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THE LAZARUS AMMONOID FAMILY GONIATITIDAE, the TETRANGULARLY COILED ENTOGONITIDAE, and MISSISSIPPIAN BIOGEOGRAPHY Author(S): DIETER KORN, CHRISTIAN KLUG, ROYAL H

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THE LAZARUS AMMONOID FAMILY GONIATITIDAE, the TETRANGULARLY COILED ENTOGONITIDAE, and MISSISSIPPIAN BIOGEOGRAPHY Author(S): DIETER KORN, CHRISTIAN KLUG, ROYAL H THE LAZARUS AMMONOID FAMILY GONIATITIDAE, THE TETRANGULARLY COILED ENTOGONITIDAE, AND MISSISSIPPIAN BIOGEOGRAPHY Author(s): DIETER KORN, CHRISTIAN KLUG, ROYAL H. MAPES Source: Journal of Paleontology, 79(2):356-365. Published By: The Paleontological Society DOI: http://dx.doi.org/10.1666/0022-3360(2005)079<0356:TLAFGT>2.0.CO;2 URL: http://www.bioone.org/doi/full/10.1666/0022-3360%282005%29079%3C0356%3ATLAFGT %3E2.0.CO%3B2 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. J. Paleont., 79(2), 2005, pp. 356±365 Copyright q 2005, The Paleontological Society 0022-3360/05/0079-356$03.00 THE LAZARUS AMMONOID FAMILY GONIATITIDAE, THE TETRANGULARLY COILED ENTOGONITIDAE, AND MISSISSIPPIAN BIOGEOGRAPHY DIETER KORN,1 CHRISTIAN KLUG,2 AND ROYAL H. MAPES3 1Museum fuÈr Naturkunde der Humboldt-UniversitaÈt zu Berlin, Invalidenstraûe 43, D-10115 Berlin, Germany, ,[email protected]., 2PalaÈontologisches Institut und Museum, UniversitaÈt ZuÈrich, Karl-Schmid-Str. 4, CH-8006 ZuÈrich, Switzerland, ,[email protected]., and 3Department of Geological Sciences, Ohio University, Athens 45701, ,[email protected]. ABSTRACTÐA small early Late ViseÂan (Mississippian) ammonoid assemblage with Entogonites saharensis new species and Goniatites lazarus new species is described from the eastern Anti±Atlas of Morocco, being the ®rst African record of Entogonites. The family Goniatitidae is a typical Lazarus taxon, which, after a gap representing approximately 10 million years, reappears in the fossil record. The genera Entogonites and Goniatites have a wide paleogeographic distribution (northwest Laurentia, northern and southern Variscides, north Gondwana). This indicates ammonoid cosmopolitism at the genus level at the end of the middle ViseÂan, before late ViseÂan ammonoid provinces formed. Entogonites with tetrangularly coiled juvenile whorls and with a low aperture probably had a planktonic life- style. INTRODUCTION stratigraphic declarations can be made. What can be said is that HE MISSISSIPPIAN is a time in which the evolution of the Am- Entogonites does not co-occur in the same rocks with Maxigon- T monoidea is characterized by phases of rapid diversi®cation, iatites. Based on the presence of the genus Entogonites, however, but signi®cant character stasis is known from several lineages. the age of the assemblage described here can be determined as Adaptive radiations were probably triggered by sea level changes, B1 Zone or, more probably, earliest B2 Zone of Riley (1990). especially regressions, and subsequent partitioning of habitats. The examined material is housed in the Museum fuÈr Natur- Dispersal of species was probably forced by transgressive events. kunde, Berlin, catalogue numbers MB.C.5301 to MB.C.5326 and Mississippian ammonoids are well known from Late ViseÂan (cf. in the Institut fuÈr Geowissenschaften, TuÈbingen, catalogue num- Korn, 1988, for more literature) and Serpukhovian (cf. Gordon, ber GPIT 1851±97. 1965; Ruzhencev and Bogoslovskaya, 1971, for more literature) Abbreviations in the text are as follows: D, conch diameter; W, strata. In these periods, the ammonoids became extremely diverse, whorl height; H, whorl height; U, umbilical width; A, apertural height; WER, whorl expansion rate (i.e., the expansion rate of the but comparatively little is known about their Early and Middle 2 ViseÂan ancestry. Also, little is known about the paleogeographic whorl spiral; calculated [D/(D 2 A)] ). distribution of Early and Middle ViseÂan ammonoids, since oc- THE OLDEST RECORDS OF GONIATITES currences outside of England (Riley, 1996) are very rare. Different species of Goniatites appear almost synchronously Occurrences in North Africa have a signi®cant potential to ®ll within the various geographic regions in the Late ViseÂan. Strati- some gaps in the knowledge of Mississippian ammonoids. Faunas graphically well-dated occurrences are known from the Rheno- are already known from the base of the Mississippian to the Mos- hercynian (especially the Rhenish Massif and Harz Mountains), covian, and many genus zones are represented by rather diverse where Goniatites hudsoni Bisat, 1934 occurs immediately above assemblages. DeleÂpine (1941) published a monograph on Missis- the so-called Grimmeri Bed. This is a thin cherty shale horizon sippian ammonoids from Morocco, Pareyn (1961) described nu- that contains Entogonites grimmeri (Kittl, 1904a) in large quan- merous Late Tournaisian, ViseÂan, and Serpukhovian assemblages tities. A co-occurrence of these two species is known from only from northwest Algeria, and Conrad (1984) illustrated Late Tour- one locality (Medebach±Bromberg; Nicolaus, 1963), where they naisian and Early ViseÂan material from Central and South Algeria. were collected from manganese nodules immediately above the Recently, Korn et al. (1999) showed that well-preserved Late Vi- Grimmeri Bed. This suggests that in the Rhenish Massif the over- seÂan ammonoids occur in the vicinity of Erfoud and Taouz (east- lap of Entogonites and Goniatites is limited to a very thin horizon ern Anti±Atlas, Morocco). Middle and Late Tournaisian assem- of approximately 10 cm. blages also occur in this area (Korn et al., 2002, 2003). Co-occurrences of Entogonites and Goniatites are also known In the following, a low diversity fauna with species of Ento- from the Kiruktagiak River in the Brooks Range of Alaska (Gor- gonites Kittl, 1904b and Goniatites de Haan, 1825, most probably don, 1957), but the specimens of Entogonites borealis Gordon, of early Late ViseÂan age (B1 Zone or earliest B2 Zone of Riley, 1957 and Goniatites americanus Gordon, 1971 were collected 1990), is described. This fauna contains one of the stratigraphi- from an approximately 15 m thick rock unit and it is not clear if cally oldest known representatives of Goniatites, which co-occurs only one fossil horizon is represented there. In Utah these two with the distinctive Entogonites with its unique tetrangularly species were found in the Chainman Shale (Gordon, 1971) where coiled inner whorls, a morphology unduplicated in the history of Goniatites americanus occurs approximately 15 m above Ento- the Ammonoidea. gonites borealis. Titus and Riley (1997) reported a succession of Entogonites species from Utah, a lower bed with E. borealis, MATERIAL which they regard as a synonym of E. nasutus (Schmidt, 1941), All the material comes from 12 km southeast of the Dar Kaoua and a higher occurrence with E. grimmeri. Titus and Riley (1997) Oasis, 32 km southeast of Erfoud, eastern Anti±Atlas, Morocco found specimens of the genus Goniatites in both horizons, and (Fig. 1); it was surface collected and is most probably of early pointed out the cryptic origin of the genus. Late ViseÂan age (Fig. 2). It is the same locality from which the A second assemblage in which Entogonites and Goniatites co- fauna with Maxigoniatites Korn et al., 1999 and Beyrichoceras occur in the same horizon is the new Moroccan assemblage de- Hyatt, 1884, described by Korn et al. (1999), was recorded. Ex- scribed here. In this assemblage Entogonites saharensis n. sp. was posures in this area are virtually nonexistent, and hence, no exact found in the body chambers of Goniatites lazarus n. sp. 356 KORN ET AL.ÐGONIATITIDAE AND ENTOGONITIDAE 357 FIGURE 2ÐStratigraphic distribution of the Mississippian genera of the Goniatitidae, displaying the gap in the record that spans from the latest Tournaisian to the Middle ViseÂan. FIGURE 1ÐMap of the eastern Anti±Atlas with the fossil locality indi- cated by a star located approximately midway between the communi- ties of Rissani and Taouz and about 5 km west of Erg Chebbi. Goniatitidae. Evidence is missing from the latest Tournaisian and the early and middle ViseÂan rocks (Fig. 2). This pattern of strati- graphic distribution of representatives of the family Goniatitidae displays the characteristic expression of the Lazarus Effect. The THE ANCESTRY OF GONIATITES gap in the record, spanning approximately 10 Ma (Menning et The stratigraphically distant early Late Tournaisian Progonia- al., 2000), remains problematic. tites Korn et al., 2003 displays a conch and suture morphology which is very close to Goniatites. Unfortunately, no ornament is EVOLUTIONARY TRENDS WITHIN GONIATITES known from Progoniatites, except for almost straight, coarse un- All species of Goniatites have a similar conch morphology, and dulations on steinkerns. The suture line is, except for the wider differences are mainly expressed by the ornament and suture. external lobe with its straight ¯anks, very similar to Goniatites Some species possess only crenulated growth lines, while in oth- (Fig. 3). ers there is a very strong
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