Ci!ocias.da Terra (UNL)

Late Quaternary warm marine mollusks from Santa Maria (Azores) paJeoecoJogic and paJeobiogeographic considerations

P. CallapeZ< 1) & A. Ferreira Soares(2)

1 . Departameolo de Ciheias da Teml da Universiclade de Coimbra, 3049 Coimbra Codex; eal1a~i.ue.pl 2 _ Dcpartamcnlo de Ci~nei a s cia TC1TI1 cia Universidade de Coimbra, 3049 Coimbra Codex;

RESUMO

Palavras-<:have: Plistocenico; invertebrados marinhos; Macaronesia; paleoecologia; paleobiogeografia.

A fauna marinha sub-f6ssil dos depOsilos quaternarios de Santa Maria e composta por mais de SO especies de gastr6podes e de bivalves, 19 das quais foram recolhidas reccntemente e pela primeira vez oa costa Norte da ilha (Baia de lagoinhas). As conehas sub-f6sseis encontram-se em depOsitos corn ardas de praia, situados 2 a 3 metros acima do nlve! de preia-mar actual. Nos nive is basais, bastante eimentados, identifieou-se uma assoei~o aut6etone corn fo!3.dios, dominada peJo biva lve Myo[orceps arislala (Dillwin, 1811). As areias marinhas incluem concentrac5es de beach drif/, compreendendo conchas bcm preservadas de gastr6podes supra-litorais e litorais, entre os quais numerosos rissoldeos. Nos bivalves predominam coneentrar,:Oes de valvas desarticuladas de Ervifia castanea (Montagu, 1803), uma pequena especie infaunal eolonizadora de substratos arenosos m6veis. Na composir,:ao total da fauna intervem esseneialmcnte especies do litoral cia Europa Ocidental, muitas delas existcntes na costa portuguesa. Todavia, estilo tambem presentcs algumas especies exoticas, "quen tes", caracleristicas das faunas de moluscos da costa ocidental de Africa ou das Caralbas. Este elenco aprescnla semelhanr,:as significativas corn diversas associar,:3es sub-f6sseis encontradas em dep6sitos plistocenicos do Mediterrineo Ocidental, considerados coma "tirrenianos".

ABSTRACT

Key.words: Pleistocene; marine invertebrates; Macaronesia; paleoecology; paleobiogeography.

The sub· fossil fauna from the Late Quaternary marine deposits of Santa Maria is made of more than SO species of gastropods and bivalves, 19 of them collccted recently and for the first lime in the northern coasl of the island (Lagoinhas Bay). The sub-fossil shells are found in deposits of beach sands, situated 2-3 mClers above the present low tide. The carbonated saT!ds from the basal part of the succession yield an autochthonous association of borers domina!ed by the bivalve Myo[orceps aristala (Dillwin, 18 17). Upwards, the marine sands contain concentrations of beach dri ft shells, inclUding well -preserved supratidal and intertidal gastropods, among them a large number ofRissoidae. The bivalve fauna is dominated by disarticulated valves of Ervi/ia castanea (Montagu, 1803), a small infauna! coloniser of mobile sandy substrates. The composition of the fauna is made essentially of West European species, many of them common to the West Coast of Portugal. However, a few "wann guests" wi th West African or Caribbean affinities were also found, suggesting a close relation with some of the ''Tyrrhenian'' wann associations found in the Western Mediterranean.

INTRODUCTION described near the Bay of Prainha, a few kilometers eastward Vila do Pprto, on the SOU lhem coast of the The Pleistocene marine deposits from Santa Maria are island (fig. I). After Zbyszewski & Ferreira (op. eit.), known for a long time, after the works of Mayer (l864) the succession observed near Prainha is situated and Bertho is (1951, 1953), revised by Zbyszewski about 2· 3 meters above the present low tide, and built & Ferreira ( 1961) and Zbyszewski et al. ( 1961). of a basal c onglomerate wi th algal.limcstone Some of these deposits proved to be exceptional sites to concreti ons followed by 0.80 to 1.50 m of fossilifcrous collect sub·fossil shells, with special emphasis to those beach sands.

313 1° Congresso sabre 0 Cenoz6ico de Portugal

bryozoans, decapods, Balanus sp. and the sea urchin Sphaerechinus granuiaris (Lamarck, 1822). The collected species are (pI. 11):

Classe Gastr-opoda

Fam. Haliotidae Haliolis luberculala Linne, 1758

Fam. Patellidae Patella ulyssiponensis Gmelin, 1791

Fam. T rochidae 9_2bn CalliosJoma zizyphinum (Linnc, 1758) • Jujubinus exasperatus (Pennant, 1777) Fig. I • Simplified map of Santa Maria, showing the location of Jujubinus strialus (Linne, 1758) the upper Pleistocene sites ofPraia, Prainha and Lagoinhas. Fam. Littorinidae Melarhaphe neritoides (Linne, 1758) In the northern coast of Santa Maria, a new deposit correlative with Prainha has been identified and sampled, Fam. supplying several hundreds of sub-fossil shells, mostly Alvania cancel/ala (da Costa, 1778) micro-gastropods. This deposit is best exposed near the Alvania cimicoides (Forbes, 1844) cliffs ofLagoinbas, between Ponta do Norte and the islet Cingula cingi/lus (Montagu, 1803) of Lagoinhas. Near this point, the succession begins with Manzonia unifasciata DaulZemberg, 1889 cemented and bored carbonated sands, overlaid by O.sOm Rissoa simi/is Scacchi, 1836 of beach sands with concentrations of shell debris, and a cover of torrential sediments (pI. I). Fam. Rissoinidae As in Prainha Bay, the fossiliferous beds yield an Zebina vi/rea A. Adams, 1854 abundant and diversified fauna of sub-fossil gastropods and bivalves, most of them belonging to small-sized Fam. Verrnetidae species. Besides its systematic and ecological relevance, Vermetus sp. this fauna is of special interest for biogeographic purposes due to the particular iocation of the Azores in the central Fam. Cerithiidae Atlantic. The bulk of the Quaternary marine mollusks from Bittium reticulatum (da Costa, 1778) this archipelago of Macaronesia show strong West European affinities (Morton, et al., 1998), with many Fam. Fossaridae species being common to the West Coast of Portugal Fossarus ambiguus (Linne, 1758) (Nobre, 1930, 1940). However, some West African and Caribbean influences can be noted by the introduction of Fam. Thaididae warm·guests in the sub-fossil faunas, suggesting the Thais haemastoma (Linne, 1767) occurrence of positive fluctuations in the mean temperatures of the Atlantic surface waters, during late Fam Coralliophilidae Pleistocene interglacial events. Coralliophila meyendorffi (Calcara, 1845) This account pretends to summarise the paleontologic data carried out after Mayer (1864) and Zbyszewski & Fam. Buccinidae Ferreira (1961), together with a comparison with a new Cantharus variegalus (Gray, 1839) collection of specimens recently collected in Santa Maria and housed in the Earth Sciences Departmcnt of the Fam. Columbellidae University of Coimbra (DCTUC). The same faunas are Mitrella broderipii (Sowerby, 1844) analysed in the point of view of their paleoecologic, paleobiogeographic and climatic significance. Fam. Cymatiidae Charonia lampas (Linne, 1758)

FAUNAL COMPOSITION Fam. MargineUidae Gibberula sp. The systematic analysis of the newbulk sample was concluded with thc identification of26 species of marine Fam Turridae gastropods and 5 bivalves. Mixed with the shell debris Mange/ia costata (Donovan, 1803) were also found a few fragmcnts of indeterminable erect Mangelia sp.

314 Ciincia$ da Terra (UNL), 14

Fam. Conidae Rissoa variabilis (Muhlfeldt, 1 824) Conus ermineus BOrn, 1778 Rissoa dolium Nyst, 1845 Rissoina bruguierei (pyraudeau, 1826) [referred Fam. Triphoridae as eanariensis d'Orbigny, 1839] Monophorus perversus (LiMe, 1758) Polinices lacteus (Guilding, 1834) Zonaria pyrum (Omelin, 1791) Fam. ElIobiidae Phalium undulatum (GemHn, 1791) Ovate/la (Mysotel/a) myosotis (Drapamaud, Cabestana cutaeea (Linne, 1767) 1805) Mitra comieula Linne, 1758 Mangelia nebula (Montagu, 1803) C1asse 8ivalvla Cerilhiopsis nalla JeffreyS, 1867 SeUa trilineala (Philippi, 1836) Fam Arcidae Area tetragona Poli, 1795 Classe Bivalvia

Fam. Mytilidae Lithophaga lithophaga (Linne, 1758) [probably a MyoJorceps aristata (Dillwin, 18 17) MyoJoreeps aristata (Dillwin, 1817)] Neopycnodollte cfr. cochlear (poli, 1795) Fam. Pectinidae Lyropecten efr. nodosus (LiMe, 1758) Chlamys distorta (da Costa, 1778) Laevicardium norvegieum (Spengler, 1797) Solen efr. marginatus (pennant, 1777) Fam. Carditidae Cardita calyculata (Linne, 1758) With the contribution of these 22 taxa the diversity of the invertebrate fauna from the Pleistocene of Santa Maria Fam. Mesodesmatidae reaches 57 species, most of them gastropods (43) and Ervilia castanea (Montagu, 1803) bivalves (11). However, this systematic list should be reviewed in some aspects, with special attention to the A comparison with the systematic lists already known micro-shells (Rissoidae and others), because there are from the researches of Mayer (1864) and Zbyszewski & scvera1 species mentioned by Mayer ( 1864) and never Ferreira (1961) shows that 19 of the taxa are, for the fIrst collected again. The ~ssoidae and Rissoinidae, they were time, referred to the Pleistocene deposits of Santa Maria. object of detailed reviews in the last decades, and some of These taxa are; Calliostoma zizyphinum (Li~e, 1758), the taxa listed by this worker are Mediterranean species Jujubinus exasperalus (Pennant, 1777), Jujubinus striatus without distribution in the adjacent Atlantic areas. (Linne, 1758), Melarhaphe neritoides (Linne, 1758), The borer bivalve Lithophaga lithophaga (Linne, Alvania caneelfata (da Costa, 1778), Cingula cingillus 1758) described by Zbyszewski & Ferreira (1961) as one (Montagu, 1803), Manzonia unifaseiata Dautzemberg, of the dominant elements from the fauna of Prainha is 1889, Zebina vilrea AAdams, 1854, Vermetus sp., Mitrel/a also a typical Mediterranean species. The specimens of broderipii (Sowerby, 1844), Charonia lampas (Linne, borers from the recent collection of the ocruc belong to 1758), Gibberula sp. Mangelia costata (Donovan, 1803), MyoJoreeps aristOla (Dillwin, 18 17), which is a close Mangelia sp., Conus ermineus Born, 1778, Ovatella species from the West European and East American coasts. (Mysotella) myosotis (Draparnaud, 1805), Area tetragona It can be differentiated by its small size, together with two Poli, 1795, Myoforeeps aristata (Dillwin, 18 17) and pointed and curved tips'in the posterior ends of the valves. Chlamys distorta (da Costa, 1778). These tips are difficult to observe in the sub-fossil shells, The taxa previously mentioned by the same workers since most of them are strongly incrusted in the carbonate but not collected this time are; concretions. The limpet Patella eaerulea Linne, 1758 listed by Zbyszewski & Ferreira (1961) is surely the macaronesian Classe Patella eandei d'Orbigny, 1840, an endemic species from Azores, Madeira and Canaries. On the contrary, the recent Patella eaerulea Linne, 1758 [Palella candei distribution of Patella caerulea is restricted to the d'Orbigny, 1840] Meditcrranean (Poppe & Goto, 1991). The occurrence of Palella vulgata Linne, 1758 [probably a Patella vulgala Linne, 1758 also needs confmnation since Patella eandei d'Orbigny, 1840] the sub-fossil shells of this Northwest European limpet Gibbula eineraria (Linne, 1758) [referred as can be confused with young specimens of the highly "Trochus "strigosus Omelin, 1791] variable Patella candei. Tricolia pullus (Linne, 1758) The Conidae from the fauna ofPrainha were compared Manzonia erassa (Kanmacher, 1798) [referred by M. Brebion (in Zbyszewski & Ferreira, 196 1) with the as costata Adams, 1796] West African Conus ambiguus Reeve, 1844 and Callus Alvania pagodula (8ucquoy, Dautzenberg & testudinarius Hwass, in Bruguiere, 1792. However, DoJl fus, 1884 ) "Iestudinarius" is a synonymous frequently used to

315 I- Congresso sob£e 0 Cenoz6ico de Ponugal designate the Eastern Atlantic populations of Conus of mobile sandy substrates, through the growth of dense ennineus Born, 1778 (Korn, 1987). The morphology of populations in the shallow sandy bars that surround coastal these shells, including the specimcn recently housed in cliffs (Morton, et aI., 1998). The shell debris from this the collection of the DCTUC, is closer to weakly developed species contribute with a significant volume to the neigh­ forms of the amphiatlantic Conus ennineus. After Bumay bourhood beach sands, being one of the most common & Monteiro (1977) and Walls (1979) these diagnostic elements of the fossil associations found on the Pleistocene features include a shoulder roundly angled, a low to deposits. moderate spire with 2-3 spiral ridges and very week and Figure 2 summarises an ewlogical section representing few axial threads, and a body whorl wi th 6-10 low spiral the sub-fossil marine molluscs from Santa Maria. ridges restricted to the area above the base. Conus ambiguus can be easily .(fistinguished because of the pattern of the spire, which has sometimes a cancellate PALEOBIOGEOGRAPHIC AND CLIMATIC appearance. Usually the shoulder is more roundly and the SIGNIFICANCE body whorl is irregularly spirall y sculptured. Just as the recent marine fauna of molluscs from the Azores, most species from the sub-fossil fauna of Santa BIOSTRATINOMIC AND PALEOECOLOGIC Maria are common to the Atlantic coasts of Western FEATURES Europe, from the Channel to Portugal. Some of the known ranges of these species also reach the Western The sub-fossil malacofaunas from Prainha Bay and Mediterranean, the coastline of Morocco, and the Madeira Lagoinhas are the result of an heterogeneous mixing of and Selvagens islands. Nevertheless, the particular shells with distinct grades of abrasion, fragmentation and interest of Pleistocene malacofauna from Santa disarticulation, belonging to species typical of rocky, sandy Maria arises from the existence of West African and or algal substrates from the supralinoral to upper sublittoral Caribbean warm-guests, today unknown from the marine environments. Two different types of concentra­ Azores waters. ti ons can be defined: Cantharus variegatus (Gray, 1839) is a tropical West (I) Autochthonous concentrations of borer and cryptic African species, reaching today the Canaries islands. bivalves found articulated in sandy carbonated substrates, During the Pleistocene its northern range area was strongly cemented and partly of algal origin. prolonged to North, into the littoral of Morocco (2) Parautochthonous concentrations of gastropods and (Harounian and Ouljian ofLecointre, 1963, and Brebion, bivalves, including some mixed allochthonous elements 1979) and the Mediterranean (Tyrrhenian 11 and III of represented by abraded and fragmented shells of sublittoral Charrier, 19.6 1 and Dieny & Massari, 1966). species. These concentrations can be interpreted as the Conus ermineus BOrn, 1778 is a widespread tropical result of beach drift accumulated during high tides, amphiatlantic species, common on the West African coasts especially during storm events or equinoctial tides. from the Sahara to Angola (Walls, 1979), but also on the The upper intertidal biota of exposed rocky substrates Brazilian coast,Antilles, Central America, Gulfof Mexico is represented by Melarhaphe neritoides (LiMe, 1758) and and Bahamas. As in the previous species, several occur­ Ovalella (Mysotella) myosolis (Draparnaud, 1805). rences are known from the Tyrrhenian of the Western and Many other species from this fauna are typical Central Mediterranean, together with the classical elements of middle or lower intertidal shores. The gastro­ "SJrombus bubonius" and Mytilus senegalensis warm pods Patella ulyssiponensis Gmelin, 1791, Calliostoma faunas (Charner, 1961; Vazzana, 1988). zizyphinum (Linne, 1758), Vermelus sp ., Bittium Polinices lacteus (Guilding, 1834) is one of the best reticulatum (da Costa, 1778), and the bivalves Myoforceps examples of tropical amphiatlantic gastropods, ranging arislala (Dillwin, 1817) and Cardita calyculata (Linne, from Mauritania, Madeira and Canaries to Angola and 1758) inhabit some of these hard intertidal substrates colo­ Santa Helena, and from the Brazilian coast to the Carib­ nised with algae and sponges. Many of the small Rissoidae bean (Abbott, 1968; Dixon & RyaU, 1986; Gofas et al., listed above are herbivorous feeders that can be found in 1987; Poppe & Goto, 199 1). This naticid is occasionally lower intertidal or upper sublittoral algal prairies and mats. found in the Azores (Morton, et al. , 1998). It is also cited The algal and rocky bottoms from the upper part of in the Mediterranean Pleistoccne, together with the the sublittoral stage are represented by the gastropods previous species and Strombus faunas. Halioris tuberculata Linne, 1758, Jujubinus exasperatus The small Zebina vitrea A. Adams, 1854 is a warm (Pennant, 1777), J. stria/us (Linne, 1758), CtJralliophila temperate North-Atlantic gastropod, known 'from the meyendorffi (Calcara, 1845), Cantharus variegalus (Gray, Antilles and also found in the Canaries and Selvagens 1839), Mitrella broderipii (Sowerby, 1844), Charonia (Poppe & Goto, 1991). lampas (Linne, 1758), Gibberula sp., Mangelia costata Lyropecten nodosus (Linne, 1758) is a West Atlantic (Donovan, 1803), and Conus ermineus Born, 1778. pectinid that range from the Brazilian coasts to the Occasionally, the same species can be found in intertidal Caribbean and the Southeast littoral of USA (Abbott, rocks. . 1968). The small but very abundant infaunal bivalve Ervilia This association of wann-temperate species suggcsts castanea (Montagu, 1803) is a very successful coloniser that the 2-3 meters marine deposits of Santa Maria can be

316 Ciblcias da Terra (UNL), 14

317 1° Congreiso sobre 0 Cenowieo de Portugal related with a Pleistoeene warm period, marked by the The geographic range of the species from the migration northwards of species traditionally found in Pleistocene of Santa Maria also emphasises the particular lower latitudes. A comparison with the Mediterranean importance of the Azores archipelago, as a crossing and classical faunas, containing "Strombus bubonius" and mixing area between the Western and Eastern Atlantic other sub-tropical components, seems evident. Both faunal molluscan faunas. During warmer interglacial events, the associations from Santa Maria and from the Mediterranean positive oscillations in the mean temperatures of the yield Cantharns variegatus, Conus ermineus and Polinices surface waters promoted migrations of warm species lacteus. In the "Tyrrhenian" deposits of Italy, Southeast northward, reaching the Azores for some times. From Spain and Canaries (Brebion, 1979), these species can be the Caribbean Province, a few other species with collected together with "Strombus bubonius", Mytilus prolonged veliger development were also able to extend senegalensis, Cardila senegaiensis, and other molluscs its geographic range eastwards, colonising the littoral of of warm-temperate waters. Santa Maria and other of the Central Atlantic shallow areas.

REFERENCES

Abbott, R. (1968) - Seashells of North America, Golden Press, New York, 280 p. Berthois, L. (1951) - Sur la presence de basses terrasses marines dan I' Archipel des Avores. Comptes Rendus du 76'- Congres des Societes Savantes a Rennes, 10[-106. Berthois, 1. (1953) - Contribution it I'etude lithologiquc de l'Arehipel des Acores. Comunicar;i5es dos Servir;os Geol6gicos de Portugal, 34,198 p., 13 pIs. Brebion, P. (1979) - Etude biostratigraphique et paleoecologique du Quaternaire marocain. Anna/es de Paleantologie (Inverlcbres), 65: 1-42. Bumay, 1. & Monteiro, A. (1977) - Scashclls from Cape Verde Islands. edition of the authors, Lisbon, 88 p., 66 figs. Charrier, G (1961) - Nuove osselVazioni sui Tirreniano di Cala Liberolto (Regione Sos Alinos) nel Golfo di Orosei (Sardegna Centro-Orientale). Bolleltino del Sen'izio Ge%gico d'Italia, 81: 557-580. Dicny, I. & Massari, F. (1966) - 11 Neogene e il Quaternario dei Dintorni di Orosci (Sardegna). Memorie del/a Societa Ilaliana di Sciellze Naturali e del Museo Civico i Storia Natura/edi Milano, IS: 123- 141, pI. 16. Dixon, M. & Ryall, P. (1986) - Naticidae of West Africa, part 11. La Conchiglia, 202-203: 4-5, 8-11. Korn, W. (1987) - COIIUS ermineus Born, an amphiatlantic species? La Conchiglia, 2 16-217: pp. 4-5. Lccointre, G (1963) - Rechcrchcs sur le Ncogene et le Quatemaire marins de la cOle atlantique du Maroc. Notes et Memoires du Sen'ice Geologique du Marac, 174: 75 p. Mayer (1864) - Palaontologische Verhaltnisse. Systematischcs Vcrzcichniss dcr fossilen Resle von Madeira, Porto Santo und Santa Maria nebst Beschreibung der neuen Arten. In Hartung, G (1864) - Geologische Beschreibung der Inseln Madeira und Porto Sanlo. Leipzig: 183-285,8 pis. Monon, 8.; Brilton, J. & Martins, A. (1998) - Ecologia Costeira dos Ar;ores. Ed. Sociedade Afonso Chaves, Ponta Oelgada, 249 p. Nobrc, A. (1930) - Materiais para 0 estudo da Fauna dos A(:oret. Instituto de Zoologia da Universidade do Porto, 108 p. Nobre, A. (1940) - Fauna malaco/Ogica de Portugal, 1- Moluscos marinhos e das aguas sa/abras. Companhia Editora do Minho, Porto, 806 p., 87 pIs. Poppe, G & GOIO, Y. (199 1) - European Seashells, vol. I (Po/yplacophora, Caudojoveala, So/cnogastra, Gastropoda). Verlag Christa Hemmen, 352 p., 40 pis. Zbyszewski, G & Ferrcira, O. (1961) - La faune marine des basses plages quatemaires de Praia et Prainha dans l'iJe de Santa Maria (A ~ores). Comunicar;i5es dos Servir;os Geol6gicos de Portugal, 45: 467-478.

Zbyszewski, G; Ferreira, O. & Assun~ao, C. (1961) - Carta Geol6gica de Portugal, na escala 1/50000; Nodcia explicativa da folha de Ilha de Santa Maria. Sen'ir;os Geo/Ogicos de Portugal, Lisboa, 28 p. Vazzana, A. (1988) - The shells of the Tyrrhenian period around Rcggio Ca labria (Italy). La Canchiglia, 234-235: 25-27. Vaught, K. (1989) - A classification o/the living . American Malaeologists Inc., Melbourne, USA, 190 p. Walls, J. (1979) - Cone Shells. A synopsis o/the living Canidae. T. F. H. Publications, Hong Kong, 1011p.

318 Ciellcias da Terra (UNl.), 14

Plate I

Fig. I - The northern eoast of Santa Mafia, near the bay and islet of Lagoinhas, with the 2-3 meters platform exposed at low tide.

Fig. 2 - Detailed view of the Upper P1cistcx:ene fossiliferous sands.

319 1° Cangressa sabre 0 Cenoz6ico de Portugal

Plate U

1, 2 - Thais haemastoma (Linne, 1767), back and aperture views. (xl,S)

3 - Melarhaphe neritoides (Linne, 1758), aperture view. (x2)

4 - Calliostoma zizyphinum (Linne, 1758), aperture view of incomplete specimen. (xl,5)

Sa-b - Mangelia costala (Donovan, 1803), back and aperture views. (x5)

6 - Biuium reticulatum (da Costa, 1778), back view. (x5)

7 - Myojorceps aris/ala (Dillwin, 1817), bored algal limestone with articulated shell. (x I ,5)

8, 9 - Ervi/ia castanea (Montagu, 1803), left and ri ght valves. (x5)

10, 11 - Cardita calyculata (Linne, 1758), left and right valves. (x5)

12a-b - Mangelia sp., back and aperture views. (x5)

13a-b - Fossarus ambiguus (Linne, 1758), back and aperture views. (x5)

14a-b - Conus ermineus Born, 1778, back and aperture views. (x 1,5)

lSa-b - Cantharus variegatus (Gray, 1839), back and aperture views. (xl,S)

16 - Haliotis tuben;ulata Linne, 1758, external view. (x 1,5)

17 - Patella ulyssiponensis Gmelin, 1791, fragment ofa large shell with a colony of Vermetus sp. (xl,S)

18a-b - Charonia Jampos (Linne, 1758), back and aperture views of young specimen. (x 1)

320 Cienciasda Terra (UNL), 14

Sa

2

6

l2a l2b

9

13a

14a 14b 15a 15b

321