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The Mode of Life of Devonian Entomozoacean Ostracods and the Myodocopid Mega-Assemblage Proxy for Hypoxic Events

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The Mode of Life of Devonian Entomozoacean Ostracods and the Myodocopid Mega-Assemblage Proxy for Hypoxic Events BULLETIN DE L’INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE SCIENCES DE LA TERRE, 74-SUPPL.: 73-80, 2004 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN AARDWETENSCHAPPEN, 74-SUPPL.: 73-80, 2004 The mode of life of Devonian entomozoacean ostracods and the Myodocopid Mega-Assemblage proxy for hypoxic events by Jean-Georges CASIER C asier , J.-G., 2004 - The mode of life of Devonian entomozoacean Introduction ostracods and the Myodocopid Mega-Assemblage proxy for hypoxic events. Bulletin de l’Institut royal des Sciences naturelles de Belgique, Sciences de la Terre, 74 suppl.: 73-80, 2 text-figs., 1 PL, Bruxelles- Ostracods are ecologically sensitive crustaceans, and Brussel, December 15, 2004- ISSN 0374-6291. their study provides valuable information about environ­ mental changes, particularly in the Palaeozoic. For ex­ ample, studies of more than 40,000 ostracods across the Frasnian - Famennian (F/F) boundary from several sec­ Abstract tions worldwide confirms that the upper Devonian event was one of the largest Phanerozoic extinctions for ostra­ The mode of life of entomozoacean ostracods, which belong to thecods. Approximately 75 percent of all marine ostracod Myodocopida, is controversial. It is generally assumed that they are species went extinct close to the F/F boundary(L e t h i e r s pelagic, more precisely planktonic or nektoplanktonic, and that their & C a s i e r , C a s i e r & L e t h i e r s , presence is indicative of deep environments. However, a purely mor­ 1999a; 2001), as a result phological study is not sufficient to determine their mode of life. Their of sea-level changes(C a s i e r & D evleeschouwer , 1995) relationship with the substrata and other animals, and their response to and a hypoxic event (C a s i e r , 1987a). The survival of major environmental changes must be taken into account. These data several species demonstrates that very shallow marine confirm a swimming mode of life for all the entomozoaceans, and suggest that a nektobenthic lifestyle, in poorly oxygenated, but not environments and surface waters were not affected during necessarily deep, environments is more probable than a planktonic the extinction(C a s i e r , 2003). mode of life. The presence of a Myodocopid Mega-Assemblage proxy Entomozoacean ostracods played an important role for hypoxic events, seems more useful that the differentiation of an “ Entomozoid ecotype” . during the Late Devonian mass extinction. However, their mode of life is controversial. Entomozoaceans are usually Key-words: Ostracods - Entomozoacea - Devonian - Mode of life -regarded as pelagic, more precisely planktonic or nekto­ Palaeoecology. planktonic. The goal of our paper is to demonstrate that a nektobenthic mode of life in poorly oxygenated environ­ ments is more probable for this group of ostracods. The Super-family EntomozoaceaP r i b y l , 1951 belongs Résumé to the Order MyodocopidaS a r s , 1866. Entomozoaceans are characterized by relatively large carapaces with Les Entomozoacea constituent une super-famille d’ostracodes apparte­ weakly calcified sub-equal valves (Fig. 1 and PI. 1, Figs nant à l’ordre des Myodocopida. Ils ont en commun une taille relative­ ment grande, des valves égales peu calcifiées, et une ornementation 9, 10, 12-15 show some species of entomozoaceans), the souvent comparée à des empreintes digitales. Elle leur confère une presence in some species of an adductorial sulcus (= nu­ grande valeur biostratigraphique. La plupart des ostracodologistes chal furrow), and a ribbed ornamentation frequently com­ estiment qu’ils étaient pélagiques et plus précisément planctoniques ou necto-planctoniques, et que leur abondance dans les sédiments pared to fingerprints (seeO l e m p s k a , 1992, for a detailed témoigne par conséquent d’une grande profondeur de dépôt. Mais study of their shell structure). Entomozoaceans were l’étude morphologique seule ne suffit pas pour déterminer leur mode abundant from the Silurian to the Upper Carboniferous. de vie. Leur relation avec le substrat et avec les autres organismes, ainsi que leur comportement lors des changements environnementaux ma­ jeurs doivent aussi être pris en considération. Ils montrent que les Entomozoacea étaient plutôt necto-benthiques et opportunistes dans Previous statements concerning the mode of life of des milieux pauvres en oxygène dissous, mais pas nécessairement entomozoaceans profonds. Un Mega-Assemblage à Myodocopida caractérisé par la présence d’Entomozoacea et (ou) de Cypridinacea (= Assemblage V de C a s ie r et al., 1985;C a s ie r , 2003) peut servir d’indicateur pour les M a t e r n (1929) considered the majority of entomozoa­ événements hypoxiques dans le Dévonien. ceans to be active swimmers. K u m m e r o w (1939) and Mots-clefs: Ostracodes - Entomozoacea - Dévonien - Mode de vie - R a b i e n (1954) compared entomozoaceans to modem Paléoécologie. Halocypridacea and surmised a nektonic or planktonic 74 Jean-Georges CASIER (1987a,b, 1988c) arrived to the conclusion that entomo­ zoaceans were probably nektobenthic, and adapted to survive in poorly oxygenated, but not necessary deep, environments. However this statement was not supported by G r o o s -U f f e n o r d e & S c h i n d l e r (1990), for whom a nektobenthic mode of life for entomozoaceans cannot be assumed because they have not been affected during global events, particularly the Upper Kellwasser Event in the Late Devonian. Their abundance in the Upper Kellwasser Horizon, which according toG r o o s -U f f e ­ n o r d e & S c h i n d l e r (1990) lacks benthos, and their rapid evolution are supplementary arguments against a nekto­ benthic lifestyle. Conversely, and also in the Proceedings of the Tenth International Symposium on Ostracoda, W i l k i n s o n & R i l e y (1990) arrived at the conclusion that entomozoaceans present in five marine horizons in the Namurian of Northern England were nektobenthic com­ Fig. 1 — Richterina (Volkina) zimmermanni ( V o l k , 1939). ponent of the dysaerobic community. The fauna asso­ An example of streamlined entomozoacean in late­ ciated with the Namurian entomozoacean in northern ral, dorsal and anterior view. Matagne Formation. England represents low diversity high-abundance assem­ Boussu-en-Fagne, Belgium. Collection IRScNB blages typical of a late Palaeozoic dysaerobic environ­ n° a l211. x50. ment, and conclusivelyW i l k i n s o n & R i l e y (Ibid.) con­ sidered the entomozoacean an integral opportunistic va­ grant nektobenthic component of the dysaerobic assem­ blage, colonizing during brief spells when the oxygen life habit for this group. On the contrary, the discovery of level was suitably high. thick shelled silicifted carapaces of entomozoaceans led Furthermore in the Proceedings of the same sympo­ B lumenstengel (1965, 1973) to suggest that they were sium, B e c k e r & B l e s s (1990) considered C a s i e r heavily calcified and consequently that they were benthic ( 1987a,b)’s argument that entomozoaceans may have a organisms. B e c k e r (1971) discovered a highly calcified nektobenthic, nearshore lifestyle to be inconclusive. They carapace of entomozoacean in the Belgian Frasnian, anddid not observe mixed Eifelian and entomozoacean as­ also concluded that some entomozoaceans displayed a semblages, and consider this to be evidence that the benthic mode of life.K o z u r (1972) arrived to the con­ entomozoaceans did not live in near-shore environments. clusion that they were benthic psychrosphaeric ostracods. Finally, O l e m p s k a ( 1992) suggested that the extremely Referring to T r i e b e l (1941), G r ü n d e l , (1962, 1965) fragile thin ornamentation over the whole carapace of and B e c k e r (1976),G o o d a y (1983) summarised the entomozoaceans is apparently inconsistent with a benthic morphological evidence for a planktonic life habit formode of life, and that the extremely high flanges may be entomozoaceans: 1. the presence of a thin, weakly calci­ an adaptation to a planktonic life habit. However, for fied, often streamlined carapace; 2. a convex ventral several species with rather heavily calcified shells and margin, and 3. the tendency of the carapace to be rounded relatively short flanges,O l e m p s k a (1992) suggested that with a high surface-volume ratio. In addition,G o o d a y an epiplanktonic as well as a planktobenthic life habit is (Ibid.) suggested that the presence of flange-like ribs of conceivable. It is herein suggested that the presence of some species may increase the surface area of the shell longitudinal canals in the carapace of some entomozoa­ and retard its sinking velocity, and that their worldwide ceans may have assisted in the exchange of respiratory distribution, together with their abundance in facies gases and such a function could have been important for which are often virtually devoid of other fossils, are entomozoaceans in stagnant and possibly dysaerobic supplementary arguments for a planktonic existence. waters. However, G o o d a y (Ibid) noted that the carapace mor­ phology of entomozoaceans provides no clear evidence for either a planktonic or benthic existence, and he con­No argument relevant for a benthic mode of life for cluded that both these life habits may have been repre­ the entomozoaceans sented within the group. He compared some species of entomozoaceans with modem species belonging to theNo particular evidence points conclusively to a genuine Thaumatocyprididae and considered that there is no con­ benthic mode of life for the entomozoaceans. Generally, vincing morphological basis for rejecting the hypothesisthe principal argument for a benthic life habit is the that at least some entomozoaceans were benthic or nek­presence of well calcified and silicifted carapaces in some tobenthic.
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