(
Ciencias da Terra(UNL) , <i ~I'
Lisboa _I N° 14 pp.323-334
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2000...
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Miocene marine mammals from Portugal
Paleogeographical and paleoecological significance
Mario Estevens
Centrode &ludoaGcol6gicos,FaculoaiedeCi&lcia!le Tecnologia,Quinta
ca
TOfTe, P-l 825-I !4Capariea,PortugalPbonc:(35 I) 2 12948573, Fax: (35 I) 212948556, E-mail:[email protected]:
ABSTRACT
Key words: Marine manvnals; Miocene; Portugal; geographic and stratigraphic distriburion; correlation with environmc:ntal changes; paleogeognphical and paleoecological significance.
Neogene marine mammalsarestill incompletelyknownin Portugal. However, a general overviewof the geographic and stratigraphic distribution of marine mammal localities in the Miocene of Portugal is already possible.Anattempt of correlation between the trends shown by these distributions and the horizontal and vertical environmental shifts is presented In general, sirenians occur in deposits representing shallow,
warm.
low energy aquatic environments; while cetaceans are more frequent in more open, deep and temperate marine environments.RESUMO
Palavras-chave: Mamlferos marinhos; Miocenico; Portugal;d i stribui~ o geografica e esrratigraflca;conet acsc com variacces ambienlais, significadopaleogeografico e paleoecologico.
o conhecimento dos mamiferos marinhosneogenicoseainda incompletoemPortugal.Ccnrudc, e pcssiveldelineara disrribuiego geografica e estratignifica das jazidas de mamiferos marinhos' do MiocCnico de Portugal. Tentou-se, igualmente, relacionar as tendencies cxibidas par estes distribuilj'Oes com varialj'Oes horizontais e verticals das condi¢es paleoambicntais.Demodo geral, cs Sirenia ocorrem em ambientes pouco profundos, de aguas quentes e fraca energia; enquanto que os Cetacea sio mais frequentes em ambicntes marinhoseberros,demaierprofundidadee aguas maistempcradas.
INTRO DUCTIO N
Thi s paper
is an
updated vers ion o f a preliminaryre-port on the same subject (Estevens, 1998a).It includes new data onthecolle ctions of the "M useu Nac ional de Hist6ria Natural da ljniversidade de Lisboa" (Na tional M useum o f Natural History oftheUni versity of Lisbon) . Th e co llections of
the
" lns tituto Geol6gico e Mineiro" (Geo logicaland
Mining Institute) were reevaluated, andsome newspecimens were added to the collections de-posi ted at the "Departamento de Ciencias da Terra da Faculdad e de
Ciencies e
Tecnol ogia daUniversidade
Nova de Lisboa" (Department ofEanh
Sciences oftheFaculty of Sciences and Technology o f the New University of Lisbon).Although long known in Portugal (Escbwege, 183 1), Mioce ne marin e mammals have not been mucb studiedin
this
country (Antunes, 1984).Onlya few sbort descrip-tive notes have bee n published (Vandelli, 1831 ; Kellogg. 1938-40 ; Zbyszewski, 1944, 1953, 1954; Antunes , 1959, 1979; Mata, 1963; Jonet, 1980-81 ).Aconsiderab le number o f undescribed specimens can befound in Portuguese collec tions.Th e systematic study of the Neogene marin e mam-mals ofPortugal
was resumed
bytheauthorat
the "Centro de Es tu dos Geol6gicos da Facu lda de de Ci encias eTecnologie",
underthesupe rvision ofProf. Miguel TellesAntunes .The
first
results about a sma ll asse mb lage o f odontocetes from the Lower Tagus Basin Miocene will soonbepresent ed. M eanwhile, based on published dataThe grea t majo rity (85 ,7% ) of the 56 localities are located in the distal reg ion of the Lowe r Tagus Basin (LTB ). This incl udes Lisbon reg ion (LR) and Setubal Peninsula (SP), located respectiv ely north and south of Tagus River (Fig. 2). Few localities can be found southwards ofthis area : only 5 in the vast region encompassing Atcacer do Sal, the littoral ofAlentejo andAlvalade Basin (Fig . 3), and a mere 3 in Algarve (Fig. 4 ). A single additional record (not disc ussed here), concerns the Santa Maria Island, in the Azo res Archi pelago (Cotter, 1888-92).
The predomi nance of mari ne mammal localities in the LTB results largely from the fact that this bas in conta ins the most co mp lete Miocen e mar ine seri es in Portugal (Antunes
et aI.,
1996). As a consequence, the LTB has been both the most stud ied an d the one from where more fossils were co llected. Barnes (1976: 338) noted that for the eastern Nort h Pacific fossil cetacean assemblages a biased collec ting might be part ly on the origin o f such diversity or abundance trends. A more detailed prospection of the other Tertiary Portuguese basins may lead to the disco very o f new localities. However, this wou ld j ust att enu a te the obs e rv ed geogra p hic t re nd wi thout produ cing any dra matic changes . Th e LTB will certainly rem ain the mo st im po rtan t area in Por tu gal in wha t concerns Miocene fauna in general, and marine mammals in pa rticular.A significant geographic segrega tion can be observed between Sircnia and Cetacea locali ties within the distal region of the LTB (Fig. 2).
10Congresso sabre
0Cenozoico de Portugal
and on the inventory of specimens from co llections, it is already possible to present a list o f Portu guese M iocene loca lities where marine mammal remains were registered . Pliocene reco rds are scarce and somewhat dubious (see below). At this poi nt, only the presence of Cetacea, Sirenia or "Pi nnipedia" will be ac knowledged for eac h locality, with no indication of subordinal assignment. Such a list allows to outli ne the geographic and stratigraphic distri-bution of each group, focusing on the main differences between them. The paleogeographical and paleoecol ogic al significance of such differenc es was eva luated by corre-lating the det ected trends with hor izontal and vertica l changes of the paleoenvironmental conditions .
The conc lusions herein presented were hampered b y some difficulties concerning both the p ublished data an d the inventoried co llections.
Pub lished records of marine mammals are not always reliable in what concerns system atics, even at an ordinal level. Many older references of cetacea ns for the Early M iocene of Lisbon probably concern sirenians (pa rtic u-larly those in Cotte r, 19 56 and Choffat, 1950a). Another example is a record for the Burdigalian of Foz da Fonte (48 ), originally reported to Cetacea (Zbyszewski
et al.,
1965) and later reassigned to Sirenia (Zbyszewski, 1967). As most ofthesc records correspondtofragmentary material not always collected, it is difficult to determi ne their tru e taxonomic nature. Nevert heless , the scarce biblio graphi c referenc es for which a positive corres pondence could be established with material deposited in the studied collec-tions ind ica te tha t mos t (i f no t all) Early M io ce ne published records of marine mammals corres pond in fact to sirenians (as noted by Antunes, 1959, 1969-70, 1984 ). The systematic nature of some references, however, could not be de finitely asce rtained. In these cases, the taxonomic uncertainty will be indicated either by a broken outline of the symbols repre senting each marine mammal orde r (in Figs. 2 and 5), either by hatched areas (in Fig. 6).The inverse prob lem was fou nd when pro ceeding to the inventory offossil marine mammals in Portuguese co l-lections. In this cas e it was not the systema tic assignment of the specime ns that constituted a problem (at least for the majority of them), but rather the definition of the ir provenance. A large part of'the collections of the "I nstitute Geologico e Minciro" raised too many problems in th is respect. Man y undescribed specimens are not labeled, and even for those sitting in labeled trays there is a strong possibi lity of having been misplaced. l f for some it is still possible to ded uce their original locality of provenan ce (through the adhering matrix or their relatio n to other well-known specimens), for others this was impossib le.
The refore, the list of localities presented and the conclusions therefrom derived arc based only on those rec or ds for whic h the geograp hic an d stratigrap hic occurrence could be safely determined.
GEOGRAPHIC DISTRIBUTION
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Ciencias da Terra (UNL) , 14
Marine mammal loc alities
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10Congressc scbre
0Cenczcico de Portugal
Sir enia localities arc almost exclus ively located in LR, many of them within the limi ts of Lisbon. There are only two localities to the west(S-Barreiro da Maruj a and 10 -Pra ia de Carcavelos ) and ano ther two near the north ern ou tskirts (8 - Casal do s Cucos and 13
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de s Fctais). Santa lria daAzcia
(23) co uld constitute the northernmost reco rd of Sirenia for the Portuguese Miocene, but see be-low discussion on uncert ain taxon omic ass ignme nt of re-mains therein fou nd. Sirenians are scan t south of Tagus River, occ urring only at Pene do Sui (47) and Foz da Fonte (48).Cetace an s, on the other hand, predominate in SP (86,7% of marine mamm al localities in this area) . Those loc ated north of Fagu s Ri ver are co nc en trated i n easternm ost Lisbon. Tune! do Rossie (1 ) would cons titute th e exception, hut the rea der is aga in re ferred to the discussi on on unce rtain taxonomic assi gnments below.
The single "p inniped" record cons ists of phocid re-mains from Pen edo Norte (46) (Antuneset af.,1995).
In the Alvalade Basin ( Fig. 3), marine mammals were reco rded at the three localities that have produced the rich -est fossil assemblages of this region ( Antuneset al., 1986 ).
G EO LOGY Of AL CA CIEA 00 SAL RIEGIO N .
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Cunctas da Terra (UNL), 14
Fig. 4 • Geogra phic distri bution of Miocene marine mammal localities in the Algarve. Geology ada pted from Oliveiraet al.
( 1992 ), revised byJ.Pais. Only cetaceans were found at Esbarro ndadoiro (52), whilst
there were only sirenians at Vale de Zebro (53 ). Acco rding to a personal co mmunication from
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Pais, sirenia n rib fragments arc to be added to Santa Margarida do Sado (5 1), fro m where only cetacean re ma ins were kno wn (Estevens, 1998a ). T he latte r thus beco mes one ofthe few Portuguese loca lities where fossils of these two groups occur simultaneously.Marine mamm als were also recorded at Ce rrado da Pedra (4 9) (SireniainAntunes& Mein, 1983), located in the sout heastern extension o f the LT B (Alcace r do Sal re-gion ), and in M elides (50) (Cetacea
in
Zbyszewski, 1954). Only one loca lity in Algarve (Fig. 4) had so far been referred in the literature as yielding marine mammal remains: Ol hos de Agua (55), from where both Sirenia and Cetacea were described (Ant unes, 1979). T his and Santa Margarida do Sado are the only Portuguese localities where sirenians and cetaceans were simulta neously recorded. The inven-tory of the P ort uguese collections revea led fur ther undescribed speci mens of cetaceans from the M iocen e o f Algarve, collected at Cerro das Mo s (54) and Cacela (56) .STRATIGRA PHI C DISTRIBUTION
T he s tr atigra phic distri bution of the 56 Neogene localities w here marine mammals were recorded is summarized in Fig. 5.
Marine mammals occur in almost every Portuguese M iocene marine unit, from the earli est ones at the distal region of the LTB (Aquitanian DS A I) , to the latest ones at the Alvalad e Basin (la te Me ssinian, Esbarrondadoiro For mation ). Inversel y, only the littor al marine sands at O lhos de Ag ue (5 5), in Algarve, could be referred to the Pliocene, according to a recent87 S r ~ 6 S r isotopic da tation of a mollusk bed locat ed j ust above the sands (An tuneset
aI.,
2000b). T he ve rtebr ates fou nd in a lower stratigrap hic level( a ~ ng which the marine mamm als), seem how ever to indicate a M iddle to La te M iocene age (Antunes, 1979). These may thus constitut e a redeposited ass ociation (Antunes , personal co mmunicatio n).Fig. 5 also shows signi fica nt differenc es between the grou ps o f mari ne mam ma ls i n what concern s their stratigraphic distri bution .
Within the distal region o f the LTB , sirenians arc freq uent in LR fro m the Aquitanian to the Langhian, nearly disa ppearing afterwards (Antunes, 1984). The few midd le Serravallian recormidds (Esteve ns, 1998a) are now con -side red too do ubtful and were consequently disregarded. Only two sirenian records are known for SP, both of them Burd igalian (loca lities 47 - Penedo Su i and 48 - Foz da Fonte) . A rib fragmen t co llected at Santa Id a da Azoia (23) was questiona bly ascri bed to Cetacea (Choffat, 195 0a: 120). Th e fragment see ms to indicate a sirenian instead, and in that case it would cons titute the latest record of the grou p (early Se rravallian) for the dis tal region of the LTB. Conversely, in the so utheas tern extension of th is same basin (Alcacer do Sal region), sirenians oc cur later (la te Serravallian) at Ccrrado da Pedra (49) (Antunes& Mein, 1983). Further to the sou th, in the Alvalade Basin, they
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occur even later (Messinian) at Santa Margarida do Sado (5 1) and Vale de Zebro (53). The only sirenian record in Algarve is from Olhos de
Agua
(55) (Ant unes, 1979 ), supposedly of Pliocene age (see Antuneset al.,2000b and above) but probably older (Middle to Late Miocene) and redeposited.Inversely to sirenians, cetaceans only became common from the Langhian-Serravallian boundary upwards. At the LTB they occ ur mostly in the Se rravallian and early Tortonian (Antunes, 1984). Outside this basin, there is one record for Melides (50) (Zbyszewski, 1954), correlative of the most modem well-dated levels of the LTB. On the other hand, cetaceans are known from latest Miocene beds at the Alvalade Basin (Messinian localities 51 - Santa Margarida do Sado and 52 - Esbarro ndadoiro) (Antunes
et al., 1986), and Algarve (late Tortonian ofCacela - 56). Also in Alga rve , ce tacea ns were found at O lhos de Agua (5 5) (Antunes, 1979) and Cerro das M6s (54). The latter (on a sma ll hill near Lagos) is now recog nized to be Serravallian in age, according to recent isotopic datations of nearby co rrelative deposits (Antuneset al.,
2000b).
So far, the only Early Miocene locality that unques-tionably yielded cetacean remains is Palcnce de Baixo (36) (middle Burdigalian) , in SP (LTB). There is a doubtful record for the Aquitanian
ofTunel
do Rossio (1) by Choffat ( 1889: 40): "os de grand e taille, incomplet, peraissant appartenira
un crane de Cetace". As no specimens possi-bly related to this occurrence were found, it must be here held questionable.Th e "pin niped " fro m Pe nedo Nor te (46) is la te Serrava llian in age (Antuneset al., 1995).
PALEOGEOGRAPII ICAL AND
PALEOECOLOGICAL SIGNIFIC ANC E
The geogra phic and stratigraphic trends summarized above can be correlated to environmental changes well documented in the literature.
The most striking case concerns the marine mammal record of the LTB Miocene, which was already the object of general considerations (Estevens, 1998b). Fig. 6 shows the e lo se correla tion be twee n va r iati ons of the paleoecological cond itions (namely sea temperature and depth) and the number of Sirenia and Cetacea localities for the distal region ofthe LTB.
Sirenia occur from the earliest Mioeene (Aquitanian) to the early Middl e M iocene (Langhian), when sea tem-peratures stayed over :::::24°C (ranging from tropical to sub-tropical environments according to Antunes&Pais, 1984). Sirenia are present whene ver very shallow near -shore environ me nts a re co ncerned, an d ab sent wheneve r environments became slightly deeper (never attaining the circalittora l stage). The maximum abundance of Sirenia occurred during the Langhian, when temperatures were highest in shallow, ncar-shore to fluvial tropical environ-ments (Antunes& Pais, 1984, Antunesetal.,1996). After the Langhian, a marked drop in temperature and increase in depth led to the disappearance of sirenians.
Cunctas da Terr a (UNL), 14
This change of envirorunental conditions turned the balance to the Cetacea favor. Cetaceans became frequent and reach maximum abundance at events of greater depth (circa or ncar-circalittoral stages ) during the Scrravallian and early Tortonian. Itis noteworthy that the only Early Miocene Cetacea occurrence correspo nds to the apogee of the Burdigalian transgression, another maximum depth event. The number of localities that yielded cetacean remains became much reduced at the late Serravallian.
The stratigr aphic nea r-e xcl usion of sirenians and cetaceans in the distal region ofthe LTB is thus a result of important environmental changes that took place during the Langhian (Antunes& Pais, 1984). The subtropical to tropical climate and shallow, protected marine gulfs that prevailed from the Early to early Middle Miocene (Antunes
etal.,1996) were an idea l habitat for sirenians (Domning, 1976, 19 78). The more tempe rate and deep er marine envirorunents that followed (Antuneset al.,1996, 2000a), were instead more favorable to the occ urrence of cetaceans (Barnes, 1976; Fordyceet al, 1994) .
The geograp hic distribution of marine mammals in the d istal region of the LTB is also co ntro lled by the se paleoecological factors. The prevailing oc cu rrenc e of siren ians in LR and cetaceans in SP is related to the predom inance of esse ntia lly Early Mioce ne shallower depos its in the former, in contrast to the deeper marine Middle to Late M iocene sediments that cover most of the latter (Antunesetal.,1996). The occurrence of a phocid in Penedo Norte (SP) agrees with these assumptions.
The scarce representation of Miocene marine mam-mals outside the distal region of the LTB is related to the lesser development of marine units.
The permanen ce of sirenians in the Alcacer do Sa l region and Alvalade Basin (until the Late Miocene in the latter) is an indic ation of the persistence of shallow protected gulfs and moderately warm waters (Ant unes, 1983; Antuneset al.;1986). This took place well after the increase in depth and coo ling verified by the end of the Langhian in the distal regio n of the LTB, that eventually led to the disappearance of sirenians from this area. The southwar ds with drawal of Sireni a in th e Portuguese Miocene is in agreement with the accentuation of the ther-mal latitud inal grad ient recognized for the northwestern Atlantic dur ing this epoch (Lauriat-Rageet al., 1993).
The cetacean record of Melides (50) is compatible with the mar ked ly op en an d litt oral ma rine env ironment recognized for both these deposits (Prates & Carvalho,
1983-85) and their correlatives in the LTB.
In the Alvalade Basin, the contemporaneous occur-rence of open marine envir onments (either shallower as in Santa Marg arida do Sado, either relatively deeper as in Esbarrondadoiro) with the restricted inner gulf of Vale de Zebro (Antu nesetal .,1986), made possible for cetaceans and sire nians to occur simultaneo usly d uring the late Messinian.
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.
C"<t.,
Fig. 6 - Correlation between the variation ofsea temperature and depth and the numberofSiren ia and Cetacea localities at the Lower Tagus Basin (Lisbon region and Setubal Peninsula) during the Miocene. DS stands for deposi cional sequences (Antune set al.,
2000a) and LLU for locallithostratigraphic units (Cotte r, 1904).
01.
(2000b). Vertebrates found in the same deposits point out. however, to near-tropical cond itions (Antuneset 01.,
1981 ). Cacela (56) co rrespo nds to a litt ora l marine environment (Antunes et01.,
1981), and Olhos deAgua
(55) to a shallow coastal environment witha nearby estuary (Antunes
et al.,
2000b) .In conclusion, the.distribution of marine mammals in the Portuguese Miocene is closely controlled by ecologi-cal restrictions. Sirenia occur mostly inshallow, warm, low energy aquatic environmen ts (a s po st ulated by Domn ing, 197.6, 1978). while cetaceans are more frequent
in more open, deep and temperate marine environments (as recognized by Barnes, 1976 and Fordyce
et al.,
1994). The paleogeographical and paleoecological sig-nificance of marine mammals, and their utility in the reconstitution ofmarine paleoenvironments, is thus re-asserted by the present study.ACKNOWLEDGMENTS
First and foremost to Prof. Miguel Telles Antunes, who from the beginning of this study has provided all
the oppo rt unitie s and ongoi ng support, as well as a thoro ugh review of this manuscript. Also to Doctor M. Magalhaes Rama lho (Vic e-Presi dent of the " Institu te Geo l6gic o e M ineiro" ) and Prof. A. M. Galop im de Carv alho (Director of the "M useu Nacional de Histor ia Natura l da Universidade de Lisboa") , who have granted acc ess to the fossil coll ections under their care.
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