journal of african earth sciences - sadabe

Journal of African Earth Sciences 100 (2014) 409–417

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Journal of African Earth Sciences

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Miocene benthic foraminifera from Nosy Makamby and Amparafaka,Mahajanga Basin, northwestern Madagascar

http://dx.doi.org/10.1016/j.jafrearsci.2014.07.0131464-343X/� 2014 Published by Elsevier Ltd.

⇑ Corresponding author. Tel.: +261 330492390.E-mail address: [email protected] (T.N. Ramihangihajason).

Tolotra N. Ramihangihajason a,⇑, Tsiory H. Andrianavalona a, Rachel Razafimbelo b, Lydia Rahantarisoa b,Jason R. Ali c, Karen E. Samonds d

a Département de Paléontologie et d’Anthropologie Biologique, Faculté des Sciences, Université d’Antananarivo, BP 906, Antananarivo, Madagascarb Département des Sciences de la Terre, Faculté des Sciences, Université d’Antananarivo, BP 906, Antananarivo, Madagascarc Department of Earth Sciences, University of Hong Kong, Hong Kong, Chinad Department of Biological Sciences, Northern Illinois University, DeKalb, IL, USA

a r t i c l e i n f o a b s t r a c t

Article history:Received 22 March 2014Received in revised form 20 July 2014Accepted 21 July 2014Available online 31 July 2014

Keywords:ForaminiferaMioceneNosy MakambyAmparafakaMadagascar

Madagascar is well known for its fossil deposits and hosts one of the world’s most important UpperCretaceous terrestrial faunal sites (in the Mahajanga and Morondava Basins in the west and northwestof the island). Cenozoic marine fossils are also described from Madagascar, but these have received farless attention from the paleontological community, with most of this work dating from the 19th and early20th centuries. Our study reports a new comprehensive microfossil assemblage from a Miocene sequenceon the island of Nosy Makamby. After washing, sieving and sorting (�30 kg), twenty-five genera offoraminifera were identified including Alveolina, Ammodiscus, Ammonia, Archaias, Bolivina, Borelis,Cassidulina, Cyclammina, Cycloforina, Dentalina, Elphidium, Hauerina, Lagena, Lepidocyclina, Nodosaria,Nonion, Nonionella, Peneroplis, Pyrgo, Quinqueloculina, Rhabdammina, Spirillina, Spirolina, Spiroloculinaand Triloculina. Ostracods are found in association with the foraminifera, as well as many other macroin-vertebrate fossils (including bivalves, gastropods, and echinoids) in addition to vertebrate fossils.Together, the assemblage indicates that during the late Miocene, Nosy Makamby was a tropical,near-shore environment, probably similar to that seen today. Furthermore, the existence of epiphyticforaminiferans (e.g., Elphidium) suggests that sea-grass beds were likely present.

� 2014 Published by Elsevier Ltd.

1. Introduction

In Madagascar, fossil foraminifera remain relatively poorlyknown, with nearly all published studies describing Paleozoicand Mesozoic forms (e.g., Besairie and Collignon, 1971; Ujiie andRandrianasolo, 1977). This gap in knowledge reflects the fact thatMadagascar’s fossil record is nearly entirely constrained to eitherthe Jurassic/Late Cretaceous (Krause et al., 2006) or the recent LatePleistocene/Holocene (extending back a mere 80,000 years;Samonds, 2007). This critical gap spanning most of the Cenozoichas remained largely unknown; this time interval is particularlysignificant because this is when the ancestors of the modernMalagasy forms are thought to have arrived (Samonds et al.,2012, 2013).

Nearshore marine carbonate deposits from Madagascar’sCenozoic are mapped, but few have been systematically explored.Previous studies include a few published reports from the Eocene

and Miocene, listing both invertebrate (foraminifera, bivalves,gastropods, echinoids) and vertebrate fossils. Vertebrates includebony fish, sharks, rays, turtles, crocodylians, and mammals, includ-ing a primitive dugongid skull and ribs belonging to the Malagasyseacow Eotheroides lambondrano (Collignon and Cottreau, 1927;Samonds et al., 2009; Andrianavalona, 2011).

Historically, Miocene rocks were first described from westernand northwestern Madagascar (Perrier de la Bathie, 1921;Collignon and Cottreau, 1927; Hourq, 1949; Besairie, 1956;Besairie and Collignon, 1971). Perrier de la Bathie produced the firstdetailed work describing the age of these post-Cretaceous forma-tions, and published a geologic section of Nosy Makamby and CapTanjona, two of the best known Cenozoic localities (Perrier de laBathie, 1921). The largest and most comprehensive work to dateis that of Collignon and Cottreau (1927); their work is the founda-tion for our current understanding of the paleontological andgeological history of Nosy Makamby. Further west, Baron andMouneyres (1904) were the first to publish a comprehensive geo-logical description for Amparafaka. However, this region remainspoorly described. Some authors have noted the ‘‘occurrence of

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Fig. 1. Map of the study region.

410 T.N. Ramihangihajason et al. / Journal of African Earth Sciences 100 (2014) 409–417

Miocene deposits’’, without any specific locality information(Collignon and Cottreau, 1927; Besairie and Collignon, 1971).

A preliminary list of foraminiferan fossils and their associatedinformation was first described by Lemoine and Douvillé (1904);this study and subsequent work has focused largely on the genusLepidocyclina (a large benthic foraminiferan within the Family Orb-itoïdidae; Lemoine and Douvillé, 1904; Douvillé, 1908; Beretti,1973). Collignon and Cottreau (1927) mention the occurrence ofonly one species (Miogypsina irregularis), and Lavocat published asimple description of the foraminiferans (Lavocat et al., 1960).Despite a few other species listed by Besairie and Collignon(1971), the microfauna of Madagascar’s Miocene remains incom-plete and poorly understood.

Here, we report the first detailed study of Miocene microfossilsfrom Madagascar, some of which are reported from the island forthe first time. In contrast to the relative paucity of MalagasyCenozoic vertebrate fossils, foraminiferans are frequently well-preserved and recovered in high abundance, representing anuntapped source of valuable paleoecological information. Thisinformation has great potential to elucidate this virtually unknowntime period in Madagascar’s past history.

2. Geological setting

The study region is located within the Mahajanga Basin,northwestern Madagascar (Fig. 1). This region contains the mostcomplete marine sedimentary section of Miocene rocks fromMadagascar, with lateral extensions that outcrop in the regionsof Nosy Makamby, Cap Tanjona, Cap Sada, and Amparafaka(Collignon and Cottreau, 1927; Samonds, personal observations).We report here microfossils from two localities: the island of NosyMakamby, and the coastal region of Amparafaka (Fig. 1).

2.1. Nosy Makamby

Nosy Makamby (or Mahakamby1) is a small island (about1.6 km � 0.4 km) SSW–NNE aligned offshore at the broad part of

1 Both names are used in the literature.

the delta at the mouth of the Mahavavy River, in the north-westernpart of Madagascar, approximately 50 km west along the coast fromthe regional capital of Mahajanga (Andrianavalona, 2011; Figs. 1 and2). The island’s sediments are mostly medium to fine-grained calcar-eous sandstones that accumulated in a coastal near-shore marineenvironment, overlain by a Pliocene continental unit comprise ofred beds and quartz grains, etc. (Collignon and Cottreau, 1927;Besairie, 1972; Fig. 3); these units are approximately 15 m thick.One horizon is particularly rich in fossilized Kuphus tubes, whichclearly demarcates the sediment horizons between exposures onthe northern region of the island (Fig. 2). In addition to foraminifer-ans, a diverse assemblage of both invertebrates and vertebrates havebeen collected from Nosy Makamby, including bivalves, gastropods,echinoids, crabs, bony fish, sharks, non-diagnostic reptiles (turtlesand crocodylians), and sirenian mammals (Collignon and Cottreau,1927; Besairie, 1972; Samonds et al., 2007).

2.2. Amparafaka

Within the Baly Bay, Amparafaka is the most southern of thesites (Fig. 1); no fossils have previously been reported from thisregion. Like Makamby, Amparafaka is characterized by steep cal-careous cliffs covered by red continental Pliocene sandstones.These deposits are interpreted as lateral extensions of thosereported at Makamby (Collignon and Cottreau, 1927).

3. Material and methods

3.1. Sampling and processing

In June and July 2010, 47 rock/sediment samples were collectedfrom Nosy Makamby and Amparafaka. Samples were washed overa succession of four sieves (2 mm, 1 mm, 0.5 mm, 0.2 mm), andresidue was examined for foraminiferal contents using the meth-odology described in Neumann (1967). A USB microscope (VEHOVMS-001) was used to photograph specimens. Species distribu-tions and stratigraphic ranges were compiled for each genus (seeLuczkowska, 1974; Murray, 2006). Identification of genera wasmade using descriptions and comparative material following

Fig. 2. Two photographs of Kuphus tubes, a genus of shipworm (marine bivalve molluscs) within the family Teredinidae.

Fig. 3. Geological section of Nosy Makamby.

T.N. Ramihangihajason et al. / Journal of African Earth Sciences 100 (2014) 409–417 411

Loeblich and Tappan (1964, 1988). A conservative approach wasused when identifying specimens (e.g., Family, Order, or Classonly), as a precise determination was not always possible (seeBignot, 1982).

4. Results

We report here 25 genera of benthic foraminifera from theMiocene of Madagascar collected from both sites (Amparafakaand Nosy Makamby). Together, this assemblage suggests that thesediments from Makamby and Amparafaka were deposited inwarm tropical shallow water. Indeed, almost all the sedimentsfrom both sites present a very clear association of shallow waterforms. Considerable epiphyte genera support the existence ofseagrass.

Genera recovered include the following: Quinqueloculina,Cycloforina, Hauerina, Triloculina, Pyrgo (Family: MILIOLIDAE); Spi-roloculina (Family: NUBECULARIDAE), Borelis, Alveolinella (Family:ALVEOLINIDAE), Peneroplis, Spirolina, Archaias (Family: SORITIDAE),Cyclammina (Family: LITUOLIDAE), Ammodiscus (Family: AMMO-DISCIDAE), Rhabdammina (Family: ASTRORHIZIDAE), Ammonia

(Family: ROTALIDAE), Elphidium (Family: ELPHIDIDAE), Nonion,Nonionella (Family: NONIONIDAE), Cassidulina (Family: CASSIDU-LINIDAE), Lagena, Dentalina, Nodosaria (Family: NODOSARIIDAE),Bolivina (Family: BOLIVINITIDAE), Lepidocyclina (Family: LEPIDO-CYCLINIDAE), Spirillina (Family: SPIRILLINIDAE). In addition to theforaminiferans recovered, ostracods are also found, as well as othermacroinvertebrate and vertebrate fossils including bivalves, gas-tropods, echinoids, bony fish, sharks, turtles, crocodylians, andsirenian mammals.

Miliolids are the most abundant form in this association; theyrepresent more than 50% of the foraminiferans collected in thisstudy. No planktonic foraminiferans were recovered.

5. Discussion and conclusions

5.1. Faunal diversity and biogeography

Foraminifera are valuable paleoenvironmental indicators due tothe fact that some species preferentially inhabit different environ-ments. Specifically, the proportion of planktonic species (floating inthe upper layers of the ocean waters) to benthic species (those that

Table 1Paleoenvironmental characteristics of Miocene foraminiferal genera found at Nosy Makamby and Amparafaka, Mahajanga Basin, Madagascar (adapted from Murray (2006)).

Genera Temperature Salinity Depth Substrate Zonation

Alveolinella 18–26 �C Normal marine 5–100 m Algal-covered carbonate gravels Inner shelf, lagoonAmmonia Warm temperate–tropical Brackish to

hypersaline0–50 m Muddy sand Inner shelf, lagoon

Archaias More than 22 �C Normal marine 0–20 m Phytal Inner shelfBolivina Cold–warm Normal marine – Muddy sediment Inner shelf, bathyalBorelis 18–26 �C Normal marine 0–40 m Algal-coated surface, seagrass and coarse

sedimentLagoon, reef

Cassidulina Cold–temperate Normal marine – Mud, sand Shelf–BathyalCyclammina – Normal marine >100 m Mud, sand Outer shelf, abyssalElphidium Warm to temperate 0–70 g/L 0–50 m Sand, vegetation Inner shelfNonion Cold–warm 30–35 g/L 0–180 m Mud, silt ShelfNonionella Warm to temperate Normal marine 10–

1000 mMud Shelf–upper bathyal

Peneroplis 18–27 �C 35–53 g/L 0–70 m Plants and hard substrates Lagoons and innermost shelfPyrgo Temperate–warm Normal marine – Plants or sediment Shelf–bathyalQuinqueloculina Cold–warm 32–65 g/L – Plants or sediment Hypersaline lagoons, marine marsh and

shelfSpirillina Cold–temperate Normal marine 0–100 m Hard substrates Inner-mid shelfSpirolina 18–26 �C 37–50 g/L – Plants Lagoons, nearshoreSpiroloculina Temperate–warm Marine–

hypersaline0–40 m Sediments or plants Lagoons, inner shelf

Triloculina Temperate–warm, cold (bathyalspecies)

32–55 g/L – Mud, sand, plants Hypersaline lagoons or inner shelf, somebathyal


inhabit the sea bed or just beneath the surface) is especially infor-mative – from 0% planktonic forams in shallow marine waters tomore than 90% in deep marine. The absence of any planktonicforaminiferans in our Miocene localities is interesting, and sug-gests a shallow water environment. Future fieldwork and samplingmay be able to specify whether or not any planktonic foram fossilsare indeed present.

5.2. Paleoenvironment

As the distribution and abundance of foraminiferans is con-trolled by environmental factors, including bathymetry, sedimenttexture, and physiochemical characteristics of sediment and water(see Table 1), these organisms are a good proxy for both oceanicand climatic information (Devi and Rajashekhar, 2009).

5.2.1. SalinityThe foraminiferans from Makamby and Amparafaka do not

show considerable salinity variation during the Miocene. Indeed,foraminiferans recorded appear to prefer normal salinity environ-ments (euhaline) from 33 g/L to 37 g/L.

5.2.2. TemperatureMiliolids are abundant in almost all the deposits of Makamby

and Amparafaka. Distribution of the Miliolidae is generally notedin tropical regions and those of warm water all over the world(Luczkowska, 1974). Also, most of the genera in all the formationstolerate temperate to warm water (�18–27 �C; e.g., Peneroplis,Borelis).

5.2.3. SubstrateNature and granulometry of sediments in both Makamby and

Amparafaka are nearly constant (medium to fine-grained calcare-ous sandstones) in all sections. This suggests that the substratedid not exert much influence on composition changes of the ben-thic foraminiferal association. However, the nature of the substrateis often regarded as an important factor limiting the distribution ofbenthic foraminifera (Van der Zwaan, 1982). The occurrence of epi-phytic taxa (e.g., Elphidium, Peneroplis) suggests the existence ofseagrass (Margerel, 2009).

5.2.4. DepthThe dominance of Miliolids corroborates that deposits in both

regions were from the inner shelf. Quinqueloculina is the most com-mon genus (it represents nearly 50% of Miliolids in both regions);this further indicates a coastal environment (Luczkowska, 1974).However, some few deep-sea forms like Cassidulina, Cyclammina,Rhabdamina, were also recovered intermixed with inner-shelf gen-era. This may have resulted from an upwelling, which could alsoexplain the abundance of other invertebrate fossils seen in associ-ation with foraminiferans, or possibly reworking or bioturbationbetween layers, though no direct evidence was visible.

In Makamby, foraminiferans are present especially in the baseof the section. On the top three layers, foraminiferans are rare orabsent. Foraminiferans are also present throughout the entire for-mation at Amparafaka, but are also scarce at the top of the section.Further work is needed to determine the paleoenvironmental set-ting associated with these two sites.

This descriptive work represents the first comprehensivereport of microfossils from Madagascar’s Miocene. As relativelylittle is known about this time period in Madagascar (due to thepaucity of the Cenozoic fossil record), foraminiferans represent asignificant source of information, as they are often well-preservedand recovered in high density. The microfauna from Makamby isdominated by Miliolids, and suggests a tropical near-shorepaleoenvironment. This research was conducted in the island’snorthwest, but future studies from other regions will help betterunderstand the biostratigraphy and paleoenvironment of theisland.

5.3. Institutional and other abbreviations

UAP, Université d’Antananarivo, Antananarivo, Madagascar.

6. Systematic paleontology

Phylum PROTISTA Haeckel 1866Subphylum SARCODINA Schmarda 1871Class RHIZOPODA Von Siebold 1845Subclass GRANULORETICULOSA De Saedeleer 1934Order FORAMINIFERIDA Eichwald 1830Suborder MILIOLINA Delage & Hérouard 1896Superfamily MILIOLACEA Ehrenberg 1839


Family MILIOLIDAE Ehrenberg, 1839Subfamily QUINQUELOCULININAE Cushman 1917Genus QUINQUELOCULINA d’Orbigny 1826Quinqueloculina sp.Fig. 4; 1

Location: Makamby and Amparafaka.Material: UAP-10.553; UAP-10.561; UAP-10.564; UAP-10.567;

UAP-10.571; UAP-10.580; UAP-10.585; UAP-10.586; UAP-10.592;UAP-10.598; UAP-10.602; UAP-10.604; UAP-10.608; UAP-10.619;UAP-10.625; UAP-10.631; UAP-10.639.

Description: Test free, coiled and alternating regularly in 5planes. 3 Chambers visible from exterior of one side of test, 4 vis-ible from opposite side. Wall is calcareous, porcelaneous. Somespecimens show some agglutinated grain on exterior. Aperture isterminal, with simple (occasionally bifid) tooth.

Remarks: This genus is described from inner shelf and open gulfenvironments, in warm and shallow tropical and subtropicalwaters, from zero to �100 m. The genus is a good indicator of con-ditions in proximity to past coastlines (Luczkowska, 1974).

Stratigraphic range: Cretaceous to Holocene (Loeblich andTappan, 1988).

Genus CYCLOFORINA Luczkowska 1972Cycloforina sp.Fig. 4; 2

Location: Makamby.Material: UAP-10.571A; UAP-10.580A.Description: Test is porcelaneous, oval, nearly elongated and

protracted at both ends. Chambers are tubular, narrow and dis-tinctly curved at the basal end. Rolling is quinqueloculine typewith always five chambers visible externally. Aperture is at theend of a short narrowing neck, and possesses a short bifid tooth.

Remarks: Many species of Cycloforina are commonly found inassociation with Quinqueloculina species, typically occurring incoastal facies.

Stratigraphic range: Jurassic to Holocene (Luczkowska, 1972;Loeblich and Tappan, 1988).

Genus HAUERINA d’Orbigny 1839Hauerina cf. compressa d’Orbigny 1846Fig. 4; 3

Location: Makamby.Material: UAP-10.556; UAP-10.559.Description: Test is almost discoidal (flat and subcircular in out-

line) and concave in the central part. Early stages are quinquelocu-line. Later several chambers are planispiral, 3 or 4 in the last whorlaround the previous chambers, and covering the initial part of thetest. Sutures bent backwards, and depressed. Aperture is a narrowand oval, with trematophore.

Remarks: Many species of Hauerina are associated with middleneritic water depths or reef environments (Havach and Collins,1997).

Stratigraphic range: Eocene to Holocene (Hauerina; Loeblich andTappan, 1964); H. cf. compressa: Poland: Upper Tortonian (GliwiceStare, Weglinek). Austria: the vicinity of Vienna. Romania: Torto-nian, Buitur. ex-USSR: Upper Tortonian, the Volhyn-Podolian Plat-form (Luczkowska, 1974).

Subfamily MILIOLINELLINAE Vella 1957Genus TRILOCULINA d’Orbigny 1826Triloculina sp.Fig. 4; 4

Location: Makamby and Amparafaka.Material: UAP-10.553A; UAP-10.561A; UAP-10.564A; UAP-

10.567A; UAP-10.586A; UAP-10.598A; UAP-10.602A; UAP-10.608A; UAP-10.619A; UAP-10.639A.

Description: Test free, oval. Only three chambers are visibleexternally. Wall calcareous and porcelaneous. Aperture is terminalwith bifid tooth.

Remarks: Good indicator of coastal conditions. This genus pre-vails over the inner shelf, generally in association with Quinquelo-culina; neither of these genera are common in the region of centraland outer shelves (Luczkowska, 1974).

Stratigraphic range: Middle Eocene to Holocene (Loeblich andTappan, 1988).

Genus PYRGO Defrance 1824Pyrgo sp.Fig. 4; 5


UAP-10.565; UAP-10.568; UAP-10.572; UAP-10.581; UAP-10.599;UAP-10.615; UAP-10.624; UAP-10.626; UAP-10.632.

Description: Test free. Only two chambers visible externally.Aperture is terminal with bifid tooth.

Remarks: Pyrgo contains various groups of species in the bathyalzone, specifically where other Miliolidae are scarce. However, thisgenus occurs also in the inner shelf (Luczkowska, 1974).

Stratigraphic range: Upper Eocene to Holocene (Loeblich andTappan, 1988).

Family NUBECULARIIDAE Jones 1875Subfamily SPIROLOCULININAE Wiesner 1920Genus SPIROLOCULINA d’Orbigny 1826Spiroloculina sp.Fig. 4; 6

Location: Makamby and Amparafaka.Material: UAP-10.553B; UAP-10.561B; UAP-10.627.Description: The test is elliptical and shows a wide central part

which is compressed on each side. Planispirally rolled; wall is por-celaneous. Aperture terminal with a bifid tooth.

Remarks: The genus is generally found in the inner shelf orlagoon, preferring temperate to warm water conditions (Murray,2006).

Stratigraphic range: Upper Cretaceous–Holocene (Loeblich andTappan, 1988).

Family ALVEOLINIDAE Ehrenberg 1839Genus BORELIS de Montfort 1808Borelis melo Fitchell & Moll 1798Fig. 4; 7a

Location: Makamby.Material: UAP-10.551; UAP-10.558.Description: Test porcelaneous, spheroidal. Chambers divided

into chamberlets. Aperture shaped by aligned pores.Remarks: The genus prefers algal-coated substrates, specifically

between 8 m and 40 m depth, and 18–26 �C in temperature(Murray, 2006).

Stratigraphic range: Upper Eocene to Holocene (Borelis; Loeblichand Tappan, 1988).

Borelis cf. pulchra d’Orbigny 1808Fig. 4; 7b


Location: Makamby.Material: UAP-10.551A.Description: Test porcelaneous, elongated. Chambers separate

into chamberlets. Aperture by ranged pores.Remarks: This species is found in low abundance in lagoons and

reefs of Bermuda and Florida-Bahamas (Javaux and Scott, 2003).Prefers warm water (18–26 �C), epifaunal on hard substrates andepiphytic on seagrass, in relatively shallow water (0–40 m;Loeblich and Tappan, 1988).

Stratigraphic range: Upper Eocene to Holocene (Borelis; Loeblichand Tappan, 1988).

Genus ALVEOLINELLA Douvillé 1906Alveolinella sp.Fig. 4; 8

Location: Makamby and Amparafaka.Material: UAP-10.573; UAP-10.585A; UAP-10.586A.Description: Test elongate, fusiform, porcelaneous. Chambers

divided into many alternate chamberlets. Apertures in numerousrows, on the oral face, decrease in number toward the poles.

Remarks: Epifaunal, near-shore marine; 18–26 �C; 5–100 m;inner shelf, lagoon (Murray, 2006).

Stratigraphic range: Miocene–Holocene (Loeblich and Tappan,1964, 1988).

Family SORITIDAE Ehrenberg 1839Subfamily PENEROPLINAE Schultze 1854Genus PENEROPLIS de Montfort 1808Peneroplis sp.Fig. 4; 9


UAP-10.609; UAP-10.616; UAP-10.633.Description: Test planispiraled, and narrowly coiled in the first

development stages. Adult stage is entirely or partially uncoiled.Chambers are whole (not divided). The aperture is terminal, con-sisting of a series of pores.

Remarks: Specifically linked to a very low depth, vegetal sub-stratum, and clear water. Distribution of this genus is generallynoted in the inner shelves, lagoons, reefs, and back-reefs. Peneroplisis abundant in littoral shallow waters which are clear and withinthe photic zone, warm and possessing high salinity.

Stratigraphic range: Miocene to Holocene (Loeblich and Tappan,1988).

Genus SPIROLINA Lamarck, 1804Spirolina sp.Fig. 4; 10


UAP-10.606; UAP-10.610; UAP-10.634.Description: Test calcareous, porcelaneous. Initial part is flat,

coiled planispirally, followed by an uncoiled final portion whichis short and cylindrical. The test is smooth. Aperture is terminaland rounded.

Remarks: Spirolina tolerates high salinity water of lagoons ornear the shore, generally between 18 and 26 �C (Murray, 2006).

Stratigraphic range: Eocene to Holocene (Loeblich and Tappan,1988).

Subfamily ARCHAISSINAE Cushman 1927Genus ARCHAIAS de Montfort 1808A. cf. angulatus Fitchell & Moll 1803

Fig. 4; 11

Location: Makamby.Material: UAP-10.574.Description: The procelaneous test is planispiral and lenticular

in the first stages of development. Chambers become flaring, andare subdivided into many chamberlets. Apertures with rows ofpores on terminal face of final chamber.

Remarks: The extant species, Archaias angulatus, is epifaunal andepiphytic, living on seagrass; lives with chlorophyte symbionts,from 0 to 20 m depth with a salinity from 35 to 42 g/L. Currentlyconfined to the Atlantic Ocean, mainly Caribbean (Murray, 2006).

Stratigraphic range: Miocene to Holocene (Archaias; Loeblichand Tappan, 1988).

Suborder TEXTULARIINA Delage & Hérouard 1896Superfamily LITUOLACEA De Blainville 1825Family LITUOLIDAE De Blainville 1825Subfamily CYCLAMMININAEMarie 1941Genus CYCLAMMINA Brady 1879Cyclammina sp.Fig. 4; 12

Location: Makamby and Amparafaka.Material: UAP-10.575; UAP-10.589; UAP-10.620.Description: Test compressed planispiral and involute. Umbilical

side depressed. Last spire contains 10–11 chambers.Remarks: This genus prefers the outer shelves, or even abyssal

zone (i.e., more than 100 m; Murray, 2006).Stratigraphic range: Paleocene to Holocene (Loeblich and

Tappan, 1988).

Superfamily AMMODISCACE, Reuss 1862Family AMMODISCIDAE Reuss 1862Subfamily AMMODISCINAE Reuss 1862Genus AMMODISCUS Reuss 1862Ammodiscus sp.Fig. 4; 13

Location: Amparafaka.Material: UAP-10.597.Description: The test is discoid and agglutinated. One undivided

chamber planispirally enrolled constitute the test after the prolo-culus. The aperture is at the end of this chamber.

Remarks: Often found in littoral environments.Stratigraphic range: Silurian to Holocene (Loeblich and Tappan,

1988).

Family ASTRORHIZIDAE Brady 1881Subfamily ASTRORHIZINAE Brady 1881Genus RHABDAMMINA M. Sars 1869Rhabdammina sp.Fig. 4; 14

Location: Amparafaka.Material: UAP-10.611.Description: Test subcylindrical and ramified with three radiat-

ing tubes; wall agglutinated. Aperture at open ends of tubularbranches.

Remarks: Deep sea water form, from 700 m to 4800 m (Loeblichand Tappan, 1988).

Stratigraphic range: Holocene (Loeblich and Tappan, 1988).

Suborder ROTALIINA Delage & Hérouard 1896Superfamily ROTALIACEA Ehrenberg 1839Family ROTALIIDAE Ehrenberg 1839


Subfamily ROTALIINAE Ehrenberg 1839Genus AMMONIA Brünnich 1772Ammonia cf. beccarii Linnaeus 1758Fig. 4; 15


UAP-10.635; UAP-10.640.Description: Test is hyaline, with trochospiral coil; spiral side is

evolute and umbilical side, involute. Umbilical side is lightly con-vex, and spiral face is always strongly convex. The last whorl con-tains 10–12 chambers that gradually increase in size. Sutures aredepressed and radials are present on umbilical side. Apertureappears to be an arch bordered by a lip.

Remarks: Mostly found in association with Quinqueloculina andElphidium. Generally, this species occurs into mangroves, lagoonsand also in low energy zones such as protected bays. Tolerate largesalinity (even living in brackish water).

Stratigraphic range: Lower Miocene to Holocene (Loeblich andTappan, 1988).

Family ELPHIDIIDAE Galloway 1933Subfamily ELPHIDIINAE Galloway 1933Genus ELPHIDIUM de Montfort 1808Elphidium sp.Fig. 4; 16


UAP-10.603; UAP-10.607; UAP-10.612; UAP-10.617; UAP-10.622;UAP-10.629; UAP-10.641.

Description: Test planispirally coiled, involute and bilaterallysymmetrical, hyaline. The last whorl shows several chambersalmost equal in size. Sutures are distinctly depressed. Aperture,pore(s) at the base of the oral face.

Remarks: Generally found associated with Quinqueloculina andAmmonia cf. beccarii; the genus is associated with littoral sur-rounding such as lagoons, back-reefs and reefs, and inner shelf.Stratigraphic range: Eocene–Holocene (Loeblich and Tappan, 1988).

Superfamily CASSIDULINACEA d’Orbigny 1839Family NONIONIDAE Schultze 1854Subfamily NONIONINAE Schultze 1854Genus NONION Monfort 1808Nonion sp.Fig. 4; 17

Location: Makamby and Amparafaka.Material: UAP-10.576; UAP-10.613; UAP-10.636.Description: Test planispiral throughout, general shape is ovate

to circular. More or less involute (coiling is involute to slightly evo-lute). Sutures are curved, depressed, and external surfaces aresmooth. Several chambers. Wall calcareous hyaline. Aperture, a slitat the base of the apertural face.

Remarks: This genus is often found in shallow-water marinedeposits. Like Elphidium, this genus prefers littorals zones.

Stratigraphic range: Campanian to Holocene (Loeblich andTappan, 1988).

Genus NONIONELLA Cushman 1926Nonionella sp.Fig. 4; 18

Location: Makamby and Amparafaka.Material: UAP-10.577; UAP-10.638.

Description: Test free, subtrochospiral and hyaline. Dorsal side issemi-involute, ventral side completely involute. Ventral side devel-ops long distinct chamber, which develops a flap-like projectionoverhanging the umbilicus. Aperture is a small arch.

Remarks: This genus is found in both shallow-water and deep-sea environments.

Stratigraphic range: Coniacian (upper Cretaceous) to Holocene(Loeblich and Tappan, 1988).

Family CASSIDULINIDAE d’Orbigny 1839Genus CASSIDULINA d’Orbigny 1826Cassidulina sp.Fig. 4; 19

Location: Makamby.Material: UAP-10.557A.Description: The test is subglobular, narrowly coiled and glob-

ally involute. Chambers are smooth and alternate in both sides,biserially arranged. The aperture is elongate and placed close tothe periphery (=virguline).

Remarks: Almost always found over 75 m depth. Cassidulina pre-fers cold water in bathyal zone (Murray, 2006).

Stratigraphic range: Upper Eocene to Holocene (Loeblich andTappan, 1988).

Superfamily NODOSARIACEA Ehrenberg 1838Family NODOSARIIDAE Ehrenberg 1838Subfamily NODOSARIINAE Ehrenberg 1838Genus LAGENA Walker & Jacob 1798Lagena sp.Fig. 4; 20

Location: Amparafaka.Material: UAP-10.594.Description: The test, smooth, calcareous, formed by only one

chamber (unilocular) with a reduced neck. The aperture is terminaland appears to be rounded and radiate.

Remarks: Most of the Lagenidae prefer warm and shallow water;this genus, however, is a cold and deep-water form. Due to a lot ofvariation, species within this genus are difficult to distinguish.

Stratigraphic range: Jurassic to Holocene (Loeblich and Tappan,1988).

Genus DENTALINA Risso 1826Dentalina sp.Fig. 4; 21

Location: Makamby.Material: UAP-10.578.Description: The test is bent, elongate and calcareous. Several

chambers are arranged in linear series and are enlarged graduallyas they are added. Sutures are oblique. Aperture is terminal andcentral.

Remarks: Tubular taxon; reported from <400 m (Murray,2006).

Stratigraphic range: Lower Cretaceous to Holocene (Loeblich andTappan, 1988).

Genus NODOSARIA Lamarck 1812Nodosaria sp.Fig. 4; 22


UAP-10.630; UAP-10.637; UAP-10.642.


Description: The test is composed by several chambers arrangedin linear series (uniserial and rectilinear). Contains distinct suturesperpendicular to the test axis. Test is smooth, with a circular sec-tion. Aperture terminal.

Remarks: Genus is found from a range of depths.Stratigraphic range: Lower Jurassic to Holocene (Loeblich and

Tappan, 1988).

Superfamily BULIMINACEA Jones 1875Family BOLIVINITIDAE Cushman 1927Genus BOLIVINA d’Orbigny 1839Bolivina sp.Fig. 4; 23

Location: Amparafaka.Material: UAP-10.596; UAP-10.614.Description: The test is elongated, hyaline with biserial cham-

bers. The shape is tapering such as the greatest width is towardthe oral face. Aperture elongate and terminal.

Remarks: This genus is found in a very large range of zones; spe-cies can live in both cold to warm water, and some species caneven tolerate a lack of oxygen. This genus is found in the bathyalzone as well as in the inner shelf (Murray, 2006).

Stratigraphic range: Upper Cretaceous (Maastrichtian) to Holo-cene (Loeblich and Tappan, 1988).

Superfamily ORBITOIDACEA Schwager 1876Family LEPIDOCYCLINIDAE Scheffen 1932

Fig. 4. Cenozoic foraminiferans from the Mahajanga Basin. 1. Quinqueloculina; 2. Cyclofori7b. Borelis cf. pulchra; 8. Alveolinella; 9, Peneroplis; 10. Spirolina; 11. Archaias cf. angulatuElphidium; 17. Nonion; 18. Nonionella; 19. Cassidulina; 20. Lagena; 21. Dentalina; 22. Nod

Subfamily LEPIDOCYCLININAE Scheffen 1932Genus LEPIDOCYCLINA Gümbel 1870Lepidocyclina sp.Fig. 4; 24

Location: Makamby.Material: UAP-10.557; UAP-10.566; UAP-10.570.Description: The test is circular, lenticular with ‘‘buds’’ on the

surface.Remarks: This genus is very abundant within tropical shallow-

water sediments and nearby regions.Stratigraphic range: Eocene–Lower Miocene (Loeblich and

Tappan, 1988; Eames et al., 2011).

Superfamily SPIRILLINACEA Reuss 1862Family SPIRILLINIDAE Reuss 1862Subfamily SPIRILLININAE Reuss 1862Genus SPIRILLINA Ehrenberg 1843Spirillina sp.Fig. 4; 25

Location: Makamby.Material: UAP-10.579.Description: Test calcareous and hyaline. Proloculus is followed

by a tubular second chamber which is spiral and undivided. Aper-ture terminal at the end of the second chamber.

Remarks: Genus usually found in continental shelf; 0–100 m in acold to temperate sea (Murray, 2006).

na; 3. Hauerina cf. compressa; 4. Triloculina; 5. Pyrgo; 6. Spiroloculina; 7a. Borelis melo;s; 12. Cyclammina; 13. Ammodiscus; 14. Rhabdammina; 15. Ammonia cf. beccarii; 16.osaria; 23. Bolivina; 24. Lepidocyclina; 25. Spirillina. Scale bar equals 0.50 mm.


Stratigraphic range: Upper Triassic to Holocene (Loeblich andTappan, 1988).

Acknowledgements

Fieldwork in 2010 was supported by National Geographic andin 2011 by the University of Queensland, which we gratefullyacknowledge. We thank the government of Madagascar for permis-sion to conduct this research, and the Department of Paleontologyand Biological Anthropology, University of Antananarivo, specifi-cally H. Andriamialison. Thanks to the field team: L. Raharivony,J.-L. Raharison, S. Razafisambatra, M. Irwin, Rianala, Sahondra, S.Roberts, D. Branch, C. Boutou, L. Razafiarisoa, and also villagersand fishermen of Nosy Makamby. Thanks to the staff at Madagas-car National Parks office in Soalala for their assistance and logisti-cal support. N. Teakle provided important laboratory and technicalsupport. Min Sun, head of the University of Hong Kong’s Depart-ment of Earth Sciences, is thanked for supporting JRA’s participa-tion in the 2010 fieldtrip. We also give thanks to G. Ramakaveloand her laboratory for contributing support for processing sedi-ments and for the determination of microfossils, and L. Godfreyand L. Meador for the use of their microscope. W. Renema andL. Cotton gave helpful advice.

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