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Biological Aspects of an Interesting Fossil Fish: Paramblypterus Duvernoy I (Amblypteridae, Actinopterygii)

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Biological Aspects of an Interesting Fossil Fish: Paramblypterus Duvernoy I (Amblypteridae, Actinopterygii) Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 4 (2001) 121-138 05. 1 I. 2001 Biological aspects of an interesting fossil fish: Paramblypterus duvernoy i (Amblypteridae, Actinopterygii) Kathrin Dietze’ With 10 figures Abstract Based on three-dimensional models of the skull and body of Pnrumhlyptenis dzivemoyi, function of the feeding apparatus and potential of locomotion are interpreted. Compared to most other lower actinopterygians, certain modified features regarding the suspensory apparatus, snout bones, gape, dermohyal and suborbital region. and palatoquadrate-maxillary chamber. are present in P dirvevrzyoi. These features suggest an increased versatility of the skull and/or enlargement of underlying muscles. thus improving the ability of food uptake. The weak, pin-like teeth present in P dirvemoyi suggest a main diet of small or soft-bodicd planktonic or benthic invertebrate organisms. Axial locomotion of P diivevmyi appears to lie within the range of moderate swimming speeds. k! drivernoyi probably had a flat belly indicating that this species lived close to the bottom. Key words: Lower Permian. Southwest Germany, basal actinopterygians, functional morphology, feeding, locomotion Zusammenfassung Anhand drei-dimcnsionaler Modclle des Schadels und Kdrpers von Pczrrimhlyprenis hvernoyi werdcn Funktionswcisc clcs Ernahrungsapparates und M(iglichkeitcn der Fnrtbewegung interprctiert. Im Vergleich mit anderen basalen Actinopterygiei-ti crscheint die Schadclrekonstruktion von F1 duvernoyi hinsichtlich des Suspensoriums, der Schnauzenknochen. der Muncliifl- nung. des Dermohyal- und Suborbitalbereiches, sowie der Palatoquadratum-Maxillare-Kammer abgeleitet. Dies deutrt auf eine erhdhte Bcweglichkeit in diesen Bereichen undioder eine VergroOerung der darunter liegenden Muskeln hin. W~ISvorleil- haft fur die Nahrungsaufnahme ist. Die schmalen, stiftformigen Zahne von F! drivernoyi deuten auf eine ErnBhrung \on kleincrcn oder weichkdrperigen benthischen und planktischen Invertebraten hin. Vermutlich zeichnete sich die axiale Fortbe- wegungsweise von Purarnhlypteriis durch maBige mittlere Schwimmgeschwindigkeit aus. Der Bauch von P riiivev/ioyi war wahrscheinlich abgeflacht. was darauf hinweist, dass diese Art bodenbezogen gelebt hat. Schliisselworter: Unter-Perm (Rotliegend), Sudwestdeutschland, basale Actinopterygier, Funktionsniorphologie. Erniihrung. Lo koino ti on Introduction function of their feeding apparatus (Schaeffer & Rosen 1961, Lauder 1980~1,Gottfried 1992, Ver- Basal actinopterygians, often referred to as “pa- an 1988, 1996), and even less is known about leoniscids”, are abundant in both marine and la- their potential of locomotion (Di Canzio 198s). custrine deposits worldwide (e.g., Aldinger 1937, The present study attempts to clarify the likely Blot 1966, Boy 1976, Gardiner 1963, 1967, Low- diet and food uptake, and the likely locomotor ney 1980a, Kazantseva-Selezneva 1981, Long capabilities of the Lower Permian Pflranzhlyp- 1988, Esin 1997, Daeschler 2000). Scales of these teriis duvernoyi. P dztvernoyi is a member of the fishes appear as early as the Silurian (e.g., Gross family Amblypteridae and, together with the 1968), and skeletal remains are known since the other members of this family P gelbrrti, F! tieco- Devonian (e.g., Gardiner 1984). They are found rus, and Anzblypterus lotus, has been described wcll into the Triassic (e.g., Hutchinson 1975), recently (Dietze 1998, 1999, 2000). Their mor- even though they are less abundant than in Pa- phology and intra-specific variation has been ex- leozoic times. Still, their biology is understood amined in detail and were the basis for three-di- only poorly. Few studies have dealt with the mensional models of the skull and the body. One ’ Institut fur Palaontologie, Muscuin fiir Naturkunde, Humboldt-Universitat, Invalidenstr. 43, D-10115 Berlin. Germany. E-mail: [email protected] Received January 2000, accepted December 2000 ( WILEY-VCH Verlag Berlin GmhH. 13086 Berlin. 2001 1435-1Y4~3/01/0411-0121$ l7.50+.5(1/0 122 Dietze. K.. Biological aspects of PnrLr/nhlypterus objcctivc of tk,e restoration in three dimensions Material and methods was to gain a certain understanding of the bio- logy of Pnrcin!h!\~prems diwerm!,i and its rela- All specimens of Prr,.o/iib!\lpr~'r~r.sdrivenioyi (Agassiz 1833) examined here are from lacustrine deposits from the Lower t ives. Studies of Pcr rtrri 1 hl~,preri 1.7 and A 17 1 b l~sp- Permian of southwest Germany (e.g.. Boy 1976. 1994, Dietx terii.7 are :onnected with research on 1099. 7000). For a complete list of catalogue numbers see paleoecosystel-is of lakes from the Lower Per- Diet7e (2000). Specinicns of ~trr.o/iib/~prc,r~~.sdi/vernoyi are preserved as mian of Centrid Europe (e.2.. Boy 1994. in press. coinpi-essions flattened during fossiliLalion. Surface structure, Clausing & Boy 2000. Boy et al.. Rotliegend-Pa- such as sculpturing of the skull roofing bones and scales, is Iaookologie, Unpublished report to the Deutsche present on many specimens. A Wild Mli microscope equipped with a camera lucida was used to draw the speci- Forschungsgenieinschaft [DFG]). Objectives are mens. Drauings of the best preserved specimens were used the reconstruction of environmental conditions. for threi-diinensioiinl i-cstoration. These drawings we]-e trans- paleocommunities. and food chains. To gain in- fei-I-ed oilto thin paper which was melted on thin plates of bees' \\ax. These piipei- and wax -bones" of the skull and sight in the intzractions of spccies. it is important palate \wre then cut out and assembled in a thi-ee-dimcn- to investigate their likely diet and locomotor sional model. Reconstruction of the postcranial part was un- abilities (e.g.. ( apacity to evade predators). Infor- der-taken in similar fashion and. including the restored Gns. fit onto the reconstructed skull and photographed. To obtain mation obtain :d from morphology and models thc like]! life form of I? drivcrnovi, line drawings of the serves to evalitatc the locomotory capabilities of photograpli~\sere niade. enlarged. redrawn with wax pencils Prrrrrtnhhytcw i diirertiojsi during feeding and lo- and ink and reduced to their original size. Trunk leiigtli (TL) was measured from the posterior mar- comotion. gin of the supracleithrum to the center of the caudal fin (as In functional morphology function can be in- indicated b!, thc shadcd scale in Fig. 7A). Scale rows wcrc ferred from :ertain morphological structures. counted along the lateral line between supracleithrum and the inclination of the scales anterior to the caudal fin. Where However, there is not always a direct correlate possible. fin rays were counted at the base of the fins. Draw- (Lauder 19XOz. 1995). If extinct animals are in- ing' of the tails wcrc digitized in AUTOCAD 14 and areas volved function cannot be directly observed or \\ere calculated. Aspect ratio was calculated by dividing the span (we helolv Fig. 7A) squared by the area of the caudal experimentally tested. and interpretations need fin. Hypo- and hetcrocercal angles of thc tail were measui-ed to be cstablislied carefully (Webb 1982). More- at the "chondrostean hinge" indicated by the shaded scale in over. structural elements oftcn perform a rnulti- Fig. 7. Three dimensional profiles ol the body as illustrated in plicitv of func.ions and therefore should not be 15s. 7B \\ere obtained from the wax model of P tliivernuyi. evaluated sep: rately (Liem Bi Wake 1985). One Tc I- i 11 o I o g ! way to avoid nisleading lines of evidence when m dealing with fossils. is to find closely related. and Terminology follo\\~sArratia & Schultze (1991) and Arratia & Clouticr (1096). Other terminologies set off by quotation ideally sirnilai constructed Recent species of niarks. except for the -nasal". reler to earlier works (e.g.. which empiric.11 data is available to test the va- Blot lc)66. Hcylcr 1969) and employ the common tcrminol- lidity of interpretations of certain structures. An og! for pamniblypterid fishes. Different terminologies are currently being used for dermal cranial bones (e.g.. Arratia analysis of the functional anatomy of Recent b: Cloutier 19%. based on homologization: Poplin Sr Lund lower actinopt xygians can provide data for rnor- 1997. traditional nomenclature). For bone terminology phological ink rpretations of fossil lower actinop- adopted hcrcin differing from common usage. e.g. "nasal 1 derinosphenotic 1-2. jugal 1-2, see Dietze (2000). terygians. Sucl$observations may corroborate in- ference of fuiiction from structures present in In\ t i t LI t i on a 1 a b b re vi ii t i ons fossil fishes. However. only few bass) actinopter- 'Ilie material examined belongs to the institutions listed be- ygians, the morphological traits of which may be 1015: CAS. California Academy of Science; GPIM, Geolo- used to interr ret those in Pcii.ariihl?,pro.iis Sau- ~isch-Pal~ontologischcsInstitut Mainz: KMMA, Koninklijk vage. 1888. exist today. Of these. Po!\*ptetxs (e.g.. Museum \ oor Midden-Afrika. Tervuren: ME. Museum fur Naturkunde der Humboldt-Universitat, Berlin; PMNB, Pfnlz- Bartsch 1997). Acipemcr (e.5.. Marinelli & Stren- muwum fur Naturkundc. Bad Diirkheim. See Dietze (2000) ger 1973). and Lepisosreiis (e.g.. Gemballa 1995) for a complele list of specimens. have been dealt with mostly. Even though Recent specic s are somewhat different from Plrvanzh/ypferiri, the information obtained from Constructional morphology their structure; and soft tissues not preserved in Prrrcrt7ihlyplerri i can be employcd to explain cer- Feeding apparatus
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