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Earth Science Frontiers, Vol. 17, Special Issue, Aug. 2010 ISSN 1005-2321<br />

Foram<strong>in</strong>iferal Response to the Toarcian Ext<strong>in</strong>ction Event<br />

M.B. Hart, M.D. Hylton, C.W. Smart<br />

School of Geography, Earth & Environmental Sciences, University of Plymouth, Drake Circus PL4 8AA, Plymouth, U K<br />

(E-mail: mhart@plymouth.ac.uk )<br />

The Early <strong>Jurassic</strong> was a time of rapidly ris<strong>in</strong>g sea<br />

level associated with the extensive spread of anoxic<br />

bottom waters (Kemp et al., 2005; de Schootbrugge et<br />

al., 2005; Cohen et al., 2007). Sea level, <strong>in</strong> particular,<br />

rose across the Pliensbachian-Toarcian boundary,<br />

culm<strong>in</strong>at<strong>in</strong>g <strong>in</strong> the Falciferum Zone of the Toarcian.<br />

The associated period of sea floor anoxia co<strong>in</strong>cides<br />

with a notable period of mass ext<strong>in</strong>ction of the mar<strong>in</strong>e<br />

fauna (Hallam <strong>and</strong> Wignall, 1997). High resolution<br />

sampl<strong>in</strong>g <strong>and</strong> study of the microfaunas through several<br />

sequences <strong>in</strong> the United K<strong>in</strong>gdom (<strong>and</strong> elsewhere)<br />

confirm that benthic foram<strong>in</strong>iferal assemblages were<br />

similarly affected by an Early Toarcian Falciferum<br />

Zone event (Hylton, 2000; Hylton <strong>and</strong> Hart, 2000).<br />

Evidence for the foram<strong>in</strong>iferal ext<strong>in</strong>ction event <strong>in</strong><br />

the Pliensbachian-Toarcian <strong>in</strong>terval <strong>in</strong>cludes the<br />

elim<strong>in</strong>atiuon of the important Lower <strong>Jurassic</strong> L<strong>in</strong>gul<strong>in</strong>a<br />

tenera, Frondicularia terquemi <strong>and</strong> Marg<strong>in</strong>ul<strong>in</strong>a prima<br />

plexus groups, <strong>in</strong>itiat<strong>in</strong>g a significant period of turnover.<br />

A marked change also occurred <strong>in</strong> the character of<br />

associated nodosariid assemblages: the uniserial forms<br />

of Nodosaria, Frondicularia <strong>and</strong> L<strong>in</strong>gul<strong>in</strong>a, which<br />

dom<strong>in</strong>ate the Pliensbachian assemblages, were largely<br />

replaced by coiled Lenticul<strong>in</strong>a <strong>in</strong> the Early Toarcian. A<br />

reduction <strong>in</strong> test size (Lilliput Effect?) <strong>and</strong> a decl<strong>in</strong>e <strong>in</strong><br />

species diversity, compared with Hettangian to<br />

S<strong>in</strong>emurian assemblages, reflect the development of<br />

low oxygen conditions followed by the subsequent<br />

renewal of the microfauna <strong>in</strong> the middle Toarcian.<br />

Little’s (1996) detailed biostratigraphic sampl<strong>in</strong>g<br />

of <strong>in</strong>vertebrate macrofaunas through complete,<br />

exp<strong>and</strong>ed sequences <strong>in</strong> North Yorkshire, Engl<strong>and</strong> <strong>and</strong><br />

<strong>in</strong> southwest Germany shows that a species level event<br />

occurs <strong>in</strong> the early Toarcian. Dur<strong>in</strong>g the late Pliens-<br />

bachian there was a diverse range of benthic species<br />

<strong>in</strong>clud<strong>in</strong>g <strong>in</strong>faunal <strong>and</strong> epifaunal bivalves, brachiopods<br />

<strong>and</strong> cr<strong>in</strong>oids. The equally diverse nekton <strong>in</strong>cluded<br />

ammonites <strong>and</strong> belemnites. While there was significant<br />

ecological disruption result<strong>in</strong>g from the onset of anoxia,<br />

the biotic responses of groups of organisms were<br />

markedly different. Taxa <strong>in</strong>habit<strong>in</strong>g the upper water<br />

column were unaffected by anoxia <strong>and</strong> <strong>in</strong>cluded<br />

ammonites <strong>and</strong> belemnites. In addition, epifaunal taxa<br />

adapted to low-oxygen conditions such as the posido-<br />

niids <strong>and</strong> <strong>in</strong>oceramids, flourished <strong>in</strong> the post-ext<strong>in</strong>ction<br />

environment dur<strong>in</strong>g the survival <strong>in</strong>terval. As conditions<br />

ameliorated, the biota became more diverse <strong>and</strong><br />

gradually began to resemble the pre-ext<strong>in</strong>ction biota.<br />

This pattern was not reflected <strong>in</strong> the response of<br />

the microfauna <strong>and</strong> especially not <strong>in</strong> the case of benthic<br />

foram<strong>in</strong>ifera. The post-ext<strong>in</strong>ction faunas were of a<br />

completely different character compared to the pre-<br />

ext<strong>in</strong>ction biotas <strong>and</strong> have greater aff<strong>in</strong>ities with<br />

Middle <strong>Jurassic</strong> foram<strong>in</strong>iferal faunas. The evidence<br />

from the North Yorkshire Coast, <strong>and</strong> other northwest<br />

European sections, confirmed the Hallam (1986) model<br />

that the ma<strong>in</strong> species ext<strong>in</strong>ctions dur<strong>in</strong>g the late<br />

Pliensbachian–early Toarcian time <strong>in</strong>terval occurred,<br />

not at the end of the Pliensbachian, but near the top of<br />

the early Toarcian tenuicostatum Zone. However, this<br />

event is only significant at species level; it cannot be<br />

recognised at genus level or at family level. This study<br />

shows a species ext<strong>in</strong>ction event with a 19% (9 from 47<br />

species) reduction <strong>in</strong> diversity at the top of the<br />

tenuicostatum Zone. This was followed by an 8% (3<br />

from 38 species) reduction at dur<strong>in</strong>g the falciferum<br />

Zone. The loss of foram<strong>in</strong>iferal species is low<br />

compared to the figures of Little (1996) who records a<br />

significant macro-species ext<strong>in</strong>ction event at the top of<br />

the tenuicostatum Zone with a 77% (17 from 22 species)<br />

reduction <strong>in</strong> diversity. The benthos was most severely<br />

affected, with an 88% species drop <strong>in</strong>clud<strong>in</strong>g the<br />

disappearance of the total <strong>in</strong>fauna.<br />

No major foram<strong>in</strong>iferal group suffered ext<strong>in</strong>ction<br />

over this <strong>in</strong>terval but the important Lower <strong>Jurassic</strong><br />

lageniids L<strong>in</strong>gul<strong>in</strong>a tenera, Frondicularia terquemi <strong>and</strong><br />

Marg<strong>in</strong>ul<strong>in</strong>a prima plexi were elim<strong>in</strong>ated; these seem<br />

to have been relatively shallow-water mid-shelf forms<br />

which were replaced by the very tolerant Lenticul<strong>in</strong>a.<br />

The generic diversity of the Lagen<strong>in</strong>a cont<strong>in</strong>ued to<br />

<strong>in</strong>crease through the Mesozoic <strong>and</strong> showed no signs of<br />

decrease through the early Toarcian events. Elevated,<br />

<strong>in</strong>terconnected cont<strong>in</strong>ents <strong>and</strong> restricted seaways <strong>in</strong> the<br />

Lower <strong>Jurassic</strong> resulted <strong>in</strong> the expansion of<br />

cosmopolitan smaller foram<strong>in</strong>ifera. Common <strong>in</strong> the<br />

shales of the <strong>Jurassic</strong> (<strong>and</strong> <strong>Cretaceous</strong>), the flattened<br />

<strong>and</strong> elongate tests of the Lagen<strong>in</strong>a were probably an<br />

adaption to burrow<strong>in</strong>g <strong>in</strong> the upper few centimetres of<br />

organic-rich substrates, as detrital or bacterial sca-<br />

vengers. In contrast to these larger foram<strong>in</strong>ifera of<br />

shallow water, the relatively small Nodosariidae,<br />

Vag<strong>in</strong>ul<strong>in</strong>idae, as well as smaller Robert<strong>in</strong><strong>in</strong>a, thrived<br />

<strong>in</strong> the f<strong>in</strong>e clastic sediments deposited <strong>in</strong> deeper waters.<br />

These foram<strong>in</strong>ifera with flattened, lenticular, or low<br />

trochoid tests probably fed on bacteria or other proto-<br />

zoans, or were <strong>in</strong>faunal detrital feeders.<br />

As a result, the Lagen<strong>in</strong>a were not greatly affected<br />

by the environmental changes <strong>and</strong> result<strong>in</strong>g ext<strong>in</strong>ctions<br />

dur<strong>in</strong>g the early Toarcian, but cont<strong>in</strong>ued to show a slow<br />

<strong>and</strong> steady <strong>in</strong>crease <strong>in</strong> diversity.<br />

369

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