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MORPHOSTRUCTURE AND SYSTEMATICS OF
LINDERINA BR UGESI
SCHLUMBERGER, 1893(FORAMINIFERA, EOCENE)
CARLES FERR/iA~DEZ-CAI~ADELL & JOSEP SERRA-KIEL
FERR/kNDEZ-CAI~ADELL & SERRA-KIEL J. 1999. Morphostructure and Systematics of
Linderina brugesi
SeHLUMBERGER, 1893 (Foraminifera, Eocene). [Morphostructure et syst~matique de
Linderina brugesi
SeHLUMBERGER,1893 (Foraminifere, Eocene)]. GEOBIOS, 32, 4: 525-537. Villeurbanne, le 31.10.1999.Manuscrit d~pos~ le 23.12.1997; accept~ d~finitivement le 11.07.1998.ABSTRACT - The morphostructure of the genus
Linderina
SCHLUMBERGER
893 is revised relying on a structural studyof the type species
L. bruguesi,
SCHLUMBERGER. ts test is lamellar-involute, with annular-cyclic growth of orbitoidaltype. It has only one kind of stolons which are crosswise-oblique, alternately arranged in axial rows, and correspondwith the apertures in the last chamber. The lateral thickenings of the test are formed by the superposition of the suc-cessive involute outer lamellae of the different chambers, lacking piles and granules, and the irregularity of the late-ral surfaces is due to the lateral rounded reliefs of the alternate chamberlets. The embryonic apparatus is quadrilo-cular, with three subspherical initial chambers forming a triconch, and a fourth larger, arcuate chamber with rows ofapertures in both sides from which the orbitoidal growth starts. An explanation is given for the morphogenesis ofembryos in triconch, observed in a number of foraminiferal genera.
L. brugesi
shows a close similarity to AmericanEocene species
Caudriella ospinae
(CAUDRI),which differs from
Linderina
in that it has lateral chamberlets. Becauseof this affinity,
Caudriella
is included in the Linderinidae. A redefinition of the genus
Linderina
and of the familyLinderinidae is performed. The genera
Eoannularia
and
Epiannularia,
previously classified into the Linderinidae, donot fit into the new characterization of the family and are thus grouped into a new family, named Eoannularidae.KEYWORDS:
LINDERINA,
FORAMINIFERA, MORPHOSTRUCTURE, EOCENE, SYSTEMATICS.RI~SUMt~ - La morphostructure du genre
Linderina
SCHLUMBERGER
893 est rSvis~e ~ partir de l'~tude structuralede l'esp~ce-type,
L. brugesi.
Le test de
Linderina
est annulaire-cyclique, de croissance orbitoYdale, et lamellaire-invo-lute. I1 ne poss~de qu'un seul type de stolons qui sent entrecrois~s et disposes en rang6es axiales alternantes. Cesstolons correspondent aux ouvertures dans la derni~re loge. Les ~paissements lat~raux du test sent form,s par lasuperposition de lamelles externes involutes successives, il n'existe ni piliers ni granules. Les irr~gularit~s de la sur-face externe sent produites par les reliefs arrondis des logettes alternantes. L'appareil embryonnaire est quadrilo-culaire, avec trois loges subsphSriques initiales formant une triade et une quatri~me Ioge arqu~e, plus grande, avecdes rang~es d'ouvertures de chaque c5t5. A partir de cette loge commence la croissance orbito~dale.
L. brugesi
res-semble beaucoup ~ l'esp~ce ~oc~ne am~ricaine
Caudriella ospinae
qui se distingue notamment de
Linderina
par lapresence de logettes lat~rales. En raison de cette affinitY,
Caudriella
est rattachSe aux Linderinidae. Une redefini-tion du genre
Linderina
et de la famille Linderinidae est propos~e. Les genres
Eoannularia
et
Epiannularia,
jus-qu'ici classes parmi les Linderinidae, n'entrent plus dans la nouvelle d~finition de la famille et sent, par consequent,regroup~s en une nouvelle famille, celle des Annularidae.MOTS-CL]~S:
LINDERINA,
FORAMINIFERA, MORPHOSTRUCTURE, I~OCI~NE, SYSTEMATIQUE.
INTRODUCTION
Since its definition (Schlumberger 1893) the genus
Linderina
has been described, or simply reportedfrom the Tertiary of Europe, Africa, America, Asiaand Australia, under about 15 different speciesassignments. Nevertheless, the descriptions of thisgenus are insufficient and contradictory. There isno agreement on fundamental questions such asthe shape of the embryo or the stolon system.Here, a revision of the morphostructure of thegenus is made based on the type species,
Linde-rina bruguesi
SCHLUMBERGER, 1893, basically onthe lamellar construction and the pattern andarrangement of its megalospheric embryonicapparatus, chambers and chamberlets, stolons andapertures. This structural study allowed us to cla-rify some problematic points dealing with thestructure, such as the stolon system, or the struc-ture of the embryonic apparatus. The results ofthis study imply a revision of the systematics ofthe group at genus and family level.PREVIOUS WORKThe genus
Linderina
was defined by Schlumberger(1893) from Eocene core material from Bruges(Gironde, southwestern France) with
Linderina
 
526
brugesi
as the type species. According to Freu-denthal (1969), the material probably came fromthe "Calcaires de Blaye", with an stimatedLutetian age according to data in Schoeller (1949)and Roman & Cahuzac (1992). Since then about 15species have been assigned to this genus (Tabl. 1).Specific assignation has often been justified exclu-sively basing on external features (e.g. Le Calvez1970; Colom 1971; Ferrer 1971). Descriptions of theinternal features vary between species, and evenwithin species,, depending on different authors.The genus
Linderina
has been reported fromEocene beds in Europe (England, France, Spain,Italy, Switzerland, Hungary, Romania), Turkey,Somalia, Qatar, Oman, Iran, Iraq, Afghanistan,Pakistan, India, Burma, Borneo, Celebes, Aus-tralia, New Guinea, Bismarck Islands (W Pacific),Jamaica, Florida, St. Barthelemy (French Antilles),and Venezuela. Some references to Cretaceous
Lin-derina
(Douvilld, 1906; Silvestri 1910; Visser 1951;Hofker 1958) were later identified as
Hellenocy-clina
(MacGillavry 1963),
Monolepidorbis
(Astre1927) or
Orbitoides tissoti
(ScHLUMBERGER, 1902)lacking lateral chambers (Neumann 1958), laterassigned to a new species, O.
hottingeri,
by vanHinte (1966). Some reports of Miocene
Linderina
were reassigned to
Miogypsina
or
Planorbulina
(Nuttall & Brighton 1931). Finally,
L. chapmani
HALKYARD,
1919 was assigned to
Halkyardia
(Neumann 1958; Freudenthal 1969), and
L. ovata
HALKYARD,
1919 was chosen as type species for anew genus,
Droogerinella
(Popescu & Brotea 1995).Freudenthal (1969) made a revision of the genus
Linderina
based on the biometry of the internalcharacters. He studied material from England,France (Paris and Aquitanian basins), Somalia(including topotypes of
L. buranensis
NUTTALL &
BRIGHTON,
1931), Qatar, Celebes (including typesofL.
paronai
OSIMO, 1909), and Borneo, and consi-derably reduced the number of species by establi-shing synonymies (Tabl. i). For example, he redu-ced to one the four Indian species described bySingh (1953). Freudenthal (1969) stated that thespecies of
Linderina
had mainly been based on"slight differences of the external characteristicsin comparison with the type species
L. brugesi";
moreover he affirmed that the species he recogni-zed could be "a junior synonym of
L. brugesi,
theinternal characteristics of which are completelyunknown". The aim of this work is to provide abetter characterization of the type species and,thus, of the genus, based on a structural study.
SPECIES AUTHOR LOCALITY AGE COMMENTS
L. brugesi
Schlumberger 1893 France Middle Eocene Type species
L. paronai
Osimo 1909 Celebes Upper Eocene Recognized by Freudenthal (1969) in the Paris Basin
L. chapmani
Halkyard 1919 France Bartonian =
Halkyardia
(Neumann1958; Freudenthal 1969)
L. ovata
Halkyard 1919 France Bartonian
= Droogerinella
(Popescu & Brotea1992)
L. buranensis
Nuttall & Somalia Middle Eocene Recognized also in Qatar (Freudenthal, 1969)Brighton 1931
L. floridensis
Cole 1942 Florida Late Middle Eocene Possibly
= L. paronai
(Freudendhal 1969)
L. bihilensis
Silvestri 1948 Somalia Middle Eocene
= L. buranensis
(Azzaroli 1952; Freudenthal 1969)
L. nuttalli
Silvestri 1948 Somalia Middle Eocene
= L. buranensis
(Azzaroli 1952; Freudenthal 1969)
L. rajasthanensisL. bikanerensisL. keyalatensis
Singh 1953
IndiaLutetianThe four species of Singh can be reduced toone
= L. rajasthanensis
(Freudenthal 1969)
L. kirtharensisL. visserae
Hoiker 1958 Netherlands Maastrichtian =
Hellenocyclina
(MacGillavry 1963; Freudenthal 1969
L. douvillei
Silvestri 1910 France Originally described as
Monolepidorbis douvillei
(AstreVisser 1961 Netherlands Campanian 1927) =
Orbitoides
(Neumann 1958; van Hinte 1966)
L. baldaci
Preyer 1912 Italy Eocene-Miocene "status completely obscure" (Freudenthal 1969)
L. glaessneri
McGowran & Australia Late Eocene Not figured, not described.Beecroft 1986TABLE 1 - Species
of Linderina
found in the literature, with reference to the original description, area and age of type locality andcomments on synonymies. Some species assigned to
Linderina
were later identified as
Hellenocyclina, Orbitoides, Halkyardia,Miogypsina
or
Planorbulina,
or included into a new genus,
Droogerinella.
The rest of species, were reduced by Freudenthal (1969)to four:
L. buranensis,
NUTTALL & BRIGHTON,
L. rajasthanensis
SINGH,
L. paronai
OSIMO and
L. floridensis
COLE, though he statedthat the latter species might be a junior synonim of
L paronai,
and that one of the species may be a junior synonim of
L. brugesi.Esp~ces de
Linderina
trouvdes clans la littdrature comportant la description originale, la rdgion et l'Ctge de la localitd type et des com-mentaires sur les synonymies. Quelques esp~ces attributes au genre
Linderina
ont dtd identifi&s post~rieurement cornme
Hellenocyclina, Orbitoides, Halkyardia, Miogypsina
ou
Planorbufina,
ou classifi~es dans un nouveau genre,
Droogerinella.
Lesautres esp~ces ont ~tg r~duites & quatre par Freudenthal (1969):
L. buranensis
NUTTALL & BRIGHTON,
L. rajasthanensis
SINGH,
L.
paronai
08IMo et
L. florindensis
COLE, bien qu'il air d&larg qu'une de ces esp&ies pourrait ~tre un synonymic du
L. brugesi.
 
527MATERIAL AND METHODSOur morphostructural study was carried out onspecimens of
Linderina brugesi
SCHLUMBERGER,
1893, from level IV (Douvillfi & O'Gorman 1929)of the Tuilerie of Biron (Gave du Pau region,Southern France; see Fig. 37 in Schaub 1981 forthe location of the site), collected by one of us(Serra-Kiel) in 1979. These beds contain
Num-mulites praeaturicus
SeHAUB, 1962, N.
benehar-
mensis
DE LA HARPE IN ROZLOZSNIK,
1926, N.
bous-saci
ROZLOZSNIK, 1924,
Assilina spira spira
(DERoISS¥, 1805) and
A. exponens
(SowERBY, 1840),that correspond to the middle Lutetian (Schaub1981), or to Shallow Benthic Zone SBZ14 (MiddleLutetian 1) of Serra-Kiel et al. (1998). The tuileriewas abandoned many years ago and the outcrop isnowadays completely covered by vegetation.Only 8 specimens were found in the sample, butthey were well preserved and empty of cement andsediment, facilitating their study by SEM. Obser-rations were performed using a Hitachi 2300Scanning Electronic Microscope of the
ServeisCientifico-T~cnics
of the University of Barcelona.Because of the probable loss of the type materialofL.
brugesi
(Neumann 1954; Freudenthal 1969),the assignment of our specimens to this species isbased on the original description by Schlum-berger, and on both geographic and stratigraphicrelationships of the respective samples. Accordingto Freudenthal (1969), the species of
Linderina
from the Aquitanian basin differs from that fromthe Paris basin. The specimens described fromGironde-Aquitania (Sclumberger 1893; Welch inSchoeller 1949; Neumann 1954; this work) wouldbelong to
L. brugesi
whereas those from the Parisbasin were assigned to L. sp. cf.
paronai
byFreudenthal (1969), though he stated that they"might be named
L. paronai
(or
L. floridensis)
onthe basis of our present knowledge".
REVISION OF THE DESCRIPTIONSOF
LINDERINA
As stated above, the descriptions of the differentspecies
of Linderina
are insufficient and contradic-torious. We'll summarize here the main features ofthe descriptions found in the literature, taking intoconsideration only those species which have notbeen posteriorly reassigned to other genera.EXTERNAL MORPHOLOGYMost authors agree in the description of the exter-nal features of the test of
Linderina:
discoidalshape, more or less convex; lobulate outline; andperforate lateral surface. Most descriptions alsomention numerous small pustules, granules orpapillae on the lateral surfaces of the test (Nuttall& Brighton 1931; Singh 1953; Neumann 1954;Smout 1954; Bombita & Popescu 1977; Loeblich &Tappan 1987). In his original description, Schlum-berger (1893) only mentioned small prominencesbetween the perforations that give the test a rugo-se surface. Schlumberger (1893) and Nuttall &Brighton (1931) also point out the absence of"pillars" (piles sensu Hottinger et al. 1993).EMBRYONIC APPARATUSVarious different descriptions and interpretationsof the embryonic apparatus and the early growthstages in
Linderina
have been published.Schlumberger (1893) described a large, sphericalcentral chamber in
L. brugesi.
According to
Nuttall & Brighton (1931), the new species L.
buranensis
they described from Somalia had a"large, central, subcircular chamber" with at leastfive smaller and compressed chamber's around itin equatorial section. Smout (1954), on the otherhand, speaks of the small "nucleoconch" in
L. 5ru-gesi
from Qatar, without specifying number andarrangement of the embryonic chambers.A bilocular embryonic apparatus have been descri-bed in
L. brugesi
(e.g. Neumann 1954; Dupeuble etal. 1972; Bombita & Popescu 1977), and in the origi-nal descriptions of other species such as
L. bikaren-
sis
SINGH,
1953,
L. kirtharhensis
StNGH, 1953. L. ko-
layatensis
SINGH, 1953 and
L. floridensis
COLE, 1942.Finally, the initial chambers of
L. brugesi
weredescribed as "trocospiral" by Horv~Lth-Koll~nyi(1988), yet she did not specify if the specimens shedescribed were micro- or megalospheric forms.LAMELLAR CONSTRUCTIONSchlumberger (1983) stated that the lamellae showcontinuity in both equatorial section "where theyform rings" and axial section "where they form thelateral thickenings" (see figs 3 and 4 in Schlumber-get 1893). The continuity of the lamina "of eachwhorl" had also been described by Nuttall &Brighton (1931).Neumann (1954) and Dupeuble et al. (1972) clai-med that such an interpretation was erroneous,and that the wall is not continuous in equatorialsection. However, attached figures show conti-nuous annular walls (see fig. 7a,b in Dupeuble etaI. 1972). According to these authors, the lateralthickenings of the test correspond to a "more orless thickened calcareous wall" (Neumann 1954)or to a covering formed by several lamellae (Du-peuble eta]. 1972; and also in Loeblich & Tappan1987).The only explicite reference to the lamellar struc-ture of
Linderina
is that of Freudenthal (1969)who regards the genus as monolamellar.
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