Britain, France, Switzerland, and Georgia (USSR). In Tunisia, only one hermatypic and endemic species (Phyllocoenia ferryi Coquand) has been recorded. A Malagasy Subprovince appeared during the Albian with 93% of autochtonous species.

The American Realm continued to be isolated: among the 21 species found in Venezuela, 17 are endemic, the others have been reported from in Mexico and Texas. Only two ahermatypic species of Trochocyathus occurred on both sides of the Atlantic (eastern France, the Alps and South America).

Cenomanian coral distribution (Fig. 5, Table 7) During the Cenomanian, the Scleractinia decreased in the Americas (Gregory, 1929) thus indicating the complete closure of the Tethys, a

\ Tethyan --.I

\

Texas •

- Tunisian Subprovince

,dr

\

dian

[enomanian hermafypic coral distribufion [enomanian ahermatypic coral disfrib~fion • l-%-i~Percentag e of shared species j ~, Hibolites realm (affer Sfevens 1973] Fig. 5. Distribution of the Scleractinia in the Cenomanian. The pa]aeogeographic map is by Smith and Briden (1977, mid Cretaceous reconstruction).

243

P A L A E O B I O G E O G R A P H Y O F T H E EARLY C R E T A C E O U S C O R A L S

TABLE 7 Comparison of the Cenomanian coral assemblages

Belgium Northeastern France Western France Italy Greece Czechoslovakia Algeria Tunisia Lebanon Sinai India Madagascar

6 1 50 8 97 41 1 9 10 1 55 13

Herm.

Aherm.

End.

NT

SWT

SET

(%)

(%)

(%)

(%)

(%)

(%)

100 0 26 I00 80 76 0 33 70 0 65 8

0 100 74 0 20 24 I00 67 30 100 35 92

66 100 92 25 77 72 0 100 80 0 81 100

33 2 37 4 13 100

12

4 37 7 13

2

20 100 17

For abbreviations, see Table 1.

process that had begun already in the Aptian. Henceforth, tile relatively deep Atlantic ocean was completely isolated from the Mediterranean Tethys, a fact indicated by the ammonite distribution (Matsumoto, 1973). During the same time, in northern America and in northern Europe, a boreal belemnite realm appeared with the Belemnitellidae and the genus Actinocamax. Thus we can envisage cold currents across America from the north, hindering the growth of the Scleractinia. On the contrary, in Europe, during the same period, an expansion of the colonial Madreporaria occurred. They spread both to the north and south across Belgium (D'Orbigny, 1862), northern Germany, Bohemia (Poeta, 1887) and Westphalia (Plauen region, Dresden, Pirna, Teplitz, etc.: B61sche, 1871; Felix, 1906), southwestern France (Corbi6res: Alloiteau, 1941), Italy (Ocre Mountains: Prever, 1909), Palermo (MontanaroGallitelli, 1937), Greece (Dremisa: Hackemesser, 1936: Renz, 1930, 1931), Karpathian Mountains(Morycowa, 1966), Austria (S6hle, 1896), Israel (Avnimelech, 1947), eastern Jordania, Lebanon, Syria (Felix, 1909) and southern India (Stoliczka, 1873). However, ahermatypic forms still prevailed in some parts of northern Germany, western France (Le Mans, Aix Island and Rochefort: Alloiteau's collections), Tunisia (Thomas and Omara, 1957: Gill and Lafuste, 1987), and Madagascar (Collignon, 1931: Alloiteau, 1958). This area of

coral distribution corresponds to that of the Tethyan belemnites (Parahibolites and Neohibolites), whereas no Scleractinia occurred within the boreal belemnite realm. The ahermatypic Scleractinia appear to be distributed as a northern and southern belt along the hermatypic coral area. Although a clear endemism is still observed in each region - - there are reports of 77% of endemic species in Greece, 80% in Lebanon, 81% in India, 92% in western France, 100% in Tunisia and Madagascar. The distribution of the species defines in the Tethyan Realm a North Tethyan Province and a South Tethyan Province. The northern Province consisted of Belgium, western France, northern Germany and Czechoslovakia with the species Leptophyllia patellata Michelin, Barvhelia archiaci (Milne-Edwards and Haime), Montlivaltia tourtiensis B61sche, Synhelia gibbosa Goldfuss, Micrubacia coronola Goldfuss, Parasmilia rudis Reuss, and Porites michelini Reuss. The South Tethyan Province included Greece, Algeria, Yripolitania, Sinai, Israel, Lebanon, Syria and Jordania with the species: Aspidiscus cristatus Lamarck, Aspidiscus felixi Renz, and Actinastraea pseudominima Koby. Greece showed relationships both with the South Tethyan Province (Actinastraea kunthi B61sche and Favia ambigua Michelin) and the Madagascar and India subprovinces with the shared species Actinastraea retiJera Stoliczka, Polytremacis edwardsi Stoliczka, Epismilia crassisepta

244 Stoliczka, Isastraea siva Stoliczka, Heliopora edwardsi Stoliczka, Holocoenia indica Stoliczka, Placosmilia similaris Stoliczka, Thamnasteria pullulata Stoliczka, Th. microglyphica Stoliczka and Th. induta Stoliczka. Hungary, Tunisia and Madagascar seem to have been completely isolated basins since they show an endemism rate of 100%. Tunisia may have been isolated by faults (Dercourt et al., 1984). When we compare the distribution of corals with that of the belemnites, we observe that the Duvalia Mediterranean Realm of Stevens (1973) corresponds approximately to the North Tethyan coral Realm, while the Duvalia CarpathianCaucasus Realm corresponds to the Hungarian coral area.

Summary Early Cretaceous coral genera retain a Late Jurassic aspect. Numerous Tithonian species range into the Valanginian. However, the genera Eugyra, Myakosmilia, Microseris, Hydnomorphomeandra and Felixigyra apperar in the Valanginian and are specific for the Early Cretaceous. The genus Aspidiscus is characteristic of the Albian-Cenomanian of the Mediterranean basin. During the Cenomanian, new genera appeared and ranged into the Late Cretaceous, e.g., Baryhelia, Sinaimeandra, Elasmocoenia, Antiguastraea. The Tethys appears to have been a dispersal center for the corals: the first Early Cretaceous species appeared in Europe during the Valanginian, Hauterivian or Barremian and persisted into the Cenomanian in more remote areas (East Africa, Lebanon and India). For example, Dimorphastraea bellula (D'Orbigny) is first known from the Urgonian and Hauterivian of Switzerland and Crimea and reached India during the Cenomanian. The Tethyan ocean probably served as a continuous corridor for dispersal of larvae corals along the shallow seas of its flanks. The extent of these equatorial routes diminished towards the end of the Cretaceous, with a progressive closing of the old ocean and widening of the Atlantic. Corals have not yet been found so far in the Berriasian, and very few Valanginian Scleractiniabearing rocks are known. The general retreat of

L.BEAUVAIS the seas had begun by the end of the Jurassic. It was followed by an emersion of the platforms and a concomitant deepening of the oceans events which impeded the growth of the building corals. During the Hauterivian, a transgression occurred with formation of carbonate platforms around the continents. Consequently, colonial corals could grow inside small basins isolated by geographical barriers such as islands, currents, river mouths, etc. In these small basins, isolated coral populations evolved and sympatric speciations took place, which resulted in strong endemism among the species. Thus in the Tethyan Realm and the North Tethyan Province, a Crimean Subprovince seems to have been isolated, and in the American Realm three provinces may have existed: the Texan-Mexican Province, the Argentine-Chilean Province and the Venezuelan Province. However, some more vigourous species with long-lived larvae [some scleractinian species may swim several months before their fixation (Chevalier, 1987)] were probably able to cross from one basin to another. During the Urgonian Aptian, the reef expansion that had begun in the Hauterivian continued with the development of coral formations reaching Great-Britain and Japan. During this period, endemism of species was still very high, with geographical barriers probably inhibiting the free dispersal of the planulae. Connections with the Americas became rare, a situation probably caused by the beginning of the westward closure of the Tethys and the opening of the Atlantic Ocean. During the Albian, coral-bearing rocks were still widespread but ahermatypic forms began to dominate; abundant argillaceous sedimentation and anoxic conditions during this time are evoked to explain this phenomenon. Endemism was still very strong: in the Tethyan Realm a Tunisian Subprovince and a Madagascan Subprovince have been recognized, both are distinct from the North Tethyan Province. During the Cenomanian, the madreporarian formations decreased; they were markedly reduced in America and replaced by rudists; the building corals developed in the Tethyan Realm, in which a North Tethyan and a South Tethyan Provinces may be distinguished. Important affinities have

P A L A E O B I O G E O G R A P H Y O F T H E EARLY C R E T A C E O U S C O R A L S

b e e n o b s e r v e d b e t w e e n G r e e c e a n d the s u b p r o v inces o f e a s t e r n A f r i c a a n d I n d i a . T h e d i s t r i b u t i o n o f the T e r e b r a t u l i d a e g i v e n by M i d d l e m i s s (1980) also s h o w s t h r e e areas: a B o r e a l R e a l m , a T e t h y a n R e a l m , a n d , in b e t w e e n , the neritic t e r e b r a t u l i d f a u n a o f the J u r a , l o c a t e d o n the flank o f the d e e p T e t h y s , w i t h affinities w i t h M o r o c c o ; this f a u n a s p r e a d a c r o s s the T e t h y s f r o m S p a i n to T u r k e y . A c c o r d i n g to p l a t e r e c o n s t r u c tions, d u r i n g this p e r i o d the t e r e b r a t u l i d f a u n a o f the J u r a c o n s t i t u t e d a belt w h i c h e x t e n d e d f r o m P r o v e n c e a n d e a s t e r n S p a i n , a c r o s s S a r d i n i a , the Baleares, s o u t h w e s t e r n M o r o c c o , w e s t e r n U S A to n o r t h e r n M e x i c o . T h e d i s t r i b u t i o n o f the t e r e b r a tulid f a u n a o f the J u r a c o r r e s p o n d s to the d i s t r i b u tion a r e a o f corals. T h e d i s t r i b u t i o n o f the B i v a l v i a ( K a u f f m a n , 1973) s h o w s s i m i l a r b i o g e o g r a p h i c units: a T e t h y a n R e a l m , i n c l u d i n g in the n o r t h a M e d i t e r r a n e a n P r o v i n c e , in the s o u t h an I n d o - M e d i t e r r a n e a n region and a North Indian Ocean Subprovince, a n d , in the w e s t a C a r i b b e a n P r o v i n c e w h i c h extended across Central America. The Tethyan R e a l m o f the b i v a l v e s was b o u n d e d to the n o r t h by the E u a m e r i c a n r e g i o n w h i c h i n c l u d e d a N o r t h American Province and a North European Province.

Acknowledgments | thank Mrs Benkheiri from the Universit~ Pierre et M a r i e C u r i e (Paris) w h o assisted m e in the d a t a processing, Mrs. Durand f r o m the M u s d u m N a t i o n a l d ' H i s t o i r e N a t u r e l l e de Paris w h o d r a f t e d the figures, M r s M a r d c h a l for h e r help, a n d the critical r e v i e w e r s o f m y m a n u s c r i p t , w h o g a v e m e precious counsels and remarks and particularly h e l p e d m e w i t h the E n g l i s h v e r s i o n .

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