Systematik Morphologie Palökologie Paläogeographie Stratigraphie

Verlag Dr. Friedrich PfeilPalaeo Ichthyologica Systematik Morphologie Palökologie Paläogeographie Stratigraphie

Werner SCHWARZHANS Fish otoliths from the Paleocene of Bavaria (Kressenberg) 12 and Austria (Kroisbach and Oiching-Graben)

Palaeo Ichthyologica Systematik Morphologie Palökologie Paläogeographie Stratigraphie

Werner SCHWARZHANS

Fish otoliths from the Paleocene 12 of Bavaria (Kressenberg) and Austria (Kroisbach and Oiching-Graben)

Verlag Dr. Friedrich Pfeil • München, Dezember 2012 ISSN 0724-6331 • ISBN 978-3-89937-155-0 Palaeo Ichthyologica 12 1-88 239 figs., 4 tabs. München, Dezember 2012

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ISSN 0724-6331 ISBN 978-3-89937-155-0 3

Palaeo Ichthyologica 12 1-88 239 figs., 4 tabs. München, Dezember 2012

Fish otoliths from the Paleocene of Bavaria (Kressenberg) and Austria (Kroisbach and Oiching-Graben)

Werner SCHWARZHANS*

Abstract

O toliths o f 54 teleo st species are repo rted fro m the Paleo cene strata o f Kressenberg in Bavaria and Kro isbach in Austria, representing 31 families: 33 new species are described, 7 remain in open nomenclature. Until o nly 25 years ago , descriptio ns o f Paleo cene o tolith assemblages were very scarce. The increase in knowledge since then has resulted in more than 100 valid otolith based species, with the fauna described here ranking as the largest and mo st diverse. While Kressenberg represents mainly a neritic shelf fauna, the Kroisbach association includes a number of truly open marine faunal elements, mo stly Stomiifo rmes, and is the first o f its kind from the Paleocene. The vario us Paleo cene o tolith based fish faunas sho w a remarkable degree o f regio nal diversificatio n. The data accumulated during the past 25 years from the U.S.A., Belgium, Denmark, Greenland and the Ukraine allow for a first paleo biogeographic evaluation of the Paleocene fish fauna as reconstructed from otoliths. This Paleo cene co llectio n is also remarkable fo r its transitio nal nature in the evo lutio n o f the Teleostei documenting forms that came after the K-T-bo undary extinctio n event and befo re the rise o f the mo dern Teleo stei in the Eocene after the Paleocene/ Eocene Thermal Maximum (PETM) event. Hence, there are o nly very few Paleocene species in common with either the Late Cretaceo us, Maastrichtian, which was recently described from nearby outcrops in Bavaria, or the Eocene. The main difference from the older faunal assemblage o f the Maastrichtian is the lo w level o f extinct o tolith morphologies in the Paleocene and the abundance of plesiomorphic morphologies, which represent modern groups for example of the Percoidei. O ther common groups with plesiomorphic morphologies are the Co ngridae, the O phidiifo rmes and the Stomiifo rmes (in Kro isbach). They are o ften difficult to asso ciate with living genera and in some cases even families and give rise to the large percentage o f taxa co nsidered as ‘ extinct plesio mo rphic’ o r ‘ missing links’ in teleo st phylo - geny. Gadiformes, which play an important role in the Danish Paleocene are poorly represented. O ne of the most dominant groups of the Late Cretaceo us, the Berycifo rmes, was still co mmon and specious in the Paleocene of Bavaria, but mo stly represented by genera o r families persisting until Recent. A go o d pro po rtio n of those groups can be considered as ‘living fossils’ in the Recent. The phylogenetic analyses and interpretatio n fo r several higher taxa are presented in a chapter at the end of this study.

* Dr. Werner SCHWARZHANS, Ahrensburger Weg 103, 22359 Hamburg, Germany; e-mail: [email protected]

Palaeo Ichthyologica 12 ©Palaeo 2012 IchthyologicaVerlag Dr. Friedrich 12 Pfeil, München 4

Contents

1. Introduction ...... 5

2. Geology and Locations ...... 5

3. Systematic Part ...... 9 3.1 Elo pifo rmes ...... 9 3.2 Anguillifo rmes ...... 11 3.3 Silurifo rmes ...... 20 3.4 Stomiifo rmes ...... 20 3.5 Aulo pifo rmes ...... 24 3.6 Myctophifo rmes ...... 27 3.7 Gadifo rmes ...... 29 3.8 Ophidiifo rmes ...... 37 3.9 Lophiifo rmes ...... 34 3.10 Berycifo rmes ...... 35 3.11 Zeifo rmes ...... 44 3.12 Percifo rmes ...... 45

4. Faunal reconstruction ...... 53 4.1 Comparison of the Kressenberg and the Kroisbach faunal composition ...... 54 4.2 Regional comparison and paleoecological interpretation ...... 55 4.3 Paleobiogeographic interpretation of Paleocene otolith associations ...... 57 4.4 Paleocene otolith biostratigraphy ...... 67

5. Phylogenetic analyses and evolutionary interpretation ...... 69 5.1 The Paleocene fish fauna: Linking the Late Cretaceous with the modern bloom ...... 69 5.1.1 Evo lutio nary interpretatio n ...... 69 5.1.2 Influence of the K-T boundary extinction event fo r teleo st evo lutio n (based on otolith analysis) ...... 70 5.1.3 Influence o f the PETM event fo r teleo st evo lutio n (based o n o tolith analysis) ...... 74 5.2 The Berycifo rmes: The fate o f the survivo rs from the Cretaceous (deferred extinctio n, evo lutio nary stasis leading to ‘ living fo ssils’ , adaptio n to niches) ...... 74 5.3 The Anguillifo rmes and Aulo pifo rmes early radiatio ns ...... 76 5.4 The rise o f the O phidiifo rmes (with an excursion to the Gadiformes) ...... 79 5.5 The early radiatio n o f the Percifo rmes/ Perco idei ...... 79 5.6 The Stomiifo rmes, Myctophifo rmes, Macro uridae: Early records of a bathyal fauna ...... 81

6. Acknowledgments ...... 83

7. References ...... 84

8. Addendum ...... 87

Werner SCHWARZHANS 5

1 Introduction

Even tho ugh Paleo cene o toliths were first reco rded by KO - graph o f figure 1 sho ws that the majo rity o f the species have KEN in 1885 and 1891, our knowledge of otoliths fro m that only been obtained and described during the last ten years. epoch has remained at a very low level for nearly 100 years. The steep curve of the faunal addition during that period STINTON (1965, 1977) and NO LF (1978) recorded a total ho wever also indicates that an early state o f maturity has o f 11 co nfirmed valid species fro m the Late Paleo cene Than- now been achieved and that future faunal additions are etian of England and Selandian and Thanetian of Belgium likely to level o ut. Co mparative time intervals o f the Eocene respectively. The next majo r co ntributio n to Paleo cene otoliths do rarely exceed 200 species in European deposits. Thus, from Europe, Selandian and Danian age from Denmark, the kno wledge level o f Paleo cene o toliths fro m Euro pe has is fro m SCHWARZHANS (2003) which also includes the re- now reached a level that allows a reasonable compariso n view o f KOKEN’ s and RO EDEL’s earlier papers. The Danish with Late Cretaceous and Eocene data from the same region Paleo cene no w co ntains 38 valid species. SCHWARZHANS (see chapters 4 and 5). & BRATISHKO (2011) described 21 o tolith-based species from the Middle Paleocene Selandian of Ukraine. With O utside o f Euro pe, Paleo cene o tolith data are still much the otoliths described here from the Paleocene of Bavaria mo re sparse with abo ut 28 valid species fro m No rth America and Austria the total reco rd o f o tolith-based fish taxa fro m and Greenland (FRIZZEL 1965; NO LF & DO CKERY 1993; the European Paleocene has risen to 91 species and 109 SCHWARZHANS 1985b, 2004) and two from southern otolith-based species from the Paleocene world wide. The Australia (SCHWARZHANS 1985a).

2 Geology and Locations

The o toliths described here were co llected by Friedrich tectonic unit o f the so -called ‘ Helveticum’ . The Helveticum Pfeil and Franz Traub in the years before 1982 along ar- comprises Cretaceous and Paleogene sedimentary rocks tificially enhanced o utcro ps o f the ‘ Kressenberg G raben’, depo sited alo ng the no rthern rims o f the fo rmer Penninic near Neukirchen in Bavaria and the ‘Kroisbach’ section O cean which nowadays is completely incorporated in the alo ng the Kro isbach creek in no rthern Austria (Fig. 2). The Alpine orogeny. In a narrow belt along the northern rim in intervals sampled in bo th instances were fro m the O lching fro nt o f the Alpine o verthrust, these sediments are found in Formation of Paleocene age positioned in the allochthonous intensely thrusted and tectonically repeated sectio ns (KUHN

Increase of knowledge of Paleocene otoliths KOKEN 1885 KOKEN 1891 NOLF 1978 SCHWARZHANS 1985a+b NOLF & DOCKERY 1993 ROEDEL 1930 STINTON 1965 STINTON 1977 SCHWARZHANS 2003 SCHWARZHANS 2004 BRATISHKO SCHWARZHANS & 2011 SCHWARZHANS 2012 160

140

120

100

40 number of valid species 20

0 ııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııııı 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010

Fig. 1. Increase of knowledge of Paleocene otoliths through time. Black = described species; grey = additio nal species in open nomenclature.

Palaeo Ichthyologica 12 6

c b

Mattsee Oiching S Waging Laufen a Hauns- l N z berg a c h

Freilassing Oiching Salzburg h c 441 la a a northern S 0 10 km N4 Oichinger Graben Klein- oichinger Graben Kleinoiching southern N2 Oichinger 425 Graben N1 Kroisbach Kroisbach

Kch1 Kch2 Kch3 Große Sperre

Kch11 Kch13 Craniensandstein Kch11b 60° 60° Kch12 Kch14 Kch12a Frauengrube

Bauerstatt 0 250 m

Fig. 2. Locatio n map fo r Kressenberg and Kro isbach lo calities. a, general overview, modified from KUHN 1992; b, sampling locations at “Kressenberg”, modified from KUHN 1992; c, sampling locations at “Kroisbach” and “northern O ichinger Graben”, modified fro m TRAUB 1979.

1992, EGGER et al. 2009). From shore to basin center of through Paleocene into Eocene, albeit shallowing upward the former Penninic O cean the thrusted zones include the during Eocene (HAGN et al. 1981, HEYNG 2009, KUHN No rthern Helveticum, the So uthern Helveticum, the Ultra- 1992, RASSER & PILLER 1999). Helveticum and the Alpine Flysch. The No rthern Helveticum The detailed stratigraphy o f the samples studied from the exhibits a pro minent sedimentary unco nfo rmity between the O lching Formatio n fo llo w KUHN 1992 (Fig. 3). For definitio n Middle Maastrichtian o f the G erhartsreiter Member (o toliths o f the fo rmatio ns see RASSER & PILLER (1999). described in SCHWARZHANS 2010a) and the Lutetian, while in the So uthern Helveticum o f the lo catio ns Kressenberg and At Kressenberg (Fig. 2a) the o toliths were co llected by Pfeil Kro isbach the successio n is uninterrupted fro m Maastrichtian o n suggestio n o f Pro f. Dr. Herbert Hagn, micropaleo ntolo gist

Werner SCHWARZHANS 7 at the BSPG in Munich, along two outcrop sequences of the The samples fro m Kroisbach follow the location code ‘Kressen Graben’. For more details see KUHN 1992: p. 11-15. Kch1-Kch14 o f TRAUB (in KUHN 1992) co unting fro m no rth to so uth (Fig. 2c). They co rrespo nd to the lo cations Location B is the northernmost outcrop along the western in KUHN (1992) as shown in Table 3. flank o f the graben. The sampling spo ts B1-B3 (fro m so uth to north), all of Danian age, pseudobulloides planktonic fo - O toliths were o btained fro m sampling spo ts co vering sedi- raminifera zone (P1b), correspond to the sampling numbers ments of Selandian and Thanetian age, ranging from the in KUHN 1992: tab. 1. Selandian angulata zone, P3a (1.2 m N of Kch4 and Kch4) For lo catio n details and stratigraphic age o f Locatio n B to Thanetian, upper pseudomenardii zone, P4 (Kch1 and 8 m see Table 1. S of sandstone bank) and velascoensis zone, P5 (10 m N of Kch11, Kch11, Kch11b, Kch12, Kch12a, Kch13 and Kch14). Location A refers to the main outcrop to the east of the graben. The measurements started in the south with the distances referring to the so uthernmo st co llectio n po int Table. 2. Sampling lo catio ns and stratigraphic age o f Location A. ro ughly co inciding with the lo catio ns in KUHN (1992). distance PFEIL KUHN HAG N plaktonic For lo catio n details and stratigraphic age o f Locatio n A (in m) (this study) (1992) (1981) fo raminifera zo nes see Table 2. 120.0 08 U1 P2 uncinata zone 119.5 09 U2 P2 uncinata zone This sectio n is intensely tectonized with steeply inclined 119.0 10 U3 P1c trinidade nsis zone bedding plains and several thrusted repeat sectio ns. All 118.5 11 U4 P1c trinidade nsis zone samples represent sediments of Danian age, ranging 117.5 12 U5 P1c trinidade nsis zone through the pse udo bullo ide s, trinidade nsis and uncinata 116.5 A 12.7 13 P1c trinidade nsis zone zones (P1b-P2). 116.4 14 U6 P1c trinidade nsis zone 115.3 A 11.6 15 U7 P1c trinidade nsis zone Mo st o f the o toliths fro m Kroisbach and from the nearby 115.0 16 P1c trinidade nsis zone 114.2 17 U8 P1c trinidade nsis zone ‘Oichinger Graben’ were co llected by Pfeil o n suggestio n 114.0 18 P1b pseudobulloides zone of Dr. Franz Traub. The specimens of the ‘Traub collectio n’ 113.1 A 9.6 19 U9 P1c trinidade nsis zone at the BSPG were also made available fo r this study. 112.0 A 8.3 20 U10 P1c trinidade nsis zone 111.9 21 P2 uncinata zone 110.9 22 U11 P2 uncinata zone Table. 1. Sampling lo catio ns and stratigraphic age o f Location B. 109.8 A 6.0 23 U12 P1c trinidade nsis zone 110.3 24 P1c trinidade nsis zone 108.7 25 U13 P2 uncinata zone distance PFEIL KUHN plaktonic fo raminifera zo nes 107.6 A 3.0 26 U14 P1c trinidade nsis zone (in m) (this study) (1992) 106.5 A 2.5 27 U15 P1c trinidade nsis zone 171.7 01 P1c trinidade nsis zone 106.4 A 2.0 28 P2 uncinata zone 170.2 B 3 02 P1b pseudobulloides zone 105.4 A 1.0 29 U16 P1b pseudobulloides zone 168.7 03 P1b pseudobulloides zone 104.3 30 U17 P1b pseudobulloides zone 166.8 B 2 04 P1b pseudobulloides zone 103.2 31 U18 P2 uncinata zone 163.0 05 P1c trinidade nsis zone 103.0 32 P2 uncinata zone 162.1 B 1 06 P1b pseudobulloides zone 102.1 33 U19 P1b pseudobulloides zone 161.5 07 P1b pseudobulloides zone 103.0 34 P1c trinidade nsis zone

Table. 3. Sampling lo catio ns and stratigraphic age fro m Kro isbach.

distance from sandstone bank TRAUB (1979) KUHN (1992) plaktonic fo raminifera zo nes (in m) and PFEIL (this study) 4.0 Kch1 K2 P4 upper pseudomenardii zone 8.0 8 m S of sandstone bank K6 P4 upper pseudomenardii zone 22.2 1.2 m N of Kch4 P3a angulata zone 23.4-23.8 Kch4 K12 P3a angulata zone 76.5 10 m N of Kch11 P5 velascoensis zone 86.5-87.3 Kch11 K21 P5 velascoensis zone 102.5-102.8 Kch11b P5 velascoensis zone 117-118 Kch12 K29 P5 velascoensis zone 127-128 Kch12a K30 P5 velascoensis zone 138 Kch13 K31 P5 velascoensis zone 140-141 Kch14 K32 P5 velascoensis zone

Table. 4. Sampling lo catio ns and stratigraphic age o f “no rthern O ichinger Graben”.

distance (in m) PFEIL (this study) TRAUB (1979) KUHN (1992) plaktonic fo raminifera zo nes 56 SE of water tank N4 (56 m SE of water tank) Og1 N4 P3a angulata zone 550 ESE point 441 N2 (550 m ESE point 441) (not marked) N2 P1c trinidade nsis zone 630 ESE point 441 N1 (630 m ESE point 441) Og3 N1 P1b pseudobulloides zone

Palaeo Ichthyologica 12 8

Fig. 3. G eneralized stratigraphy o f the Kressenberg and Kro isbach lo catio ns; mo dified fro m KUHN 1992. Appro ximate intervals sampled fo r o toliths are framed.

In addition a few specimens of the ‘Traub collectio n’ were For lo catio n details and stratigraphic age o f “no rthern obtained from the nearby ‘northern Oichinger Graben’ O ichinger Graben” see Table 4. (Fig. 2c). The co rrespo nding sampling lo catio ns are named variably and inco nsistent in literature. Therefo re I use here These locations are covering sediments of Danian and the exact measurements as noted by Traub (handwritten Selandian age, ranging from the Danian pseudobulloides labels, 1979) and corresponding to the naming conven- zone, P1b (N1; 630 m ESE point 441) and trinidade nsis tio n used in KUHN (1992). It seems that the lo catio ns N1 zone, P1c (N2; 550 m ESE point 441) to Selandian angulata and N2 in KUHN (1992: fig. 4) were placed further to the zone, P3a (N4; 56 m SE of water tank). southeast at the southern branch of the gorge.

Werner SCHWARZHANS 9

3 Systematic Part

All illustrated specimens and all ho lo types and paratypes Each species is accompanied with a short description are depo sited at the Bayerische Staatssammlung für Palä- (except fo r po o rly preserved specimens) co mplementing the ontologie und Geologie, München, and are indicated with figures and with the aim to o ptimize future identification of the prefix BSPG, except fo r a small number o f co mpara- similar co llectio ns. tive specimens made available fo r the private co llection of Explanatio n fo r abbreviatio ns used: O L = otolith length; Werner SCHWARZHANS. O H = o tolith height; O T = o tolith thickness; SuL = sul- The taxo no mic descriptio n o f the o toliths and the cus length; SuH = sulcus height; O sL = ostium length; mo rpho lo gical termino lo gy fo llo w that o f KOKEN (1884) CaL = cauda length; OsH = ostium height; CaH = cau- with amendments proposed by WEILER (1942) and da height; O CL = ostial co lliculum length; CCL = cau- SCHWARZHANS (1978). The metho do lo gy ado pted fo r dal co lliculum length; O CH = ostial co lliculum height; referring to species that canno t be attributed to any recent CCH = caudal colliculum height; CL = colliculum length o r fo ssil genus is described in chapter 8 (Addendum). (in case o f a single co lliculum); CH = co lliculum height The classificatio n used fo llo ws the o ne pro po sed by (in case o f a single co lliculum); aff. = affinity; vs = versus; NELSO N (2006), except for the Gonostomatidae used sensu syn = syno nymo us. lato and Paraulo pus placed in Chlorophthalmidae. Syno nymy listings are restricted to citatio ns fro m the Pal- All measurements are given in mm. All figures sho w o toliths eo cene, o ther primary citatio ns, impo rtant revisio ns relevant fro m the right side, o r mirro r imaged as if fro m the right to the described species and new or updated synonymizatio ns. side, in o rder to better facilitate co mpariso n.

3.1 Order Elopiformes

Suborder Albuloidei ventral angle located deep on ventral rim; 3, postventral angle at tip of cauda; 4, postdorsal angle. Very small Family Pterothrissidae specimens witho ut a pro minent preventral angle (Fig. 9). Genus Pteralbula SCHWARZHANS 1981 Anterio r and po sterio r tips blunt, ventral rim mo re steeply inclined than do rsal rim. Do rsal rim anterio r depressed, Pteralbula conchaeformis (KOKEN 1885) po sterio r expanded, slightly undulating. (Figs. 4-9) Inner face markedly co nvex with lo ng, stro ngly inclined sulcus. Sulcus divided in a sho rt, wide, shallo w o stium clo sely 1885 genus inc. sed. conchaeformis KOKEN – plate 5, approaching but not opening to the anterior-dorsal rim of fig. 25 the otolith, and a narrow, slightly deepened, nearly straight 1930 genus inc. sed. e rhardvo igti RO EDEL – plate 1, fig. 14 cauda with its tapering tip clo sing near the po stventral co r- 2003 Pte ralbula co nchae fo rmis (KOKEN 1885) – SCHWARZ- ner o f the o tolith. CaL : O sL = 1.2-1.5; O sH : CaH about 2. - HANS: fig. 7A J O stium with faint do rsal canal at rear upper margin. Do rsal 2004 Pte ro thrissus co nchae fo rmis (KOKEN 1885) – SCHWARZ- depressio n wide; no ventral furro w. Ventral field and rim HANS: fig. 2A-D delicately ornamented in specimens of 5 mm length and Material: 32 specimens. less. Danian (P1c), trinidade nsis zone: Kressenberg, 10 specimens: O uter face flat and smooth in large specimens, slightly 6 spec. loc. A – 2.5 m (BSPG 1984 X 1305), 2 spec. loc. A – convex and with many radial furrows in specimens of 5 mm 6 m (BSPG 1984 X 1306), 1 spec. loc. A – 8.3 m (BSPG length and less. 1984 X 1307), 1 spec. loc. A – 9.6 m (BSPG 1984 X 1308); Selandian (P3a), angulata zone: Kroisbach, 4 specimens: loc. Discussion. The extant genus Pte ro thrissus and its extinct - 1.2 m N of Kch 4 (Fig. 4a b – BSPG 1943 II 740; ex 509; relative Pte ralbula represent o ne o f the earliest co nsistent lin- Fig. 5 – BSPG 1943 II 741; ex 510; BSPG 1943 II 509; BSPG eages o f o tolith-based teleo sts in the fo ssil reco rd, reported 1943 II 510); and O ichinger Graben 1 specimen: loc. N 4 (BSPG 1943 II 737; ex 519); since Co niacian/ Santonian times (SIEBER & WEINFURTER Thanetian (P4), upper pseudomenardii zone: Kroisbach, 15 speci- 1967, SCHWARZHANS 2010a). Pte ralbula co nchae fo rmis mens: loc. Kch 1 (Fig. 6 – BSPG 1984 X 1300; Fig. 7a-b – BSPG occurs widespread in the Paleocene of the North Atlantic 1984 X 1301; Fig. 8 – BSPG 1984 X 1302; Fig. 9 – BSPG from Greenland to Denmark and now also Bavaria and 1984 X 1303; 11 spec. BSPG 1984 X 1304); Austria, but appears to missing in the early Danian (P1b) Thanetian (P5), velascoensis zone: Kroisbach, 2 specimens: 1 spec. o f Kressenberg. It represents a direct descendent o f P. fo re yi loc. Kch 12 (BSPG 1943 II 729; ex 516), 1 spec. loc. Kch 14 SCHWARZHANS 2010 fro m the Maastrichtian o f Bavaria, (BSPG 1943 II 518). fro m which it mainly differs in the mo re elo ngate shape Description. Large otoliths up to 10 mm length and more. (OL : OH = 1.35-1.75 vs 1.1) and the less deep anterior- O utline rectangular, but angles more or less rounded. ventral rim. Both species have the distinctly co nvex inner OL : OH = 1.35-1.75, decreasing with size; O H : OT = 2.7- face in common, which was considered the key character of 2.9. The four corners being: 1, anterior tip of oto lith shifted separatio n o f the extinct genus Pte ralbula fro m Pte ro thrissus do rsally and po sitio ned at lo wer margin o f o stium; 2, pre- in SCHWARZHANS 1981b.

Palaeo Ichthyologica 12 10

4 b a

8 7 a 6 b

Figs. 4-9. Pteralbula conchaeformis (KOKEN 1885). 4-5, Kro isbach, Selandian (P3a); 4, BSPG 1943 II 740; ex 509; 5, BSPG 1943 II 741; ex 510. 6-9, Kro isbach, Thanetian (P4); 6, BSPG 1984 X 1300; 7, BSPG 1984 X 1301; 8, BSPG 1984 X 1302; 9, BSPG 1984 X 1303. – 10 × ; 4b, 5 × .

Family indet. near Pterothrissidae Family Albulidae Genus Genartina FRIZZEL & DANTE 1965 Genus Albula SCOPOLI 1777

Genartina hauniensis SCHWARZHANS 2003 Albula sp. (Fig. 10) (Fig. 11)

2003 Genartina hauniensis SCHWARZHANS – fig. 8A-H Material: 1 specimen. Selandian (P3a), angulata zone: Kroisbach: loc. 1.2 m N of Kch 4 Material: 1 specimen. (Fig. 11a-b – BSPG 1943 II 717; ex 509). Selandian (P3a), angulata zone: Kroisbach: loc. 1.2 m N of Kch 4 (Fig. 10a-c – BSPG 1943 II 716; ex 509). Remarks. The single ero ded specimen is lacking its po sterio r tip. Its shallo w do rsal rim and the wide o stium resemble Remarks. The single repo rted specimen fro m Kro isbach is o toliths o f A. eppsi WHITE & FRO ST 1931 reco rded fro m the inco mpletely preserved with the anterio r part o f the o tolith Late Paleocene and Early Eocene of England (as reco rded by ero ded and the ro strum missing. The high do rsal rim with STINTON 1965). 1975 STINTON has described a number the nearly vertically cut anterio r po rtio n abo ve the o stium o f further no minally valid Albula species fro m the English is typical fo r G. hauniensis and distinguishes this species Eocene, which may represent only one or two distinct species well from the younger G. hampshirensis (SCHUBERT 1906). (NO LF 1985) and which differ from A. eppsi in a narrower Genartina hauniensis so far has only been reported from ostium. Albulid otoliths have been recorded much mo re rare the Selandian of Denmark. fro m the Paleo gene o f Euro pe than ptero thrissids, while they have been found more common and specious in North America (FRIZZELL 1965, NO LF & DO CKERY 1993).

Werner SCHWARZHANS 11

a 10 b a c 11 b

Fig. 10. Genartina hauniensis SCHWARZHANS 2003. Kro isbach, Selandian (P3a), BSPG 1943 II 716; ex 509. – 10 × . Fig. 11. Albula sp. Kro isbach, Selandian (P3a), BSPG 1943 II 717; ex 509. – 10 × .

12 c

a b

Fig. 12. Anguilla pfeili n. sp. Kressenberg, Danian (P1c), Ho lo type (BSPG 1984 X 1309) – 20 × .

3.2 Order Anguilliformes

Suborder Anguilloidei in ho rizo ntal directio n, less so vertically. Sulcus narro w, moderately deep, long, anterior open, posterior terminating Family Anguillidae mo derately clo se to po sterio r tip o f o tolith. O L : SuL = 1.2. Genus Anguilla SCHRANK 1798 Cauda much longer than ostium, straight and very slightly downward swinging; ostium short, somewhat deeper than Anguilla pfeili n. sp. cauda and slightly widened. Co lliculi po o rly defined, not (Fig. 12) differentiated between ostium and cauda. Remainder o f inner face smooth with very feeble narrow dorsal depressio n - Holotype: Fig. 12 a c; BSPG 1984 X 1309. and no ventral furro w. Type location: Kressenberg, Bavaria, lo catio n A – 2.5 m. Outer face nearly flat and smooth. Type formation: O iching Formation, Danian (P1c), trinidad- Discussion. This o tolith o f Anguilla pfe ili resembles well ensis zone. tho se o f recent representatives o f the genus Anguilla such as A. anguilla (see HÄRKÖ NEN 1986), A. ro strata (see Name: In ho no r o f Friedrich Pfeil, who co llected the o to liths NO LF 1985 and CAMPANA 2004) and A. australis and from the Kressenberg and Kroisbach locations and made A. dieffenbachi. O ther fo ssil anguillid o toliths reco rded are them available to me. Anguilla? semisphaeroides SCHWARZHANS 2003 fro m the Diagnosis. Moderately elongate and compact otolith with Middle Paleocene of Denmark, which differs in the more O L : OH = 1.6 and O H : OT = 2.4. Do rsal rim undulating, compressed and compact appearance with a nearly circu- regularly curved, without angles. Inner face convex; outer lar outline and a deep cauda, and Anguilla re ctangularis face nearly flat. Sulcus median, long, anterior open, its STINTON & NO LF 1970 from the Eocene of Belgium and cauda about three times as long as ostium, the latter slightly England, which is characterized by a pronounced post- widened and deepened. do rsal angle and the similar A. ro uxi NO LF 1977 from the O ligo cene. Ano ther similar o tolith with a po stdo rsal angle Description. Unique holotype about 2.8 mm long and con- has been described as an indeterminated Anguillidae fro m sidered diagno stically mature. O utline mo derately elongate the Paleocene of Denmark. with symmetrically po sitio ned, slightly inframedian, slightly Anguilla pfe ili is no w the earliest reco rd o f the family po inted anterio r and po sterio r tips. Do rsal rim steeper ante- Anguillidae and indicates that the family is pro bably o f rio r than po sterio r, smo o th except fo r po sterio r undulating, pre-Tertiary o rigin. So far, o nly an indistinct juvenile An- witho ut pro minent angles. Ventral rim shallo w, its median guillo idei o tolith has been repo rted fro m the Cretaceous part nearly straight ho rizo ntal, witho ut pro minent angles. in SCHWARZHANS 2010a. O tolith mo derately co mpact, its inner face stro ngly bent

Palaeo Ichthyologica 12 12

a 14 13 b a 16 b

17 ab

Fig. 13-15. Bavariconger sp. 13-14, Kro isbach, Thanetian (P4); 13, BSPG 1943 II 700; ex 430; 14, BSPG 1984 X 1310. 15, Kressen- berg, Danian (P1c), BSPG 1984 X 1311. – 30 × . Fig. 16. Protanguilla palau JOHNSON, IDA, SAKAUE, SADO, ASAHIDA & MIYA 2011. Paratype, leg. USNM 396016, Palau Blue Ho le, standard length o f fish 65 mm. – O tolith 30 × . Fig. 17. Conger illaesus SCHWARZHANS 2003. Kressenberg, Danian (P1c), BSPG 1984 X 1312. – 10 × .

Suborder indet. Description. Small and thin otoliths not exceeding much 1.5 mm length, po ssibly representing juveniles. O L : OH Family Protanguillidae = 1.7-1.8; O H : OT = 2.8. O utline regularly o val with ro unded anterio r and po sterio r tips, rather shallo w ventral The family Pro tanguillidae was recently established fo r the rim and mo re irregular do rsal rim. The latter o ccasio nally newly described marine cave dwelling Pro tanguilla palau with po stdo rsal angle and incipient lo be at level o f o stial JO HNSO N, IDA, SAKAUE, SADO , ASAHIDA & MIYA 2011. channel. JO HNSO N et al. co nsider it a ‘ living fo ssil’ amo ngst Anguilli- Inner face smo o th, slightly co nvex, with sho rt, shallo w formes representing ‘one of the most basal, independent sulcus filled with a single undivided co lliculum. Sulcus in- lineages of the true eels’ and as having retained ‘ primitive clined at angle of 10-15°. Colliculum anteriorly and pos- morphological features’ even more than those of the known terio rly reduced (O L : CL = 2.25-2.3), somewhat extending fo ssil skeletal data o f Cretaceo us anguillifo rms. An o tolith into bro ad base o f do rsally directed, sho rt o stial channel. of a paratype kindly made available by Dave Johnson Small but distinct do rsal depressio n. No ventral furro w. sho ws that the fo ssil o tolith-based genus Bavarico nge r O uter face nearly flat, smo o th. SCHWARZHANS 2010a described fro m the Maastrichtian o f Bavaria in fact belo ngs to the same family (Fig. 16). The Discussion. The available o toliths o f this Paleo cene spe- main, tho ugh significant differences o f Bavarico nge r fro m cies o f Bavarico nge r do not exceed much 1.5 mm length Pro tanguilla are the longer and more anterior reaching and probably represent juveniles. When compared with sulcus (vs anterio rly reduced) and the wider o stial channel B. po lle rs po e cki fro m the Maastrichtian o f Bavaria, which (vs sho rt and reduced). reaches nearly 3 mm length, and the recent otolith o f Pro tanguilla palau (Fig. 16) stemming from a juvenile specimen of 65 mm SL (the holotype and only adult specimen is 175 mm Genus Bavariconger SCHWARZHANS 2010 SL), these o toliths must be regarded as stemming from juveniles. For this reaso n I have refrained fro m establishing Bavariconger sp. a new species. These Bavarico nge r o toliths also resemble (Figs. 13-15) the much larger o toliths o f the parallel o ccurring Conger illae sus, but differ in the much smaller sulcus. Therefo re, it Material: 3 specimens. can no t po ssibly represent juveniles o f C. illae sus, although Danian (P1c), trinidade nsis zone: Kressenberg, 1 specimen: loc. A such are not yet known. – 12.7 m (Fig. 15 – BSPG 1984 X 1311); Thanetian (P4), upper pseudomenardii zone: Kroisbach, 2 specimens: loc. Kch 1 (Fig. 13a-b – BSPG 1943 II 700; ex 430; Fig. 14 – BSPG 1984 X 1310).

Werner SCHWARZHANS 13

18 ab

22 23

Fig. 18-23. Gnathophis probus n. sp. 18-22, Kressenberg, Danian (P1c). 18, Ho lo type, BSPG 1984 X 1314. 19-23, Paratypes. 19, BSPG 1984 X 1315; 20, BSPG 1984 X 1316; 21, BSPG 1984 X 1317; 22, BSPG 1984 X 1318. 23, Kro isbach, Thanetian (P4), Paratype, BSPG 1984 X 1320. – 20 × .

Suborder Congroidei Genus Gnathophis KAUP 1860

Family Congridae Gnathophis probus n. sp. Genus Conger OKEN 1817 (Figs. 18-23)

Conger illaesus SCHWARZHANS 2003 Holotype: Fig. 18 a-c; BSPG 1984 X 1314. (Fig. 17) Type location: Kressenberg, Bavaria, lo catio n A – 2.5 m.

2003 Co nge r illae sus SCHWARZHANS – fig. 10A-E Type formation: O iching Formation, Danian (P1c), trinidadensis zone. Material: 2 specimens. Danian (P1c), trinidade nsis zone: Kressenberg: loc. A – 2.5 m Paratypes: 6 specimens. (Fig. 17 – BSPG 1984 X 1312; BSPG 1984 X 1313). Danian (P1c), trinidade nsis zone: Kressenberg (same data as holotype), 5 specimens: loc. A – 2.5 m (Fig. 19 – BSPG 1984 X 1315; Elo ngate, thick o toliths with almo st sym- Description. Fig. 20 – BSPG 1984 X 1316; Fig. 21 – BSPG 1984 X 1317; metrical o utline. O L : OH = 2.0. Do rsal and ventral rims Fig. 22 – BSPG 1984 X 1318; BSPG 1984 X 1319); shallo w; anterio r and po sterio r tips ro unded to mo derately Thanetian (P4), upper pseudomenardii zone: Kroisbach, 1 specimen: pointed. lo c. Kch 1 (Fig. 23 – BSPG 1984 X 1320). Inner face slightly co nvex with slightly inclined, wide, Further material: 43 specimens. shallo w sulcus. Faint o stial channel leading to the anterio r- Danian (P1b), pse udo bullo ide s zone: Kressenberg, 1 specimen: do rsal rim. Sulcus with single undivided co lliculum. Do rsal lo c. B3 (BSPG 1984 X 1324); depression moderately well developed; now ventral furro w. Danian (P1c), trinidade nsis zone: Kressenberg, 38 specimens: 31 spec. O uter face moderately convex, smooth. Rims sharp. loc. A – 2.5 m (BSPG 1984 X 1321), 3 spec. loc. A – 9.6 m (BSPG 1984 X 1322), 4 spec. loc. A – 12.7 m (BSPG 1984 X 1323); Discussion. Co nge r illae sus is a rare species in the Paleo - Selandian (P3a), angulata zone: Kroisbach, 1 specimen: loc. “close cene of Denmark and Bavaria. to” Kch 4 (BSPG 1943 II 720; ex 512);

Palaeo Ichthyologica 12 14

a a 25 26 24 b

Fig. 24-26. Rhynchoconger angulosus (SCHWARZHANS 2003). Kressenberg, Danian (P1c); 24, BSPG 1984 X 1328; 25, BSPG 1984 X 1329; 26, BSPG 1984 X 1330. – 20 × .

Thanetian (P5), velascoensis zone: Kroisbach, 3 specimens: loc. from the Middle Eocene of New Zealand finally shows a 10 m N of Kch 11 (BSPG 1943 II 722; ex 513). very similar o utline including o f the massive po stdo rsal - Name: pro bus (Latin) = humble, referring to the inco n- pro jectio n, but a sho rter sulcus (O L : SuL = 2.0 vs 1.5 1.7) spicuous appearance of the otoliths. and a more distinct and wide ostial channel. Ano ther species similar in o utline and general ap- Diagnosis. O toliths mo derately co mpressed (O L : OH pearance has recently been described as Heteroconger = 1.5-1.7). O utline with pronounced postdorsal projection. astro ble maticus SCHWARZHANS & BRATISHKO 2011 fro m Sulcus mo derately sho rt (O L : SuL = 1.5-1.7) and inclined the Paleocene of Ukraine. These otoliths, however, show no at angle of 10°. O stium not widened; undivided colliculum. ostial channel and an anterior widened ostium. Indistinct o stial channel; no do rsal depressio n.

Description: Mo derately large, ro bust o toliths to abo ut Genus Paraconger KANAZAWA 1961 3 mm length. O H : OT = 2.3. O utline o val with o btuse anterior tip and expanded, often pointed, dorsally shifted, Paraconger vetustus n. sp. massive po sterio r tip. Do rsal rim shallo w anterio r, po sterio r (Figs. 37-39) with pronounced postdorsal projection, without prominent angles; ventral rim deeper than do rsal rim, regularly curved, Holotype: Fig. 37 a-b, BSPG 1984 X 1325. deepest anterio r o f its middle. Inner face smooth, strongly convex, with moderately Type location: Kressenberg, Bavaria, lo catio n A – 9.6 m. sho rt, markedly inclined and slightly deepened sulcus filled Type formation: O iching Formation, Danian (P1c), trinidad- with a single undivided colliculum. O stium and cauda un- ensis zone. divided, of equal width. Colliculum not reduced anterio r o r Paratypes: 2 specimens. po sterio r. O stial channel indistinct. No do rsal depressio n. Danian (P1c), trinidade nsis zone: Kressenberg 2 specimens: 1 spec. No ventral furro w. loc. A – 9.6 m (same data as holotype) (Fig. 38 – BSPG 1984 X O uter face slightly co nvex o r flat, smo o th. 1326); 1 spec. loc. A – 12.7 m (Fig. 39 – BSPG 1984 X 1327). Variability and Ontogeny. The variability o bserved in the Name: vetustus (Latin) = ancient, referring to the early oc- o toliths o f this species is lo w, co nfined to mild variatio ns o f currence and the plesiomorphic appearance of this species the index O L : O H and the expressio n o f the po stdo rsal rim. within the co ngrid subfamily Bathymyrinae. Most specimens are of similar sizes between 2 and 3 mm Diagnosis. O toliths co mpressed (O L : OH = 1.25-1.35). length, except for a single small specimen of about 1 mm Do rsal rim shallo w, witho ut pre- o r mid-do rsal lo be. Sulcus length, which differs fro m the larger specimens in exhibiting long (OL : SuL = 1.4-1.6), inclined at angle of 10°. Colliculi a near opening of the sulcus to the anterior-dorsal rim. undivided, shallo w; lo wer sulcus margin slightly swinging. Discussion. O toliths o f the genus Gnathophis are recog- Distinct o stial channel; no do rsal depressio n. nized by the relatively shallo w, undifferentiated sulcus, the Description. Small, compact otoliths to about 2 mm length. convex inner face, absence of a dorsal depression and O H : OT = 2.5. Anterio r tip bro adly ro unded, very o btuse; presence of an ostial channel. Gnathophis probus repre- po sterio r tip ro unded, shifted do rsally. Do rsal rim shallo w sents the earliest o tolith based reco rd o f the genus, differ- witho ut pro minent angles o r lo bes; ventral rim very deep, ing from the species in the Eocene as follows. Gnathophis regularly curved, deepest anterio r o f its middle. dissimilis (FRIZZELL & LAMBER 1962) and G. yazooensis Inner face smooth, moderately convex, with long, mark- (NO LF & STRING ER 2003) (as Paraconger yazooensis) edly inclined and relatively shallo w sulcus filled with a single fro m the Eocene o f the U.S. G ulf Co ast differ in the more undivided colliculum. O stium and cauda undivided, lower compressed appearance without postdorsal projection but a sulcus margin slightly swinging, expanded belo w o stial marked predo rsal lo be and a very sho rt sulcus. Gnathophis channel and with narrowed, not expanded or bent caudal ro se nblatti NO LF 1988 from the Early Eocene of SW-France tip. O stial channel distinct, inserted very clo se to anterio r and G. schepdaalensis STEURBAUT & NO LF 1990 fro m the tip o f sulcus. No do rsal depressio n. No ventral furro w. Early Eocene o f Belgium are mo re similar in pro po rtio ns, O uter face nearly flat, smo o th. but lack the do rsal shift o f the massive po sterio r tip (o r po stdo rsal pro jectio n) and sho w a mo re distinct and wide Discussion. The o toliths o f genus Paraconger represent one ostial channel. Gnathophis flemingi SCHWARZHANS 1980 of the more plesiomorphic morphologies in the congrid

Werner SCHWARZHANS 15

28 a

27 b c

a 31 b 33 32

34 35 36

Fig. 27-36. Rhynchoconger intercedens n. sp. 27, Kressenberg, Danian (P1c), Ho lo type, BSPG 1984 X 1332. 28-36, Paratypes. 28, Kressenberg, Danian (P1b), BSPG 1984 X 1333. 29-35, Kressenberg, Danian (P1c); 29, BSPG 1984 X 1334; 30, BSPG 1984 X 1335; 31, BSPG 1984 X 1336; 32, BSPG 1984 X 1340; 33, BSPG 1984 X 1337; 34, BSPG 1984 X 1338; 35, BSPG 1984 X 1339. 36, Kres- senberg, Danian (P2), BSPG 1984 X 1341. – 20 × ; 27b-c, 10 × .

subfamily Bathymyrinae, with the genus Ariosoma repre- Genus Rhynchoconger JORDAN & HUBBS 1925 senting the most advanced one. The otoliths of P. ve tustus are pro bably no t fully mature, but nevertheless are read- Rhynchoconger angulosus (SCHWARZHANS 2003) ily reco gnized by their flat do rsal rim witho ut any pre- or (Figs. 24-26) mid-do rsal expansio n, the inclinatio n o f the sulcus and the simple shape of the caudal tip. 2003 Rhechias angulosus SCHWARZHANS – fig. 10F-L Material: 5 specimens. Danian (P1c), trinidade nsis zone: Kressenberg: loc. A – 2.5 m (Fig. 24a-b – BSPG 1984 X 1328; Fig. 25 – BSPG 1984 X 1329; Fig. 26a-b – BSPG 1984 X 1330; 2 not fig. BSPG 1984 X 1331).

Palaeo Ichthyologica 12 16

a 37 b 38 39

a b

c 41 d

a 42 43 b c

Fig. 37-39. Paraconger vetustus n. sp. Kressenberg, Danian (P1c). 37, Ho lo type, BSPG 1984 X 1325. 38-39, Paratypes; 38, BSPG 1984 X 1326; 39, BSPG 1984 X 1327. – 20 × . Fig. 40-41. Gorgasia? turgidus n. sp. Kro isbach, Thanetian (P4). 40, Ho lo type, BSPG 1984 X 1353; 41, Paratype, BSPG 1984 X 1354. – 30 × . Fig. 42. Congridae indet. 1. Kroisbach, Thanetian (P4), BSPG 1984 X 1355. – 30 ×. Fig. 43. Congridae indet. 2. Kroisbach, Thanetian (P4), BSPG 1984 X 1356. – 30 × .

Description. Moderately elongate otoliths up to about Rhynchoconger intercedens n. sp. 5.5 mm length. OL : OH = 1.55-1.75; OH : OT about (Figs. 27-36) 2.5. Anterio r tip slightly po inted; po sterio r tip do rsally pro - nounced. Ventral rim gently and regularly curved, deepest Holotype: Fig. 27 a-c, BSPG 1984 X 1332. anterio r o f the middle; do rsal rim shallo w, po stdo rsally Type location: Kressenberg, Bavaria, lo catio n A – 2.5 m. pronounced. Rims smooth. Inner face convex, smooth. Sulcus narrow, moderately Type formation: O iching Formation, Danian (P1c), trinidad- sho rt, inclined at abo ut 10°. Sulcus margins simple oval, ensis zone. anterio rly reduced, with faint o stial channel, which is no t Paratypes: 9 specimens. filled with co lliculum. Do rsal depressio n wide, so mewhat Danian (P1b), pse udo bullo ide s zone: Kressenberg, 1 specimen: deepened. Ventral field smo o th, with faint ventral furro w lo c. B3 (Fig. 28 – BSPG 1984 X 1333); clo se to ventral rim o f o tolith. Danian (P1c), trinidade nsis zone: Kressenberg, 7 specimens: Outer face convex and smooth. 6 spec. loc. A – 2.5 m (same data as holotype) (Fig. 29 – BSPG 1984 X 1334; Fig. 30 – BSPG 1984 X 1335; Fig. 31a-b – BSPG Discussion. O toliths o f R. angulo sus differ fro m o ther Rhyn- 1984 X 1336; Fig. 33 – BSPG 1984 X 1337; Fig. 34 – BSPG choconger o toliths like R. eocenicus fro m the Early Eocene 1984 X 1338; Fig. 35 – BSPG 1984 X 1339); 1 spec. loc. A – of England and New Zealand in the much reduced nature 9.6 m (Fig. 32 – BSPG 1984 X 1340). o f the o stial channel, which led SCHWARZHANS (2003) to Danian (P2), uncinata zone: Kressenberg, 1 specimen: loc. A – place this species in the genus Rhechias JORDAN 1921, a 2.0 m (Fig. 36 – BSPG 1984 X 1341). genus now understood to be a junior synonym of Bathycon- Further material: 180 specimens. grus O G ILBY 1898 (see ESCHMEYER, 1998). The reco rds Danian (P1b), pse udo bullo ide s zone: Kressenberg, 11 specimens: o f Rhynchoconger sp. (NO LF & DO CKERY, 1993) show a 1 spec. loc. A – 1.0 m (BSPG 1984 X 1342); 1 spec. loc. B1 related species but with less reduced o stial o pening and (BSPG 1984 X 1343); 9 spec. lo c. B3 (BSPG 1984 X 1344); with a po inted po sterio r tip o f the o tolith.

Werner SCHWARZHANS 17

Danian (P1c), trinidade nsis zone: Kressenberg, 154 specimens: Genus Gorgasia MEEK & HILDEBRAND 1923 119 spec. loc. A – 2.5 m (same data as holotype) (BSPG 1984 X 1345), 6 spec. loc. A – 6.0 m (BSPG 1984 X 1346), Gorgasia? turgidus n. sp. 1 spec. loc. A – 8.3 m (BSPG 1984 X 1347), 8 spec. lo c. A – (Figs. 40-41) 9.6 m (BSPG 1984 X 1348), 3 spec. loc. A – 11.6 m (BSPG 1984 X 1349), 17 spec. loc. A – 12.7 m (BSPG 1984 X 1350); Danian (P2), uncinata zone: Kressenberg, 4 specimens: loc. A – Holotype: Fig. 40 a-d, BSPG 1984 X 1353. 2.0 m (BSPG 1984 X 1351); Type location: Kro isbach, Austria, lo catio n Kch 1. Selandian (P3a), angulata zone: Kroisbach, 5 specimens: loc. 1.2 m N of Kch 4 (BSPG 1943 II 710; ex 504); Type formation: O iching Formation, Thanetian (P4), upper and Oichinger Graben 1 specimen: loc. N 4 (BSPG 1943 II 733; pseudomenardii zone. ex 519); Thanetian (P4), upper pseudomenardii zone: Kroisbach, 5 speci- Paratype: 1 specimen. mens: loc. Kch 1 (BSPG 1984 X 1352). Thanetian (P4), upper pseudomenar dii zone: Kroisbach: loc. Kch 1 (same data as ho lo type) (Fig. 41 – BSPG 1984 X 1354). Name: intercedens (Latin) = interjacent, intermediate, referring to the intermediate mo rpho lo gy o f the o to liths o f Name: turgidus (Latin) = swo llen, referring to the thick this species. appearance o f the o toliths o f this species.

Diagnosis. O toliths mo derately co mpressed (O L:O H = 1.4- Diagnosis. Small, thick, co mpressed o toliths (O L : OH - 1.5). O utline regular oval without prominent angles o r tips. = 1.35 1.45). O utline irregular o val witho ut pro minent Sulcus inclinatio n feeble, less than 5°. Unifo rm co lliculum angles. Sulcus wide, somewhat deepened, without ostial shallo w, po sterio rly reduced, anterio rly with faint or no channel. Uniform colliculum large, partly deepened. No extensio n into feeble o stial channel; lo wer sulcus margin do rsal depressio n. straight. Pro minent do rsal depressio n. Description. Small otoliths to about 1.5 mm length. Description. Mo derately large o toliths to abo ut 4 mm O H : OT abo ut 2.0. O utline irregularly o val witho ut promi- length. O H : OT about 2.5. Anterior tip rounded; po sterio r nent tips or angles and deepest at the middle of the o tolith. tip bro adly ro unded, so metimes shifted do rsally. Do rsal Rims smo o th. rim moderately high and gently curved, without prominent Inner face markedly co nvex, with rather large, so mewhat angles o r lo bes; ventral rim deep, regularly curved, deepest deepened sulcus filled with a single undivided colliculum. at abo ut its middle o r slightly anterio r. Rims smo o th. Sulcus margin o val o r slightly sinuate. No o stial channel Inner face smo o th, markedly co nvex, with sho rt, slightly develo ped. No do rsal depressio n. No ventral furro w. inclined and shallo w sulcus filled with a single undivided O uter face less stro ngly co nvex, smo o th. co lliculum. Sulcus margin straight, with ro unded caudal tip. Variability. The single paratype differs fro m the ho lo type Co lliculum o ften terminating significantly prio r to po sterio r in a somewhat less wide sulcus which also shows some tip o f sulcus. O stial channel feeble, o ften barely visible, degree o f sinuo sity. Both is assumed to represent intraspe- inserted very clo se to anterio r tip o f sulcus. Co lliculum cific variatio n. rarely extending into o stial channel. Distinct, but rather small do rsal depressio n. No ventral furro w. Discussion. These small o toliths resemble certain o toliths Outer face markedly convex, smooth. o f the co ngrid subfamily Hetero co ngrinae, particularly of the genus Gorgasia MEEK & HILDEBRAND 1923 (see Discussion: The o toliths o f R. intercedens are intermedi- SCHWARZHANS & BRATISHKO, 2011). I have, ho wever, ate in morphology between those of other species of the placed G.? turgidus o nly tentatively in the genus until mo re genus Rhynchoconger, like R. eocenicus, and of the genus recent o toliths o f the subfamily have beco me available. Bathyco ngrus, like B. te re do philus SCHWARZHANS 2010b, through the regular outline and the weakly developed o stial channel. In most species of the genus Rhynchoconger the Congridae indet. 1 ostial channel is well developed, while in Bathyco ngrus it (Fig. 42) is absent, probably due to reduction. Rhynchoconger intercedens resembles R. angulo sus, with Material: A single small specimen fro m Kro isbach, Austria, lo cation which it shares the feeble develo pment o f the o stial channel Kch 1, Thanetian (P4), upper pseudomenardii zone of about and fro m which it differs mainly in the mo re co mpressed 1.3 mm length (Fig. 42 – BSPG 1984 X 1355). shape (OL : H = 1.4-1.5 vs 1.55-1.75), the more gently Discussion: This is a truly juvenile o tolith fo rm, but o bvi- curved dorsal rim, the more median positioned deepest ously not belonging to any of the recorded congrid species po int o f the ventral rim and the less inclined sulcus (< 5° described abo ve. Characteristic is the anterio r o pening vs 10°). A further similar species has been reco rded as o f the sulcus with a small, o val co lliculum lo cated on the Rhynchoconger sp. by NO LF & DO CKERY (1993) fro m center of the inner face. There is no dorsal depressio n o r the Paleo cene o f the U.S. G ulf Co ast differing in the more ventral furro w. elongate shape with an expanded and pointed posterio r tip. Rhynchoconger intercedens is one of the most common o tolith-based species in the Danian o f Kressenberg, but Congridae indet. 2 only moderately common in the Thanetian and Selandian (Fig. 43) o f Kro isbach. Material: A single small specimen fro m Kro isbach, Austria, lo cation Kch 1, Thanetian (P4), upper pseudomenardii zo ne o f slightly more than 1.1 mm length (Fig. 43a-c – BSPG 1984 X 1356).

Palaeo Ichthyologica 12 18

b a 44 c

45 47

Fig. 44-47. Nettastoma davejohnsoni n. sp. Kro isbach, Thanetian (P4). 44, Holotype, BSPG 1984 X 1357. 45-47, Paratypes; 45, BSPG 1984 X 1358; 46, BSPG 1984 X 1359; 47, BSPG 1984 X 1360. – 30 × .

Discussion. This very small o tolith exhibits a nevertheless Name: In honor of Dave Johnson, Washington D.C., for apparently mature morphology and therefore may represent his many co ntributio ns to ichthyo lo gy and Anguilliformes a small species rather than a juvenile specimen. Charac- and for having made the otolith of the recent Pro tanguilla teristic is the po sterio rly expanded do rsal rim, the clearly palau available to me. marked sulcus entirely filled by a shallo w co lliculum, the Diagnosis. Mo derately thick, co mpressed o toliths (O L : O H lack of a sulcus opening or ostial channel and the deep and = 1.35-1.45). O utline o val, with undulating rims, witho ut wide do rsal depressio n. This o tolith very likely represents pro minent angles. Sulcus narro w, shallo w, anterio rly closed, yet another new Paleocene congrid species, but I have re- without ostial channel, inclined at 10-15°. Unifo rm co llicu- frained fro m fo rmal descriptio n because to the singularity lum anterio rly narro wing. Mo derate do rsal depressio n. and small size o f the find. Description. Relatively small o toliths to abo ut 2 mm length. O H : OT abo ut 2.5. O utline irregularly o val, with Family Nettastomatidae undulating rims, witho ut pro minent tips o r angles. Do rsal rim so metimes po sterio rly pro no unced; ventral rim deepest Genus Nettastoma RAFINESQUE 1810 anterio r o f its middle. Inner face mo derately co nvex, with narro w, shallo w Nettastoma davejohnsoni n. sp. sulcus, anterio r clo sed and witho ut o stial channel. Sulcus (Figs. 44-47) margins shallo w do rsally and slightly curved ventrally. Sulcus steeply inclined at 10-15°, filled with a unifo rm undivided, Fig. 44 a-c, BSPG 1984 X 1357. Holotype: anterio rly narro wing co lliculum. Do rsal depressio n present, Type location: Kro isbach, Austria, lo catio n Kch 1. but feeble. No ventral furro w. Outer face more strongly convex, smooth. Type formation: O iching Formation, Thanetian (P4), upper pseudomenardii zone. Discussion. O toliths o f Nettastoma davejohnsoni can po- tentially be confused with the more common Rhynchoconger Paratypes: 4 specimens. intercedens, the main differences being the total lack of an Thanetian (P4), upper pseudomenardii zone: Kroisbach: loc. Kch 1 (same data as ho lo type) (Fig. 45 – BSPG 1984 X 1358; o stial channel (vs feeble o stial channel), the anterio rly nar- Fig. 46 – BSPG 1984 X 1359; Fig. 47 – BSPG 1984 X 1360; rowed colliculum and the undulating outline. BSPG 1984 X 1361).

Further material: 6 specimens. Thanetian (P4), upper pseudomenardii zone: Kroisbach, 5 specimens: lo c. Kch 1 (same data as ho lo type) (BSPG 1984 X 1362); Selandian (P3a), angulata zone: Kroisbach, 1 specimen: loc. 1.2 m N of Kch 4 (BSPG 1943 II 718; ex 511).

Werner SCHWARZHANS 19

b a 48

a 51 50 a c b

a 52 c

a 54 b

Fig. 48-51. Arius danicus KOKEN 1891. 49, Kressenberg, Danian (P1b), BSPG 1984 X 1363. 48, Kro isbach, Thanetian (P4), BSPG 1943 II 444; 50-51, Kro isbach, Thanetian (P5); 50, BSPG 1943 II 727; ex 516; 51, BSPG 1943 II 731; ex 518. – 10 × ; 48b, 8 × . Fig. 52-54. Arius subtilis SCHWARZHANS & BRATISHKO 2011. Kressenberg, Danian (P1c). 52, BSPG 1984 X 1370; 53, BSPG 1984 X 1371; 54, BSPG 1984 X 1368. – 10 × .

Palaeo Ichthyologica 12 20

3.3 Order Siluriformes

Family Ariidae tio n. O uter face nearly flat with so me radial furro ws and few ridges. Genus Arius CUVIER & VALENCIENNES 1840 Discussion. Arius danicus is widely distributed thro ugho ut Arius danicus KOKEN 1891 the Paleocene of Europe and Greenland and is one of (Figs. 48-51) the few Paleocene species likewise known from the Late Cretaceo us, the Maastrichtian o f Bavaria. 1891 Arius danicus KOKEN – pl. 81, fig. 1 1930 Arius ro tundus RO EDEL – pl. 1, fig. 17 ?1993 genus Ariidarum sp. 1 – NO LF & DO CKERY, pl. 2, figs. Arius subtilis SCHWARZHANS & BRATISHKO 2011 - 5 6 (Figs. 52-54) 2003 Arius danicus KOKEN 1891 – SCHWARZHANS, fig. 11J-K 2004 Arius danicus KOKEN 1891 – SCHWARZHANS, fig. 3A-F 2011 Arius subtilis SCHWARZHANS & BRATISHKO – fig. 4F-H 2010a Arius danicus KOKEN 1891 – SCHWARZHANS, figs. 31-32 Material: 9 specimens. 2011 Arius danicus KOKEN 1891 – SCHWARZHANS & BRA- Danian (P1b), pse udo bullo ide s zone: Kressenberg, 2 specimens: TISHKO, fig. 4A-E lo c. B3 (BSPG 1984 X 1367); Danian (P1c), trinidade nsis zone: Kressenberg, 6 specimens: Material: 20 specimens. 4 spec. lo c. A – 2.5 m (Fig. 54a-b – BSPG 1984 X 1368; Danian (P1b), pse udo bullo ide s zone: Kressenberg, 8 specimens: loc. BSPG 1984 X 1369); 1 spec. loc. A – 6.0 m (Fig. 52a-c – B3 (Fig. 49 – BSPG 1984 X 1363), 7 spec. (BSPG 1984 X 1364); BSPG 1984 X 1370); 1 spec. loc. A – 8.3 m (Fig. 53 – BSPG Danian (P1c), trinidade nsis zone: Kressenberg, 7 specimens: 4 spec. 1984 X 1371); loc. A – 2.5 m (BSPG 1984 X 1365); 3 spec. loc. A – 9.6 m Selandian (P3a), angulata zone: Kroisbach, 1 specimen: loc. 1.2 m (BSPG 1984 X 1366); N o f Kch 4 (BSPG 1943 II 508). Selandian (P3a), angulata zone: O ichinger Graben 1 specimen: loc. N 4 (BSPG 1943 II 734; ex 519); Description. Lapilli o toliths o f a size up to 6.5 mm. O L : O H Thanetian (P4), upper pseudomenardii zone: Kroisbach, 2 speci- = 1.4-1.6. OH : OT = 3.0-4.0, decreasing with size. mens: loc. Kch 1 (Fig. 48a-b – BSPG 1943 II 444), (BSPG Inner face moderately convex, smooth, with faint 1943 II 742; ex 444); sulcus-like feature alo ng do rsal rim widening towards the Thanetian (P5), velascoensis zone: Kroisbach, 2 specimens: loc. Kch 12 (Fig. 50a-b – BSPG 1943 II 727; ex 516), loc. Kch 14 po stdo rsal pro jectio n and with marked indentatio n in large (Fig. 51a-c – BSPG 1943 II 731; ex 518). specimens before reaching the projection. O uter face nearly flat with so me radial furro ws and ridges. Description. Arius danicus o toliths are lapilli, which are recognized by the almost regular outline, except fo r a mild Discussion. Arius subtilis is easily distinguished fro m the po stdo rsal pro jectio n. O tolith size up to nearly 10 mm. contemporaneous A. danicus through the more slender OL : OH = 1.25-1.35. OH : OT = 2.5-2.8, decreasing outline (OL : OH = 1.4-1.6 vs 1.25-1.35) and the thin- with size. ner appearance (O H : OT = 3.0-4.0 vs 2.5-2.8). It is less Inner face co nvex, smo o th, with faint sulcus-like feature widely distributed, so far o nly kno wn fro m the Danian and alo ng do rsal rim widening towards the po stdo rsal projec- Selandian o f Bavaria and Ukraine.

3.4 Order Stomiiformes

Suborder Gonostomatoidei different families ever since, namely Polyipnus G ÜNTHER 1887 in Sternoptychidae and Ichthyococcus BO NAPARTE Family Gonostomatidae 1840 in Gonostomatidae, now Phosichthyidae. Therefo re the current subdivisio n and allo catio n o f genera in the four The families G o no stomatidae, Sterno ptychidae and Phos- families mentio ned abo ve is no t suppo rted by analysis o f ichthyidae are regarded as separate families after WEITZ- o tolith mo rpho lo gy. MAN (1974) and are placed in two different suborders of The wealth of data now available from the Paleocene the Stomiifo rmes; the first two families in G o no sto matoidei o f Bavaria indicates that several separate lineages have and the latter in the Pho sichthyo idei alo ng with the Stomi- indeed existed in the Stomiifo rmes since the beginning of idae. In NELSO N (2006), referring to HARO LD (1998) a Tertiary, but it is no t po ssible at this stage to define their exact further family is listed, Diplo phidae, separated from the phylogenetic position (see also chapter 5.6). The Eocene former Gonostomatidae and placed outside the two above- and O ligocene of the Aquitaine Basin (NO LF 1988, NO LF mentioned suborders. & BRZO BO HATY 2002) and the O ligocene of Piemont, There may be all good phylogenetic rationale to sub- Italy (NO LF & STEURBAUT 2004) and fro m Eger, Hungary divide what earlier was o ne family, the G o no stomatidae. I (NO LF & BRZO BO HATY 1994) have so far yielded sizeable wo uld like to stress tho ugh, that two genera with a unique Paleogene otolith-based fish faunas with a considerable specialization in otolith morphology have been placed in two wealth o f stomiifo rm o toliths.

Werner SCHWARZHANS 21

57 a a 55 56 b b

60 62 58 59

Fig. 55-62. Progonostoma primordialis n.gen. n. sp. Kro isbach, Thanetian (P4). 55, Holotype, BSPG 1984 X 1372. 56-62, Paratypes; 56, BSPG 1984 X 1373; 57, BSPG 1984 X 1374; 58, BSPG 1984 X 1375; 59, BSPG 1984 X 1376; 60, BSPG 1984 X 1377; 61, BSPG 1984 X 1378; 62, BSPG 1984 X 1379. – 30 × .

Genus Progonostoma n. gen. Progonostoma primordialis n. sp. (Figs. 55-62) Type species: Pro go no sto ma primo rdialis n. sp. Holotype: Fig. 55 a-b, BSPG 1984 X 1372. Name: Referring to the early occurrence of the genus in the supposed evolution of Gonostomatidae. Type location: Kro isbach, Austria, lo catio n Kch 1. Diagnosis. A fo ssil o tolith-based genus o f the family G o n- Type formation: O iching Formation, Thanetian (P4), upper ostomatidae with the following combination of characters: pseudomenardii zone. Mo derately lo ng, sharp o r sho rt ro strum. Regularly rounded Paratypes: 7 specimens. o utline with gently curved do rsal, po sterio r and ventral rims. Thanetian (P4), upper pseudomenardii zone: Kroisbach: loc. Kch 1 Sulcus anterio rly o pen with o stium abo ut as wide as cauda, (same data as ho lo type) (Fig. 56a-b – BSPG 1984 X 1373; bo th barely distinguished (‘ archaesulco id’ in the termino l- Fig. 57 – BSPG 1984 X 1374; Fig. 58 – BSPG 1984 X 1375; o gy o f SCHWARZHANS, 1978). CaL : O sL = 1.3-2.0. Inner Fig. 59 – BSPG 1984 X 1376; Fig. 60 – BSPG 1984 X 1377; face flat with bro ad do rsal depressio n. O uter face slightly Fig. 61 – BSPG 1984 X 1378; Fig. 62 – BSPG 1984 X 1379). co nvex, particularly towards po sterio r rim. Further material: 33 specimens. Thanetian (P4), upper pseudomenardii zone: Kroisbach: loc. Kch 1 Discussion. O toliths o f Progonostoma resemble those of (same data as ho lo type) (BSPG 1984 X 1380). recent species of the genus Gonostoma, but these differ in the pointed and slender rostrum and the rounded posterio r Name: primo rdialis (Latin) = initial; referring to the inferred rim. Progonostoma is co nsidered to represent an early, early phylo genetic po sitio n. plesiomorphic morphology to certain gonostomatid genera, Diagnosis. O utline with ro unded do rsal and po sterio r rims, where Gonostoma and Margare thia are characterized by gently and regularly curved ventral rim and sharp, long the thin, slender ro strum (much sho rter in Margare thia) and ro strum up to nearly 40 % o f O L. Ventral ro strum margin the circular o utline o f the po sterio r part o f the o tolith, and co nvex, gently curved, do rsal margin inclined, straight. Sul- Bonapartia by the reduced rostrum and the high outline. cus lo ng, straight, indistinctly divided in o stium and cauda; Similar o toliths are also fo und in o ther families o f CaL : OsL = 1.3-1.5. the Stomiifo rmes, fo r instance in the genera Diplophos (Diplo phidae), Mauro licus (Sterno ptychidae), Polymetme, Description. Small o toliths up to 1.5 mm length. O L : OH Vincigue rria and Yare lla (all Pho sichthyidae). Each o f these, = 1.25-1.5, increasing with size and length of rostrum; ho wever, exhibit specific features o f the o utline o f the o tolith OH : OT = 3.5-4.5. Rostrum long, 25 to 40 % of O L (in- that are different fro m Progonostoma. In Diplophos and creasing with size), po inted, with co nvex ventral margin and Yare lla it is the elo ngate shape with a massive po sterio r tip inclined straight do rsal margin. Do rsal rim high, regularly and a thick, long rostrum; in Vincigue rria and Polymetme it is curved, so metimes slightly undulating; po sterio r rim broad, the very slender o utline, in Vincigue rria with a thin ro strum; regularly curved; ventral rim mo derately deep, regularly and in Mauro licus the specific and sharp postdorsal and curved, deepest at about its middle. midventral angles. Inner face flat, with slightly supramedian po sitio ned sulcus. Sulcus straight, lo ng, anterio rly o pen, po sterio rly Species: Pro go no sto ma primo rdialis and P. hagni, both from reaching clo se to po sterio r tip o f o tolith, slightly deepened the Thanetian o f Kro isbach, Austria. and indistinctly divided into similarly wide, slightly sho rter

Palaeo Ichthyologica 12 22

ab 63

c 64 65

Fig. 63-65. Progonostoma hagni n.gen. n. sp. Kro isbach, Thanetian (P4). 63, Ho lo type, BSPG 1984 X 1381. 64-65, Paratypes; 64, BSPG 1984 X 1382; 65, BSPG 1984 X 1383. – 30 × . ostium and longer cauda. Wide dorsal depression; ventral P. primo rdialis. Interestingly, the index O L : O H is in the same furro w indistinct o r no t visible. range in bo th species indicating that the sho rt ro strum in O uter face slightly co nvex and smo o th; ro strum very P: hagni is compensated by a more slender overall outline. thin, po sterio r po rtio n so mewhat thickened. A similar, but much more compressed otolith was described from the Late O ligocene of Hungary – g. Gonostomatoideo- Discussion. The lo ng and massive ro strum readily distin- rum ae nigmaticus NO LF & BRZO BO HATY 1994. guishes o toliths o f P. primo rdialis from other gonostomatid and sterno ptychid o toliths o ccurring sympatrically. A similar morphology has been described as g. Gonostomatidarum Genus Cyclogonostoma n. gen. hoffmani NO LF & BRZO BO HATY 2002 fro m the Late O ligo cene o f the Aquitaine Basin that differs in having a slightly Type species: Cyclogonostoma disciformis n. sp. more compressed appearance and a thinner rostrum, but indicating, however, that the genus Progonostoma may have Name: Referring to the almost ideal round shape of the persisted into O ligo cene times. Pro go no sto ma primo rdialis o tolith o f the type species. is the second most common species in the Thanetian of Diagnosis. A fo ssil o tolith-based genus o f the fam- Kro isbach. ily G o no stomatidae with the fo llo wing co mbinatio n o f characters: Nearly perfectly ro und o utline with minute rostrum. OL : OH = 1.1-1.15. Sulcus narro w, anterio rly Progonostoma hagni n. sp. open. O stium very short, about as wide as cauda, bo th (Figs. 63-65) barely distinguished (‘ archaesulco id’ in the terminology of SCHWARZHANS, 1978). CaL : OsL = 2.2-2.7. Inner face Holotype: Fig. 63 a-c, BSPG 1984 X 1381. flat with bro ad do rsal depressio n. O uter face slightly co nvex, Type location: Kro isbach, Austria, lo catio n Kch 1. particularly towards po sterio r rim. Type formation: O iching Formation, Thanetian (P4), upper Discussion. O toliths o f Cyclogonostoma are readily rec- pseudomenardii zone. ognized by the round shape, the almost entire lack of a ro strum and the very sho rt o stium. Within G o no stomatidae Paratypes: 3 specimens, Thanetian (P4), upper pseudo- the o toliths o f Bonapartia are the mo st similar likewise with me nardii zone: Kroisbach: loc. Kch 1 (same data as holotype) a reduced rostrum, but not quite as much as in Cyclogono- (Fig. 64 – BSPG 1984 X 1382; Fig. 65 – BSPG 1984 X 1383; stoma. BSPG 1984 X 1384). O toliths with reduced ro stri, even to the extent o bserved Name: In honor of Prof. Dr. Herbert Hagn, München, for in Cyclogonostoma, are common in Sternoptychidae, but his many co ntributio ns to the paleo ntolo gy o f the Helvetic these forms are always accompanied by a very high bodied zone of the Northern Alps. and dissimilar o tolith o utline.

Diagnosis. O utline with ro unded do rsal and po sterio r rims, Species: Cyclogonostoma is a monotypic genus with C. dis- gently and regularly curved ventral rim and sho rt and blunt cifo rmis kno wn fro m the Paleo cene o f Kro isbach, Austria. ro strum o f less than 20 % o f O L. Sulcus lo ng, straight, indistinctly divided in o stium and cauda; CaL : OsL = 1.7-2.0. Cyclogonostoma disciformis n. sp. Description. Small o toliths up to 1.3 mm length. O L : OH (Figs. 66-69) = 1.35-1.45; O H : OT about 3.5. Rostrum short, blunt, less than 20 % of O L. Dorsal rim moderately high, regularly Holotype: Fig. 66 a-b, BSPG 1984 X 1385. curved; po sterio r rim bro adly ro unded; ventral rim no t very deep, rather flat at its central po rtio n. Type location: Kro isbach, Austria, lo catio n Kch 1. Inner face slightly co nvex, with slightly supramedian Type formation: O iching Formation, Thanetian (P4), upper po sitio ned sulcus. Sulcus straight, lo ng, anterio rly open, pseudomenardii zone. po sterio rly reaching clo se to po sterio r tip o f o tolith, slightly Paratypes: 4 specimens. deepened and indistinctly divided into similarly wide, short Thanetian (P4), upper pseudomenardii zone: Kroisbach: loc. Kch ostium and long cauda. Wide, indistinct dorsal depressio n; 1 (same data as ho lo type) (Fig. 67 – BSPG 1984 X 1386; no ventral furro w. Fig. 68 – BSPG 1984 X 1387; Fig. 68 – BSPG 1984 X 1388; O uter face slightly co nvex and smo o th. BSPG 1984 X 1389).

Discussion. The sho rt ro strum and with it the sho rt o stium Name: Fro m discus (Latin) = disk, referring to the ro und expressed in the high CaL : O sL ratio distinguish P. hagni fro m disk-shape o f the o tolith.

Werner SCHWARZHANS 23

a 68 66 67 69 b

a 72 73 b 71

70 76

74 75

a 77 b 81 78 79 80

Fig. 66-69. Cyclogonostoma disciformis n. gen. n. sp. Kro isbach, Thanetian (P4). 66, Holotype, BSPG 1984 X 1385. 67-69, Paratypes; 67, BSPG 1984 X 1386; 68, BSPG 1984 X 1387; 69, BSPG 1984 X 1388. – 30 × . Fig. 70-76. Argyripnus kroisbachensis n. sp. Kro isbach, Thanetian (P4). 72, Holotype, BSPG 1984 X 1390. 70-71, 73-76, Paratypes; 70, BSPG 1984 X 1391; 71, BSPG 1984 X 1392; 73, BSPG 1984 X 1393; 74, BSPG 1984 X 1394; 75, BSPG 1984 X 1395; 76, BSPG 1984 X 1396. – 30 × . Fig. 77-81. Valenciennellus kennetti n. sp. Kro isbach, Thanetian (P4). 77, Holotype, BSPG 1984 X 1398. 78-81, Paratypes; 78, BSPG 1984 X 1399; 79, BSPG 1984 X 1400; 80, BSPG 1984 X 1401; 81, BSPG 1984 X 1402. – 30 × .

Diagnosis: See diagnosis of genus. Family Sternoptychidae Description. Small o toliths o f slightly mo re than 1.0 mm Genus Argyripnus GILBERT & CRAMER 1897 length. OL : OH = 1.1-1.15; O H : OT about 4.0. Rostrum minute, excisura small. Do rsal rim high, regularly curved; Argyripnus kroisbachensis n. sp. po sterio r rim bro adly ro unded, so metimes with feeble and (Figs. 70-76) o btuse po stventral angle; ventral rim deep, regularly curved, deepest at its middle. Holotype: Fig. 72 a-b, BSPG 1984 X 1390. Inner face rather flat, with slightly supramedian po si- Type location: Kro isbach, Austria, lo catio n Kch 1. tio ned sulcus. Sulcus straight, narro w, lo ng, anterio rly o pen, po sterio rly reaching clo se to po sterio r tip o f o tolith, slightly Type formation: O iching Formation, Thanetian (P4), upper deepened and indistinctly divided into similarly wide, very pseudomenardii zone. short ostium and long cauda. CaL : OsL = 2.2-2.7. Wide, Paratypes: 6 specimens. indistinct do rsal depressio n; no ventral furro w. Thanetian (P4), upper pseudomenardii zone: Kroisbach: loc. Kch O uter face slightly co nvex and smo o th, thickest po ste- 1 (same data as ho lo type) (Fig. 70 – BSPG 1984 X 1391; rio rly. Fig. 71 – BSPG 1984 X 1392; Fig. 73 – BSPG 1984 X 1393;

Palaeo Ichthyologica 12 24

Fig. 74 – BSPG 1984 X 1394; Fig. 75 – BSPG 1984 X 1395; Genus Valenciennellus JORDAN & EVERMANN 1896 Fig. 76 – BSPG 1984 X 1396).

Further material: 54 specimens. Valenciennellus kennetti n. sp. Thanetian (P4), upper pseudomenardii zone: Kroisbach: loc. Kch 1 (Figs. 77-81) (same data as ho lo type) (BSPG 1984 X 1397). Holotype: Fig. 77 a-b, BSPG 1984 X 1398. Name: Referring to the type-lo cality Kro isbach in Austria. Type location: Kro isbach, Austria, lo catio n Kch 1. Diagnosis. High bo died o toliths, O L : OH = 0.90-0.95. Dorsal rim with rounded pre- and postdorsal angles; ven- Type formation: O iching Formation, Thanetian (P4), upper tral rim deep; po sterio r rim blunt; ro strum less than 20 % pseudomenardii zone. o f O L. Sulcus lo ng, straight, o stium slightly deepened; - Paratypes: 4 specimens. CaL : OsL = 1.0 1.25. Thanetian (P4), upper pseudomenardii zone: Kroisbach: loc. Kch Description. Small o toliths up to 1.5 mm length. O H : OT 1 (same data as ho lo type) (Fig. 78 – BSPG 1984 X 1399; Fig. 79 – BSPG 1984 X 1400; Fig. 80 – BSPG 1984 X 1401; about 3.5. Rostrum short, blunt, less than 20 % of O L. Fig. 81 – BSPG 1984 X 1402). Dorsal rim high, with rounded pre- and postdorsal angles Further material: 18 specimens, Thanetian (P4), upper pseudo- and flat area in between; ventral rim deep, deepest at its me nardii zone: Kroisbach: loc. Kch 1 (same data as holotype) middle o r slightly towards po sterio r; po sterio r rim blunt, (BSPG 1984 X 1403). almo st vertical. Inner face slightly co nvex, with markedly supramedian Name: In honor of James Kennett, Santa Barbara, for his po sitio ned sulcus. Sulcus straight, lo ng, anterio rly open, many contributions to paleontology and pale-oceanography po sterio rly reaching very clo se to po sterio r tip o f o tolith. and the recognition of the Paleocene-Eocene Temperature O stium abo ut as lo ng as cauda o r slightly sho rter, some- Maximum (PETM). what deepened, but not widened. Wide, indistinct do rsal Diagnosis. High bo died o toliths, O L : OH = 0.75-0.80. depressio n; no o r very faint ventral furro w. Do rsal rim high; ventral rim deep; po sterio r rim nearly O uter face markedly co nvex, particularly po sterio rly and vertically cut; ro strum minute. Sulcus lo ng, straight, o stium smooth. sho rt; CaL : OsL abo ut 2.0.

Discussion. The o utline o f these o toliths resemble tho se o f Description. Small o toliths up to 1.0 mm length. O H : OT recent species o f Argyripnus, such as A. atlanticus MAUL abo ut 5.0. Rostrum minute, barely visible. Do rsal rim high, 1952 or A. ephippiatus G ILBERT & CRAMER 1897 (fo r broad, without angles; ventral rim very deep, deepest at its figures see RIVATON & BO URRET, 1999), in the high body, middle o r slightly towards anterio r; po sterio r rim vertical. the sho rt ro strum, the characteristic angular shape of the Inner face nearly flat, with median to slightly suprame- dorsal rim and the general form of the outline. The o toliths dian po sitio ned sulcus. Sulcus straight, lo ng, anterio rly o pen, of the recent species however show a shorter, somewhat po sterio rly reaching very clo se to po sterio r tip o f o tolith. widened cauda that does not reach so close to the po sterio r O stium about half as long as cauda, slightly deepened, rim o f the o tolith. no t widened. Wide, indistinct do rsal depressio n; very faint A similar fo ssil species again with a sho rter cauda ventral furro w clo se to ventral rim o f o tolith. has been described from the O ligo-Miocene transitio n in O uter face slightly co nvex and smo o th. the Aquitaine Basin by STEURBAUT 1984 – A. primige nius STEURBAUT 1984. NO LF & BRZO BO HATY 1994 mentio n Discussion. The o utline o f these o toliths is typical fo r tho se an Argyripnus sp. with a sho rt, po inted ro strum fro m the of the genus Vale ncie nne llus. The recent V. tripunctulatus Late O ligocene of Hungary. (ESMARK 1871) and the two fo ssil V. brzobohatyi STEUR- Argyripnus kroisbachensis is the most common otolith- BAUT 1982 and V. ko tthausi STEURBAUT 1979 fro m the based teleo st species in the Thanetian o f Kro isbach. O ligocene and Miocene of the Aquitaine Basin all have a slightly less reduced ro strum and a less high, clearly po sterio rly expanded do rsal rim.

3.5 Order Aulopiformes

Suborder Aulopoidei Paratypes: 3 specimens (fragmentary). Thanetian (P4), upper pseudomenardii zone: Kroisbach: loc. Kch 1 Family Aulopidae (same data as holotype) (Fig. 83 – BSPG 1984 X 1405; Fig. 84 – BSPG 1984 X 1406; BSPG 1984 X 1407). Genus Aulopus CUVIER 1816 Name: Fro m praeteritus (Latin) = bygo ne, past, referring Aulopus praeteritus n. sp. to the early phylo genetic stage o f the species. (Figs. 82-84) Diagnosis. Elo ngate o toliths; O L : OH = 1.9. Do rsal rim shallo w, nearly flat; ventral rim deepest far anterior below Holotype: Fig. 82 a-b, BSPG 1984 X 1404. o stium; ro strum bro ad, ro unded. Sulcus straight, inclined Type location: Kro isbach, Austria, lo catio n Kch 1. by abo ut 5°, its o stium o nly very slightly widened. Type formation: O iching Formation, Thanetian (P4), upper Description. Small, elongate otoliths to about 2 mm length. pseudomenardii zone. O H : OT about 2.8. Anterior rim with broad, rounded rostrum, but no excisura o r antiro strum; do rsal rim shallo w,

Werner SCHWARZHANS 25

a 84 82 b 83

a 86 ba b 85

87 88

a 90 b a b 89 c e

a 91 b

Fig. 82-84. Aulopus praeteritus n. sp. Kro isbach, Thanetian (P4). 82, Ho lo type, BSPG 1984 X 1404. 83-84, Paratypes; 83, BSPG 1984 X 1405; 84, BSPG 1984 X 1406. – 30 × . Fig. 85-88. Paraulopus postangulatus (NOLF & DOCKERY 1993). 85-86, Kressenberg, Danian (P1c); 85, BSPG 1984 X 1415; 86, BSPG 1984 X 1416. 87-88, Kro isbach, Thanetian (P4); 87, BSPG 1984 X 1421; 88, BSPG 1984 X 1422. – 20 × . Fig. 89-92. Paraulopus novellus n. sp. 89-90, Kressenberg, Danian (P1c); 89, Ho lo type, BSPG 1984 X 1408; 90, Paratype, BSPG 1984 X 1410. 91-92, Kro isbach, Thanetian (P4); 91, BSPG 1984 X 1412; 92, BSPG 1984 X 1413. – 20 × .

Palaeo Ichthyologica 12 26 nearly flat alo ng much o f its length, with marked po stdo rsal O stium widened, shallower than cauda, its dorsal mar- angle at junctio n with sho rt, vertical, do rsally shifted po sterio r gin extending o ver do rsal rim o f ro strum (see Fig. 89d). rim; ventral rim deepest anterio rly belo w o stium, thereafter Cauda narrow, long, with markedly downturned tapering gently rising to meet po sterio r rim abo ve caudal tip. Rims tip terminating mo derately clo se to po stventral rim. Do rsal slightly undulating. depressio n narro w, mo derately distinct; no ventral furro w. Inner face slightly co nvex, no t twisted alo ng lo ng axis as O uter face almo st flat, smo o th. is the case in o ther species o f the genus (see SCHWARZHANS 2003). Sulcus straight, inclined at about 5°, long, not much Variability: Three stratigraphic yo unger specimens fro m deepened. O stium narrow, not much widened but slightly Kro isbach differ fro m the type-specimens fro m the Danian of deepened compared to cauda. Cauda terminating close to Kressenberg in the more compressed outline and the more po stventral rim, with ro unded tip. Do rsal depressio n narrow, flat inner face. Because they are also smaller in size (2 mm indistinct; no ventral furro w. length) they are tentatively kept with P. no ve llus. O uter face almo st flat, smo o th except fo r few sho rt radial furro ws anterio r-ventrally. Discussion: The o toliths o f P. no ve llus strikingly resemble the o toliths o f the extant P. nigripinnis (G ÜNTHER 1878) (see Discussion. Aulo pus pre ate ritus resembles A. to rtus SCHWARZHANS 1980) even to the extent o f the peculiar SCHWARZHANS 2003 from the Danian of Denmark, dif- dorsal margin of the ostium being bent over the dorsal rim fering in the lack o f a torsio n alo ng the lo ng axis of the of the rostrum. The only comparable species in the European otolith, the much more massive, rounded rostrum and the Paleo cene (and Maastrichtian) is P. po stangulatus (NO LF & blunt po sterio r-do rsal rim. Chlo ro phthalmus udo viche nko i DO CKERY 1993), which was o riginally described fro m the SCHWARZHANS & BRATISHKO 2011 fro m the Selandian o f Paleocene of the U.S. Gulf Coast and also is common in Ukraine differs in the tapering, slightly do wnturned caudal Bavaria (see belo w). The main difference o f P. no ve llus is tip, the torsion along the long axis and the shape of the the more strongly inclined caudal tip, the combination of do rsal rim. a co nvex inner and a flat o uter face (vs bico nvex) and the bro adly undulating do rsal rim in adults. Family Chlorophthalmidae Genus Paraulopus SATO & NAKABO 2002 Paraulopus postangulatus (NOLF & DOCKERY 1993) (Figs. 85-88) Paraulopus novellus n. sp. (Figs. 89-92) 1993 genus Chlo ro phthalmidarum po stangulatus NO LF & DO CKERY – pl. 2, figs. 1-2 2003 Chlo ro phthalmus po stangulatus NO LF & DO CKERY 1993 Holotype: Fig. 89 a-e, BSPG 1984 X 1408. – SCHWARZHANS: fig. 16 G- P Type location: Kressenberg, Bavaria, lo catio n A – 2.5 m. 2004 Chlo ro phthalmus po stangulatus NO LF & DO CKERY 1993 – SCHWARZHANS: fig. 4 C-G Type formation: O iching Formation, Danian (P1c), trinidad- 2010a Chlo ro phthalmus po stangulatus NO LF & DO CKERY 1993 ensis zone. – SCHWARZHANS: figs. 52-56

Paratypes: 5 specimens. Material: 44 specimens. Danian (P1c), trinidade nsis zone: Kressenberg: 3 spec. loc. A – Danian (P1c), trinidade nsis zone: Kressenberg, 36 specimens: 2.5 m (same data as holotype) (not fig. – BSPG 1984 X 1409); 24 spec. loc. A – 2.5 m (Fig. 85a-c – BSPG 1984 X 1415; 2 spec. loc. A – 12.7 m (Fig. 90a-b – BSPG 1984 X 1410; Fig. 86a-b – BSPG 1984 X 1416; 22 spec. not fig. – BSPG BSPG 1984 X 1411). 1984 X 1417); 3 spec. loc. A – 6.0 m (BSPG 1984 X 1418); 4 spec. loc. A – 9.6 m (BSPG 1984 X 1419); 5 spec. lo c. A – Tentatively assigned specimens: 4 specimens. 12.7 m (BSPG 1984 X 1420); Thanetian (P4), upper pseudomenardii zone: Kroisbach: loc. Kch 1 Thanetian (P4), upper pseudomenardii zone: Kroisbach, 7 speci- (Fig. 91a-b – BSPG 1984 X 1412; Fig. 92 – BSPG 1984 X 1413; mens: loc. Kch 1 (Fig. 87 – BSPG 1984 X 1421; Fig. 88 – BSPG BSPG 1984 X 1414). 1984 X 1422), 5 spec. not fig. – BSPG 1984 X 1423); Name: Fro m no vellus (Latin) = new, mo dern, referring to Thanetian (P5), velascoensis zone: Kroisbach, 1 specimen: loc. Kch the mo dern lo o ks o f the o toliths o f the species. 11b (BSPG 1943 II 723; ex 515).

Diagnosis. Elo ngate o tolith; O L : OH = 1.7-1.8. Dorsal Description. Medium sized, elongate otoliths to about - - rim bro adly undulating in large specimens; ro strum long. 3.0 mm length in Kressenberg. O L : OT = (1.65 )1.8 2.1. Cauda long, narrow, markedly downturned at tip; ostium O H : OT abo ut 2.5. Anterio r rim with mo derately lo ng widened with do rsal margin extending o ver anterio r-do rsal ro strum with po inted tip; no o r very feeble excisura or anti- rim o f o tolith. Inner face co nvex; o uter face flat. ro strum; do rsal rim irregularly curved, undulating, highest at its middle; po sterio r rim ro unded, so metimes slightly Description. Medium sized, elongate otoliths to about do rsally shifted; ventral rim regularly and gently curved, 3.5 mm length. O H : OT abo ut 2.8. Anterio r rim with long deepest at its middle, crenulated. ro strum with narro w, ro unded tip; no excisura o r antiro strum; Inner face moderately convex, not twisted along lo ng do rsal rim shallo w, undulating with bro ad medio do rsal axis. Sulcus slightly supramedian, no t inclined, slightly lobe and postdorsal concavity, both increasing in intensity deepened. O stium slightly widened, its do rsal margin slightly with size; po sterio r rim do rsally shifted, ro unded; ventral extending. Cauda narrow, long, with slightly downturned rim regularly and gently curved, deepest at its middle. tapering tip terminating at so me distance fro m po sterio r tip Rims smo o th. o f o tolith. Do rsal wide, indistinct; no ventral furro w. Inner face markedly convex, not twisted along long axis. O uter face about as strongly convex as inner face, with Sulcus slightly supramedian, no t inclined, slightly deepened. some ornamentation near rims.

Werner SCHWARZHANS 27

Variability. O toliths o f P. po stangulatus show a rather Discussion. For comparison see above to P. no ve llus. Pa r- wide variatio n in the expressio n o f the do rsal rim and the aulo pus po stangulatus is no t o nly kno wn fo r its wide degree O L : OH ratio. I have figured here a specimen which is o f variability (see abo ve), but also fo r its wide distributio n particularly co mpressed, (Fig. 85; O L : O H = 1.65) to dem- ranging fro m No rth America to the No rth Sea Basin (Den- onstrate the extreme that can be expected in the species. mark), Bavaria and Ukraine. It is also o ne o f the very few More “typical” and slender specimens have been figured species known to occur across the K-T boundary, namely in SCHWARZHANS 2003, 2004 and 2010a. fro m the Late Cretaceo us (Maastrichtian) and Danian to Thanetian o f Bavaria and Austria.

3.6 Order Myctophiformes

Family indet. 1984 X 1430; Fig. 99 – BSPG 1984 X 1431; Fig. 98 – BSPG 1984 X 1432; 12 spec. not fig. – BSPG 1984 X 1433; 4 spec. Genus Bavariscopelus SCHWARZHANS 2010 no t fig. – BSPG 1943 II 430).

Bavariscopelus bispinosus SCHWARZHANS 2010 Description. Moderately elongate and moderately large - (Figs. 93-99) specimens to nearly 3 mm length. O L : OH = 1.35 1.5 (rarely to 1.6); O H : OT = 2.5-3.0. Rostrum massive, long, - 2003 genus Myctophidarum sp. – SCHWARZHANS: fig. 17G-I with blunt tip, about 20 25 % of O L. Dorsal rim high, gen- 2010a Bavarisco pe lus bispino sus SCHWARZHANS – figs. 64-71 tly curving o r o ften with bro ad po stdo rsal lo be; po sterio r rim bro adly ro unded; ventral rim shallo w, o ften with little Material: 65 specimens. po stventral denticle and rarely with very small preventral Danian (P1b), pse udo bullo ide s zone: Kressenberg, 4 specimens: lo c. B3 (Fig. 96a-b – BSPG 1984 X 1424; 3 spec. not fig. – denticle. Excisura and antiro strum feeble. Rims smo oth; BSPG 1984 X 1425); do rsal rim crenulated in small specimens. and Oichinger Graben, 1 specimen: loc. N 1 (BSPG 1943 II 521); Inner face flat, with slightly supramedian, lo ng, slightly Danian (P1c), trinidade nsis zone: Kressenberg, 40 specimens: 26 deepened sulcus. O stium slightly wider than cauda, both spec. loc. A – 2.5 m (BSPG 1984 X 1426); 1 spec. lo c. A – abo ut equal in length (CaL : OsL = 1.0-1.2). O stial and 9.6 m (BSPG 1984 X 1427); 11 spec. loc. A – 12.7 m (BSPG caudal co lliculi clearly separated; caudal co lliculum with 1984 X 1428); ventral ridge similar to the ventral pseudo co lliculum of and Oichinger Graben, 2 specimens: loc. N 2 (Fig. 93a-c – myctophid o toliths. Do rsal depressio n wide; ventral furro w BSPG 1943 II 520; Fig. 95 – BSPG 1943 II 743; ex 520); Thanetian (P4), upper pseudomenardii zone: Kroisbach, 20 speci- faint o r absent. mens: loc. Kch 1 (Fig. 94 – BSPG 1984 X 1429; Fig. 97 – BSPG O uter face convex, with postcentral umbo, smooth.

b a 93 94 c

97 a 96 b

95 98 99

Fig. 93-99. Bavariscopelus bispinosus SCHWARZHANS 2010. 96, Kressenberg, Danian (P1b), BSPG 1984 X 1424. 93, 95, O ichinger Graben, Danian (P1c); 93, BSPG 1943 II 520; 95 BSPG 1943 II 743; ex 520. 94, 97-99, Kro isbach, Thanetian (P4); 94, BSPG 1984 X 1429; 97, BSPG 1984 X 1430; 99, BSPG 1984 X 1431; 98, BSPG 1984 X 1432. – 20 × .

Palaeo Ichthyologica 12 28

100 101 a 103 102 b

104 105 106

ab 109 108 107

Fig. 100-106. Bavariscopelus parvinavis n. sp. Kressenberg, Danian (P1c). 102, Ho lo type, BSPG 1984 X 1434. 100-101, 103-106, Paratypes; 100, BSPG 1984 X 1435; 101, BSPG 1984 X 1436; 103, BSPG 1984 X 1437; 104, BSPG 1984 X 1438; 105, BSPG 1984 X 1439; 106, BSPG 1984 X 1440. – 20 × . Fig. 107-109. Danoscopelus schnetleri n. gen. (SCHWARZHANS 2003). Kro isbach, Thanetian (P4). 107, BSPG 1984 X 1445; 108, BSPG 1984 X 1446; 109, BSPG 1984 X 1447. – 30 × .

Discussion: Bavarisco pe lus bispino sus was first de- Danian (P2), uncinata zone: Kressenberg, 1 specimen: loc. A – scribed fro m the Cretaceo us (Maastrichtian) o f Bavaria 2.0 m (BSPG 1984 X 1444). by SCHWARZHANS 2010a and represents one of the few Name: Co mbinatio n o f parvus (Latin) = small, and navis species evident acro ss the K-T bo undary. Its relation to the (Latin) = ship, referring to the shape o f the o toliths. Myctophidae is based on the general appearance of the otolith and the development of the caudal colliculum, but Diagnosis. Small, thin o toliths. O L : OH = 1.3-1.4; the lack of a clearly developed caudal pseudocolliculum OH : OT = 3.5-4.0. Rostrum massive, blunt, about 25- and the cauda being longer than the ostium has now 30 % of O L. Dorsal rim short, much expanded postdorsally. led me to place it in an open familial taxonomic po sitio n Ventral rim shallo w, preventrally pro no unced. Ventral furro w within the Myctophifo rmes (instead o f the Myctophidae as bro ad, distinct. in SCHWARZHANS, 2010a, see also discussio n in chapter Description. Moderately elongate, small specimens to 5.6 later). slightly abo ve 1.5 mm length. Rostrum massive, lo ng, with blunt tip, about 25-30 % of O L. Dorsal rim high, with massive pro jectio n at abo ut middle o f sho rt do rsal rim; poste- Bavariscopelus parvinavis n. sp. rio r rim bro adly ro unded; ventral rim shallo w, so mewhat - (Figs. 100 106) undulating, deepest anterio r o f the middle. Excisura and antiro strum absent o r very feeble. Rims sharp. - Holotype: Fig. 102a b, BSPG 1984 X 1434. Inner face flat, with slightly supramedian, lo ng, slightly Type location: Kressenberg, Bavaria, location A – 12.7 m. deepened sulcus. O stium slightly wider than cauda, both abo ut equal in length (CaL : OsL = 1.0-1.1). O stial and Type formation: O iching Formation, Danian (P1c), trinidad- caudal co lliculi clearly separated; caudal co lliculum with ensis zone. ventral ridge similar to the ventral pseudo co lliculum of Paratypes: 6 specimens. mo dern myctophid o toliths. Do rsal depressio n small, in- Danian (P1c), trinidade nsis zone: Kressenberg: loc. A – 12.7 m distinct; ventral furro w bro ad, distinct, clo se to ventral rim (same data as ho lo type) (Fig. 100 – BSPG 1984 X 1435; o f o tolith. Fig. 101 – BSPG 1984 X 1436; Fig. 103 – BSPG 1984 X 1437; O uter face nearly flat, smo o th. Fig. 104 – BSPG 1984 X 1438; Fig. 105 – BSPG 1984 X 1439; Fig. 106 – BSPG 1984 X 1440). Discussion: Bavarisco pe lus parvinavis differs fro m the larger contemporaneous B. bispinosus in the higher dorsal Further material: 31 specimens. - Danian (P1c), trinidade nsis zone: Kressenberg, 30 specimens: rim with its expansio n, the even lo nger ro strum (25 30 % 7 spec. loc. A – 6.0 m (BSPG 1984 X 1441), 1 spec. lo c. A – o f O L vs 20-25 %), the thin appearance (O H : OT = 3.5- 11.6 m (BSPG 1984 X 1442), 22 spec. loc. A – 12.7 m (same 4.0 vs 2.5-3.0) and the distinct and broad ventral furrow data as ho lo type) (BSPG 1984 X 1443); o n the inner face. It is also remarkable fo r the smaller size it attains as co mpared to B. bispinosus. Ano ther similar

Werner SCHWARZHANS 29 species is Neoscopelus? nuusuaqensis SCHWARZHANS but a true representative o f that family is already recorded 2004 from the Paleocene of western Greenland, which contemporaneously from the Paleocene of western Green- however has a less prominent rostrum and a shallower land – Neoscopelus nuussuaquensis SCHWARZHANS 2004. and longer dorsal rim. Bavarisco pe lus on the other hand shows more resemblance to myctophids. The first certain myctophid o tolith is repo rted fro m the Late Paleo cene o f so uthern Australia – Eokre fftia Genus Danoscopelus n. gen. prediaphus SCHWARZHANS 1985a. Species: Danoscopelus is mo no specific with D. schne tle ri Type species: Genus Myctophidarum schne tle ri SCHWARZ- known from the Selandian of Denmark and the Thanetian HANS 2003 o f Austria. Name: Referring to the first descriptio n o f these o toliths from the Paleocene of Denmark. Danoscopelus schnetleri (SCHWARZHANS 2003) Diagnosis. A fo ssil o tolith-based genus o f the o rder (Figs. 107-109) Myctophifo rmes witho ut fo rmal familial asso ciatio n with the following combination of characters: Elongate o toliths 2003 genus Myctophidarum schne tle ri SCHWARZHANS – with mo derately lo ng ro strum. Do rsal and ventral rims fig. 17A-F, J shallo w, po sterio r rim blunt. Sulcus anterio rly o pen, wide, long. O stium about as wide and long as cauda. Caudal Material: Thanetian (P4), upper pseudomenardii zone: Krois- bach, Austria, 6 specimens: loc. Kch 1 (Fig. 107 – BSPG co lliculum lo nger than o stial co lliculum, with sharp and 1984 X 1445; Fig. 108a-b – BSPG 1984 X 1446; Fig. 109 – elevated ventral margin resembling the caudal pseudocol- BSPG 1984 X 1447; 3 spec. no t fig. – BSPG 1984 X 1448). liculum o f mo dern myctophids, anterio r-ventrally so mewhat widened. Inner face flat with distinct ventral furrow. O uter Description. Small, elo ngate o toliths pro bably no t exceed- face slightly co nvex. ing 2.5 mm length. O L : O H = 1.7-1.9; O H : OT about 3.0. Rostrum mo derately lo ng, mo derately po inted (no t preserved Discussion. O toliths o f Danoscopelus resemble those of re- in the specimens fro m Bavaria). Do rsal and ventral rims cent species of the families Myctophidae and Neosco pelidae. shallo w, o ften crenulated; po sterio r rim blunt. From Myctophidae they differ in the absence of a clearly Inner face flat, with bro ad, shallo w sulcus divided in defined caudal pseudocolliculum, while those of the Neo- almost equally wide ostium and cauda. O stium slightly scopelidae are less elongate and thinner. Further more, oto- deepened and shorter. Dorsal depression narrow; ventral liths o f the two families do no t exhibit the anterio r-ventrally furro w mo stly distinct, sho rt. widened cauda, which is diagnostic for Danoscopelus. This O uter face almo st flat, smo o th o r with sho rt radial fur- character as well as the elongate outline of the oto lith is ro ws near rims. also the main difference to Bavarisco pe lus. Like with Bavarisco pe lus, the relatio nship o f Danoscope- Discussion. This is a fairly co mmo n species in the Selan- lus is uncertain. The general appearance with the narro w dian of Denmark. The few finds reported here from the o stium and the elevated ventral margin o f the elevated Thanetian o f Bavaria are mo stly fro m very small specimens caudal co lliculum are interpreted as myctophifo rm charac- and inco mpletely preserved. ters. The o utline o f Danoscopelus resembles neoscopelids,

3.7 Order Gadiformes

Family Merlucciidae Further material: 13 specimens. Thanetian (P4), upper pseudomenardii zone: Kroisbach, 4 specimens: Genus Palaeogadus RATH 1859 lo c. Kch 1 (same data as ho lo type) (BSPG 1984 X 1458); Thanetian (P5), velascoensis zone: Kroisbach, 9 specimens: 5 spec. Palaeogadus? bratishkoi n. sp. loc. Kch 11b (BSPG 1943 II 724; ex 515), 3 spec. lo c. Kch (Figs. 110-118) 12 (BSPG 1943 II 528; ex 516), 1 spec. loc. Kch 12a (BSPG 1943 II 517).

Holotype: Fig. 110a-c, BSPG 1984 X 1449. Name: In ho no r o f Andrei Bratishko , Kiev, Ukraine, in Type location: Kro isbach, Austria, lo catio n Kch 1. recognition of his contribution to the knowledge of fo ssil o toliths fro m Ukraine. Type formation: O iching Formation, Thanetian (P4), upper - pseudomenardii zone. Diagnosis. Moderately compressed otoliths; O L : O H = 1.5 1.7. Do rsal rim gently curved; ventral rim mo derately deep. Paratypes: 8 specimens. Rims finely crenulated. CaL : O sL abo ut 1.0 in specimens to Thanetian (P4), upper pseudomenardii zone: Kroisbach: loc. Kch 1 2 mm length, 1.3-1.8 in specimens from 2 to 3 mm length. (same data as ho lo type) (Fig. 111 – BSPG 1984 X 1450; Co llum width less than co llum height; o stial co lliculum Fig. 112 – BSPG 1984 X 1451; Fig. 113 – BSPG 1984 X 1452; Fig. 114 – BSPG 1984 X 1453; Fig. 115 – BSPG 1984 X 1454; anteriorly reduced; no pseudocolliculum. Fig. 116 – BSPG 1984 X 1455; Fig. 117 – BSPG 1984 X 1456; Description. Small, oval shaped otoliths to about 3.5 mm Fig. 118 – BSPG 1984 X 1457). length. O H : O T abo ut 2.8. Do rsal rim gently curved, highest anterio rly, witho ut pro minent angles; ventral rime regularly curved, deepest anterio r o f the middle; anterio r rim broadly

Palaeo Ichthyologica 12 30

ba 110 c 111

113 114

112

118 116 117 115

a a 119 b b 120

Fig. 110-118. Palaeogadus? bratishkoi n. sp. Kro isbach, Thanetian (P4). 110, Ho lo type, BSPG 1984 X 1449. 111-118, Paratypes; 111, BSPG 1984 X 1450; 112, BSPG 1984 X 1451; 113, BSPG 1984 X 1452; 114, BSPG 1984 X 1453; 115, BSPG 1984 X 1454; 116, BSPG 1984 X 1455; 117, BSPG 1984 X 1456; 118, BSPG 1984 X 1457. – 20 × . Fig. 119-120. Gadidae indet. Kro isbach, Thanetian (P4). 119, BSPG 1943 II 701; ex 430; 120, BSPG 1943 II 744; ex 430. – 30 × .

ro unded; po sterio r tip slightly mo re pro jecting. All rims high as 1.8. The latter is a typical ontogenetic development finely crenulated. observed in many gadid species and indicates diagno stically Inner face slightly co nvex with supramedian sulcus with mature specimens. Therefore, P.? bratishko i is interpreted as indistinct anterior and posterior opening. Cauda about 50 % a small gadifo rm species and no t as juvenile o toliths. longer than ostium in specimens of 2 mm and larger, about Discussion. Palae o gadus? bratishko i resembles P.? antiqu- same length as o stium in smaller specimens. Co lliculi well us SCHWARZHANS & BRATISHKO 2011 fro m the Paleo cene defined, slightly deepened, o stial co lliculum anterio rly re- o f Ukraine, differing mainly in the mo re co mpressed shape duced. Collum higher than wide, with convex lower margin, (OL : OH = 1.7-1.7 vs 1.9-2.0), the shape and ornamenta- but without pseudocolliculum. Dorsal depression moderately tion of the dorsal rim and the cauda being longer than the wide; ventral furro w mo stly distinct, clo se to ventral rim o f o stium (vs abo ut equal in length) with the o stial co lliculum o tolith. anterio rly reduced. Also similar is Archaemacruroides orna- O uter face slightly co nvex with numero us radial furro ws, tus STINTON 1965 (interpreted as a very basal gadifo rm particularly near o tolith rims. o f the family Euclichthyidae in SCHWARZHANS 2004) in Ontogeny. O toliths o f P.? bratishko i show a remarkable shape and the anterio rly reduced o stial co lliculum, but ontogenetic change at about 2 mm length in the index these o toliths sho w a mo re intense marginal crenulatio ns, O L : OH increasing fro m 1.5-1.6 to 1.6-1.7 and the in- a do rsal rim with pre- and po stdo rsal angles (vs gently creasing length o f the caudal co lliculum best reflected in curved without angles) and colliculi of equal length also at the increase of the index CaL : OsL from about 1.0 to as sizes of 3.5 mm length.

Werner SCHWARZHANS 31

Family Gadidae Discussion. The two small specimens are slender (O L : O H = 1.9-2.1), with a rather flat do rsal rim, deeply curved Genus indet. ventral rim, which is deepest anterio r o f its middle, a broadly rounded anterior tip and a dorsally pronounced Gadidae indet. po sterio r tip. O stium and cauda are o f similar size with (Figs. 119-120) two extremely small, reduced co lliculi, which likewise are of similar size. The co llum is wide and sho ws a marked ventral Material: Thanetian (P4), upper pseudomenardii zone: Kroisbach, pseudo co lliculum, a character which is very rarely found Austria, 2 specimens: loc. Kch 1, of a length of about 1.5 mm (Fig. 119a-b – BSPG 1943 II 701; ex 430; Fig. 120a-b – BSPG in Paleo cene gadifo rm o toliths. These are clearly juvenile 1943 II 744; ex 430). otoliths representing an unknown Paleocene species.

3.8 Order Ophidiiformes

Suborder Ophidioidei 6 mm length. O H : OT about 2.5. Dorsal rim shallow, nearly horizontal, with weak predorsal projection and obtuse Family Ophidiidae po stdo rsal angle. Anterio r tip bro ad, o btusely po inted at Genus Ampheristus KÖNIG 1825 its middle abo ut level o f lo wer sulcus margin. Ventral rim regularly curved, mo stly no t very deep, deepest anterio r Ampheristus neobavaricus n. sp. o f its middle belo w o stium: Posterio r tip po inted at level - (Figs. 121 129) of caudal tip. Inner face slightly co nvex with median to slightly supra- - Holotype: Fig. 124a c, BSPG 1984 X 1459. median sulcus. Sulcus clo sely reaching anterio r tip o f o tolith, Type location: Kressenberg, Bavaria, location A – 12.7 m. but terminating at so me distance fro m po sterio r tip. O stium about 50 % longer and wider than cauda, with shallo w o stial Type formation: O iching Formation, Danian (P1c), trini- colliculum. Cauda deepened with rounded and slightly bent dadensis zone. tip. Do rsal depressio n small, narro w; ventral field smooth Paratypes: 8 specimens. with o nly faint indicatio n o f ventral furro w. Danian (P1b), pse udo bullo ide s zone: Kressenberg, 1 specimen: O uter face flat, smooth in large and with some radial lo c. B3 (Fig. 125 – BSPG 1984 X 1460); o rnamentatio n in small specimens. Danian (P1c), trinidade nsis zone: Kressenberg, 6 specimens: 4 spec. loc. A – 2.5 m (Fig. 129 – BSPG 1984 X 1461; Fig. 123 Discussion. Amphe ristus ne o bavaricus resembles well – BSPG 1984 X 1462; Fig. 126 – BSPG 1984 X 1463; Fig. 122 A. bavaricus and A. brevicauda fro m the Maastrichtian o f – BSPG 1984 X 1464); 1 spec. loc. A – 9.6 m (Fig. 128 – BSPG Bavaria. It differs fro m bo th species in the shallo w, nearly 1984 X 1465); 1 spec. loc. A – 12.7 m (same data as ho lo type) ho rizo ntal do rsal rim (vs inclined) with its weak predo rsal - (Fig. 127a b – BSPG 1984 X 1466); pro jectio n and the slightly bend cauda with its ro unded Thanetian (P4), upper pseudomenardii zone: Kroisbach, 1 specimen: tip terminating at so me distance fro m the po sterio r tip of loc. Kch 1 (Fig. 121 – BSPG 1984 X 1467). the o tolith. Amphe ristus ne o bavaricus is interpreted as the Further material: 273 specimens. descendant of A. bavaricus in the Paleocene. Danian (P1b), pse udo bullo ide s zone: Kressenberg, 3 specimens: Amphe ristus ne o bavaricus co nstitutes the seco nd mo st lo c. B3 (BSPG 1984 X 1468); co mmo n species at Kressenberg, but is rare at Kro isbach. Danian (P1c), trinidade nsis zone: Kressenberg, 265 specimens: 217 spec. loc. A – 2.5 m (BSPG 1984 X 1469); 8 spec. loc. A – 2.5 m (BSPG 1984 X 1470); 1 spec. loc. A – 3.0 m (BSPG 1984 X 1471); 8 spec. loc. A – 6.0 m (BSPG 1984 X 1472); Palaeomorrhua GAEMERS & SCHWARZHANS 1973 6 spec. loc. A – 8.3 m (BSPG 1984 X 1473); 13 spec. lo c. A – 9.6 m (BSPG 1984 X 1474); 12 spec. loc. A – 12.7 m (same Palaeomorrhua sp. data as ho lo type) (BSPG 1984 X 1475); (Fig. 130) Daniian (P2), uncinata zone: Kressenberg, 4 specimens: loc. A – 2.0 m (BSPG 1984 X 1476); Material: 1 specimen. Selandian (P3a), angulata zone: Kroisbach, 1 specimen: loc. 1.2 m Danian (P1c), trinidade nsis zone: Kressenberg: loc. A – 2.5 m N of Kch 4 (BSPG 1943 II 709; ex 504). (Fig. 130a-c – BSPG 1984 X 1477).

Name: As a derived name from A. bavaricus (KOKEN Discussion. The bro ken anterio r half o f a single large 1891) fro m the Maastrichtian o f Bavaria, o f which it is specimen that o riginally must have been larger than considered as a descendant in Paleocene times. 10 mm length. It is massive with an O H : OT o f abo ut 2.5, Diagnosis. Elo ngate o toliths with bro ad anterio r and a strongly convex inner face, flat outer face, and shallo w po inted po sterio r tips; O L : OH = 1.8-2.0. Do rsal rim flat o stium and o stial co lliculum resembling well o toliths of the with weak predo rsal pro jectio n. O sL : CaL = 1.5-1.8. Cau- genus Palae o mo rrhua. There are two species so far known dal colliculum deepened and slightly bent downward. within this genus – P. faba (KOKEN 1884), the type species from the O ligocene of the North Sea Basin and P. bulbus Description. Large o toliths attaining sizes up to 9 mm (NO LF 1978) fro m the Thanetian o f Belgium. length, though most specimens are in the range of 4 to

Palaeo Ichthyologica 12 32

122

121

123 124 c

125

a 126 127 b

128 129

Fig. 121-129. Ampheristus neobavaricus n. sp. Kressenberg, Danian (P1c). 124, Holotype, BSPG 1984 X 1459. 121-123, 125-129, Paratypes. 125, Kressenberg, Danian (P1b), BSPG 1984 X 1460; 122-123, 125-129, Kressenberg, Danian (P1c); 122, BSPG 1984 X 1464; 123, BSPG 1984 X 1462; 126, BSPG 1984 X 1463; 127, BSPG 1984 X 1466; 128, BSPG 1984 X 1465; 129, BSPG 1984 X 1461. 121, Kro is bach, Thanetian (P4), 1943 II 709; ex 504. – 10 × .

Suborder Bythitioidei 504); and O ichinger Graben 1 specimen: loc. N 4 (BSPG 1943 II 735; ex 519); Family Bythitidae Thanetian (P4), upper pseudomenardii zone: Kroisbach, 33 specimens: 32 spec. loc. Kch 1 (BSPG 1984 X 1487), 1 spec. loc. Genus Bidenichthys BARNARD 1934 8 m S o f Kch1 (BSPG 1943 II 445); Thanetian (P5), velascoensis zone: Kroisbach, 1 specimen: loc. Kch Bidenichthys lapierrei (NOLF 1978) 11b (BSPG 1943 II 726; ex 515). (Figs. 132-138) Diagnosis. Elo ngate, mo derately thick o toliths; O L : OH = 1.75-1.95 increasing with size; O H : OT = 2.2-2.5. Dorsal 1978 O gilbia lapie rre i NO LF – pl. 2, figs. 2-3 rim symmetrical, with sho rt ho rizo ntal median po rtion (about Material: 120 specimens. 40-55 % of O L). Sulcus medial, moderately short, wide, di- Danian (P1c), trinidade nsis zone: Kressenberg, 78 specimens: vided in a large and wide ostium and a much smaller cauda; 60 spec. loc. A – 2.5 m (Fig. 134a-b – BSPG 1984 X 1478; OL : SuL = 1.6-1.8; SuL : SuH = 2.8-3.3; OCL : CCL = 2.5- Fig. 136 – BSPG 1984 X 1479; Fig. 137 – BSPG 1984 X 1480; - Fig. 138 – BSPG 1984 X 1481; 54 spec. not fig. – BSPG 3.0; OCH : CCH = 1.5 1.8. O uter face convex. 1984 X 1482; 2 spec. not fig. – BSPG 1984 X 1483), 2 spec. Description. Medium sized otoliths up to about 3.7 mm loc. A – 6.0 m (BSPG 1984 X 1484), 5 spec. loc. A – 9.6 m (BSPG length. Do rsal rim symmetrical, with inclined anterior and 1984 X 1485), 11 spec. loc. A – 12.7 m (BSPG 1984 X 1486); po sterio r and sho rt ho rizo ntal median sectio n. Ventral rim Selandian (P3a), angulata zone: Kroisbach, 8 specimens: 7 spec. loc. 1.2 m N of Kch 4 (Fig. 132 – BSPG 1943 II 745; ex 504; regularly curved; anterio r and po sterio r tip almo st symmetri- Fig. 133a-b – BSPG 1943 II 746; ex 504; Fig. 135 – BSPG cal, with ro unded median tip. All rims smo o th, irregularly 1943 II 747; ex 504; 4 spec. not fig. – BSPG 1943 II 711; ex crenulated in specimens smaller than 2.5 mm length.

Werner SCHWARZHANS 33

ba a 130 131 c b

a 132 133 b

a 134 b

135

136 137 138

Fig. 130. Palaeomorrhua sp. Kressenberg, Danian (P1c), BSPG 1984 X 1477. – 10 ×. Fig. 131. Ogilbia luzanensis SCHWARZHANS & BRATISHKO 2011. Kressenberg, Danian (P1c), BSPG 1984 X 1488. – 20 × . Fig. 132-138. Bidenichthys lapierrei (NOLF 1978). 134, 136-138, Kressenberg, Danian (P1c); 134, BSPG 1984 X 1478; 136, BSPG 1984 X 1479; 137, BSPG 1984 X 1480; 138, BSPG 1984 X 1481. 132-133, 135, Kro isbach, Selandian (P3a); 132, BSPG 1943 II 745; ex 504; 133, BSPG 1943 II 746; ex 504; 135, BSPG 1943 II 747; ex 504. – 20 × .

Inner face mildly convex, about the same as outer face Outer face convex, smooth. o r less. Sulcus medial, shallo w, with large o stium and small Discussion. SCHWARZHANS & BRATISHKO (2011) elabo - cauda. O stium often with ridge connecting to anterior tip of rated abo ut the reco gnitio n o f two superficially similar o tolith. Do rsal depressio n wide, narro w, indistinct; ventral bythitid species in the European Paleocene, namely Bide n- furro w mo stly indistinct, clo se to ventral rim o f o tolith.

Palaeo Ichthyologica 12 34 ichthys lapie rre i and O gilbia luzane nsis SCHWARZHANS Material: 1 specimen. & BRATISHKO 2011 (see also belo w), which were no t Danian (P1c), trinidade nsis zone: Kressenberg: loc. A – 2.5 m - recognized before. The main differences of B. lapie rre i are (Fig. 131a b – BSPG 1984 X 1488). the outer face being equally convex than the inner face Diagnosis. Elo ngate, mo derately thin o toliths; O L : OH o r mo re co nvex (vs less co nvex to flat), the less elongate = 1.9-2.2 increasing with size; O H : OT = 2.0-2.5. Dor- shape (OL : OH = 1.75-1.95 vs 1.9-2.2) the medial posi- sal rim with lo ng ho rizo ntal median po rtio n (abo ut 55- tio n o f the sulcus (vs anterio rly shifted, the narrower sulcus 65 % o f O L). Sulcus anterio rly shifted, mo derately sho rt, (SuL : SuH = 2.8-3.3 vs 3.7-4.4) and the shorter median narrow, divided in a long ostium and a short cauda; do rsal po rtio n (40-55 % of O L vs 55-65 % of O L). Most of OL : SuL = 1.7-2.0; SuL : SuH = 3.7-4.4; OCL : CCL = 2.6- these characters are rather subtle in nature, o verlapping in 2.8; OCH : CCH = 1.4-1.8. O uter face flat o r slightly co nvex. part and hence not readily recognized in all instances. In specimens smaller than 2 mm o f length particularly distinc- Description. Medium sized otoliths up to about 4.0 mm tio n o f the two species may no t always be po ssible. length. Do rsal rim mo re steeply inclined anterio rly than Bide nichthys lapie rre i is a common species both at po sterio rly and with lo ng ho rizo ntal median sectio n. Ventral Kressenberg and Kro isbach. rim shallo w, regularly curved; anterio r mo derately pointed, po sterio r tip slightly expanded, o ften with small po stdo rsal co ncavity. All rims smo o th. Genus Ogilbia JORDAN & EVERMANN 1898 Inner face moderately convex. Sulcus medial, shallow, with long ostium and short cauda. O stium often with ridge Ogilbia luzanensis SCHWARZHANS & BRATISHKO 2011 co nnecting to anterio r tip o f o tolith. Do rsal depressio n nar- (Fig. 131) ro w, indistinct; ventral furro w mo stly feeble, clo se to ventral rim o f o tolith. 2003 Bide nichthys lapie rre i (NO LF 1978) – SCHWARZHANS: O uter face flat o r slightly co nvex, but always less than fig. 29 A-J inner face, smooth. 2004 Bide nichthys lapie rre i (NO LF 1978) – SCHWARZHANS: For co mpariso n with the sympatric Bidenichthys fig. 9C-F Discussion. 2011 O gilbia luzane nsis SCHWARZHANS & BRATISHKO – lapie rre i see above. O gilbia luzane nsis is rare in Kressen- fig. 9 A-F, 1 5 H berg, Bavaria and so far absent fro m Kro isbach, Austria, while it represents the do minant bythitid in the Paleocene of Denmark, West Greenland and Ukraine.

3.9 Order Lophiiformes

Suborder Ogcocephaloidei Type location: Kro isbach, Austria, lo catio n Kch 1. Family Ogcocephalidae Type formation: O iching Formation, Thanetian (P4), upper pseudomenardii zone. Genus indet. Paratype: 1 specimen. Ogcocephalus? semen n. sp. Thanetian (P4), upper pseudome nardii zone: Kroisbach: loc. Kch 1 (same data as ho lo type) (Fig. 140 – BSPG 1984 X 1490). (Figs. 139-140) Name: Fro m semen (Latin) = seem, referring to the ap- Holotype: Fig. 139a-d, BSPG 1984 X 1489. pearance of the otolith.

ba d 139

140

Fig. 139-140. Ogcocephalus? semen n. sp. Kro isbach, Thanetian (P4); 139, Ho lo type, BSPG 1984 X 1489; 140, Paratype, BSPG 1984 X 1490. – 30 × .

Werner SCHWARZHANS 35

Diagnosis. Elo ngate, regularly o val o toliths with bro adly and po sterio rly po inted co lliculum. Few radial furrows on ro unded anterio r and po inted po sterio r tips; O L : O H = 1.6- do rsal field extending fro m crenulatio ns o f o tolith rim; in- 1.7. Do rsal rim bro adly crenulated. Small, undivided sulcus distinct do rsal depressio n; weak ventral furro w near ventral at center of inner face; O L : SuL = 2.3-2.7. Single colliculum rim o f o tolith. anterio rly po inted. O uter face slightly co nvex, smo o th except fo r few radial furro ws. Description: Small o toliths with regular o val o utline reaching abo ut 2 mm length. O H : OT = 2.5. Do rsal rim irregu- Discussion. Ogcocephalus? semen is the earliest reco rded larly and bro adly crenulated; ventral rim shallo w, regular, lo phiifo rm o tolith, and it already exhibits all typical charac- smo o th. Anterio r and po sterio r tips bro adly ro unded; ters o f such o toliths like the small, central sulcus with the anterio r tip higher than po sterio r tip. very simple “reduced” morphology and the outline of the Inner face nearly flat with centrally po sitio ned small, o tolith. O therwise, several species are kno wn since Eocene slightly deepened, undivided o tolith with unifo rm anterio rly times (see SCHWARZHANS 2007).

3.10 Order Beryciformes

Suborder Berycoidei Thanetian (P4), upper pseudomenardii zone: Kroisbach, 2 specimens: loc. Kch 1 (Fig. 143 – BSPG 1984 X 1495; Fig. 144 Family Berycidae – BSPG 1984 X 1496). Genus Centroberyx GILL 1862 Further material: 35 specimens. Danian (P1b), pse udo bullo ide s zone: Kressenberg, 1 specimen: Centroberyx apogoniformis n. sp. lo c. B3 (BSPG 1984 X 1497); Danian (P1c), trinidade nsis zone: Kressenberg, 31 specimens: (Figs. 141-146) 29 spec. loc. A – 2.5 m (same data as holotype) (BSPG 1984 X 1498); 2 spec. loc. A – 12.7 m (BSPG 1984 X 1499); 2004 Centroberyx sp. – SCHWARZHANS: fig. 10F-H Thanetian (P4), upper pseudomenardii zone: Kroisbach, 3 speci- 2011 Centroberyx sp. – SCHWARZHANS & BRATISHKO : fig. mens: loc. Kch 1 (BSPG 1943 II 702; ex 430). 11G-H Name: Referring to the percifo rm genus Apogon, to which Holotype: Fig. 142, BSPG 1984 X 1491. the otoliths resemble in their general appearance. Type location: Kressenberg, Bavaria, lo catio n A – 2.5 m. Diagnosis. Mo derately elo ngate o toliths, with o val shape; Type formation: O iching Formation, Danian (P1c), trinidad- OL : OH = 1.4-1.5. O toliths thin with convex inner face; ensis zone. OH : OT = 3.0-3.5. Dorsal rim shallow, reduced above caudal tip. O stium about as long as cauda; CaL : O sL = 0.9- Paratypes: 5 specimens. Danian (P1c), trinidade nsis zone: Kressenberg, 3 specimens: 2 spec. 1.15. Caudal tip straight, tapering. - loc. A – 2.5 m (same data as holotype) (Fig. 141a c – BSPG Description. Large, o val shaped, rather thin o toliths pro b- 1984 X 1492; Fig. 145 – BSPG 1984 X 1493); 1 spec. lo c. A ably reaching about 10 mm length (single eroded speci- – 12.7 m (Fig. 146 – BSPG 1984 X 1494); men; holotype is 5.5 mm long). Dorsal rim shallow, witho ut

ba 141 142 c

143 144 145 146

Fig. 141-146. Centroberyx apogoniformis n. sp. 142, Kressenberg, Danian (P1c), Ho lo type, BSPG 1984 X 1491. 141, 143-146, Paratypes; 141, 145-146, Kressenberg, Danian (P1c); 141, BSPG 1984 X 1492; 145, BSPG 1984 X 1493; 146, BSPG 1984 X 1494. 143-146, Kro isbach, Thanetian (P4); 143, BSPG 1984 X 1495; 144, BSPG 1984 X 1496. – 10 × .

Palaeo Ichthyologica 12 36 prominent angles, reduced above caudal tip. Ventral rim newly extracted specimen of 6 mm length (WAM P.27210- moderately deep, gently and regularly curved, deepest at 004). Unfortunately, the checking of the early reco rds fro m its middle, witho ut pro minent angles. Anterio r and po sterio r 1980 cannot be verified due to lack of noting the co llectio n tips expanded, ro unded. All rims smo o th except slightly number o f the respective fishes. undulating do rsal rim in juveniles. In any case, the ontogenetic changes in Centroberyx oto- Inner face co nvex with distinctly supramedian sulcus. liths no w appear significantly less dramatic than reported in Sulcus anterio rly o pened, po sterio rly reaching clo se to 1980. This correction may have influence on the reco gnitio n po sterio r tip o f o tolith. O stium abo ut as lo ng as cauda and o f o tolith-based Centroberyx species in the European Eocene. about twice as wide, with shallow colliculum; cauda slightly Discussion. Ce ntro be ryx fragilis usually o ccurs sympatrically deepened with straight, tapering tip. Caudal colliculum with with C. integer, but in varying proportions. In Denmark, both distinct, crest-like ventral margin. Do rsal depressio n narro w, species are about equally common with C. fragilis prevailing small, indistinct; no ventral furro w. during Danian (SCHWARZHANS 2003). In West G reenland O uter face flat to slightly co ncave, smo o th. C. integer has not been observed (SCHWARZHANS 2004). Discussion. O toliths o f C. apogoniformis are readily distin- In Ukraine, C. fragilis is more common than C. integer guished from other contemporaneous Centroberyx species (SCHWARZHANS & BRATISHKO 2011). In Kro isbach, C. fra- by their elongate shape, the shallow dorsal rim and the gilis is much less common than C. integer, and in Kressen- lo ng o stium. This species no w represents the fo urth of the berg C. fragilis was not identified. The cause of the observed genus Centroberyx in the European Paleocene. variatio ns in abundance o f the two species mo st likely is o f enviro nmental nature. It appears that C. fragilis dominates in more shallow habitats and C. integer in deeper water Centroberyx fragilis SCHWARZHANS 2003 habitats, except fo r Kressenberg. Kressenberg represents a (Fig. 152) shallower environment than Kroisbach, but here C. integer is the o nly present species o f the two . 2003 Ce ntro be ryx fragilis SCHWARZHANS – fig. 34 A-J Ce ntro be ryx fragilis is best distinguished fro m C. in- 2004 Ce ntro be ryx fragilis SCHWARZHANS 2003 – SCHWARZ- te ge r by the thinner appearance and the bent caudal tip - HANS: fig. 12 A E observed in specimens above at least 5 mm length. Also , 2011 Ce ntro be ryx fragilis SCHWARZHANS – SCHWARZHANS C. fragilis is usually somewhat more elongate than C. inte - & BRATISHKO : fig. 10 A-I, 15 K ge r (OL : OH = 1.2-1.35 vs 1.1-1.25), but the single large Material: 2 specimens. specimen fro m Kro isbach is markedly mo re co mpressed Thanetian (P4), upper pseudomenardii zone: Kroisbach, 1 specimen: than usual at a ratio O L : OH of 1.1. lo c. Kch 1 (BSPG 1984 X 1500); Thanetian (P5), velascoensis zone: Kroisbach, 1 specimen: loc. Kch 14 (Fig. 152a-b – BSPG 1943 II 732; ex 518). Centroberyx integer (KOKEN 1885) Description. Large, high bodied, moderately thin otoliths (Figs. 147-151) reaching up to about 9 mm length. O L : OH = 1.2-1.35, but the large specimen from Kroisbach has only 1.1. 1885 O tolithus (Apo go nidarum) inte ge r KOKEN – pl. 5, fig. 27 OH : OT = 3.5-4.0. Dorsal rim high, undulating, usually 1978 Trachichthodes integer (KOKEN 1885) – NO LF: pl. 2, - with broad predorsal angle. Ventral rim deep, gently curved, figs. 4 6 deepest anterio r o f its middle, witho ut pro minent angles. 2003 Centroberyx integer (KOKEN 1885) – SCHWARZHANS: fig. 33 A-J Anterio r and po sterio r tips o btuse angular. 2011 Centroberyx integer (KOKEN 1885) – SCHWARZHANS Inner face convex with supramedian sulcus. Sulcus & BRATISHKO : fig. 11 A-C, 15 L anterio rly o pened, po sterio rly reaching clo se to po sterio r tip of otolith. O stium about twice as wide as cauda, with Material: 213 specimens. shallo w co lliculum; CaL : OsL = 1.1-1.25; cauda slightly Danian (P1b), pse udo bullo ide s zone: Kressenberg, 3 specimens: lo c. B3 (BSPG 1984 X 1501); deepened with bent, tapering tip. Caudal colliculum with Danian (P1c), trinidade nsis zone: Kressenberg, 197 specimens: distinct, crest-like ventral margin. Do rsal depression wide, 173 spec. loc. A – 2.5 m (Fig. 147 – BSPG 1984 X 1502; mo derately distinct; no ventral furro w. Fig. 149 – BSPG 1984 X 1503; Fig. 150 – BSPG 1984 X 1504; O uter face flat to slightly co ncave, smo o th. 170 spec. not fig. – BSPG 1984 X 1505; 3 spec. not fig. – BSPG 1984 X 1506); 1 spec. loc. A – 5.5 m (Fig. 148a-c – BSPG Ontogeny. SCHWARZHANS 1980 elaborated on the ex- 1984 X 1507); 10 spec. loc. A – 6.0 m (BSPG 1984 X 1508); trao rdinary o ntogenetic changes o bserved in o toliths o f the 1 spec. loc. A – 8.3 m (BSPG 1984 X 1509); 6 spec. lo c. A recent Centroberyx affinis (G ÜNTHER 1859) fro m so uthern – 9.6 m (BSPG 1984 X 1510); 3 spec. loc. A – 12.7 m Australia and New Zealand. While clearly the bending of the (Fig. 148a-c – BSPG 1984 X 1511); caudal tip observed in several species of the genus represents Danian (P2), uncinata zone: Kressenberg, 1 specimen: loc. A – a character o nly visible in specimens abo ve a certain size, 2.0 m (Fig. 151 – BSPG 1984 X 1512); usually 4-5 mm length, the main part of the argumentation Thanetian (P4), upper pseudomenardii zone: Kroisbach, 11 specimens: loc. Kch 1 (BSPG 1984 X 1513); in SCHWARZHANS (and subsequent citatio ns) appears to Thanetian (P5), velascoensis zone: Kroisbach, 1 specimen: loc. Kch be related to misidentificatio n o f certain fishes from which 11 (BSPG 1984 X 1514). the o toliths were taken. Recent extractio n o f o toliths fro m the Western Australian Museum (WAM) has revealed that Description: Large, high bodied, thick otoliths reaching up the specimens figured in SCHWARZHANS 1980 as juveniles to about 9 mm length. O L : O H = 1.1-1.25. OH : OT = 2.6- o f Centroberyx affinis (fig. 330) likely represent Centroberyx 3.3. Dorsal rim high, undulating, usually with broad predor- australis SHIMIZU & HUTCHINS 1987, which was o bvio usly sal and marked postdorsal angle, the latter above caudal not recognized then. The size of these otoliths at about 5 mm tip. Ventral rim deep, with prominent pre- and postventral wo uld usually be sufficiently mature that further o ntogenetic angles and nearly straight sectio n in between. Anterio r tip changes would not be expected. This is confirmed by a blunt, po sterio r tip o btuse, do rsally pro no unced.

Werner SCHWARZHANS 37

ba 148

c 147

149 150

a 152 b 151

Fig. 147-151. Centroberyx integer (KOKEN 1885). 147-150, Kressenberg, Danian (P1c); 147, BSPG 1984 X 1502; 148, BSPG 1984 X 1507; 149, BSPG 1984 X 1503; 150, BSPG 1984 X 1504. 151, Kressenberg, Danian (P2), BSPG 1984 X 1512. – 10 × . Fig. 152. Centroberyx fragilis SCHWARZHANS 2003. Kro isbach, Thanetian (P5), BSPG 1943 II 732; ex 518. – 10 × .

Inner face convex with supramedian sulcus. Sulcus Diagnosis. A fo ssil o tolith-based genus o f the family anterio rly o pened, po sterio rly reaching clo se to po sterio r Berycidae with the following combination of characters: tip of otolith. O stium about twice as wide as cauda, with Co mpressed o utline with high do rsal field. Ventral margin shallo w co lliculum; CaL : OsL = 1.2-1.35; cauda slightly with ro unded pre- and po stventral angles sitting far apart. deepened with straight, upward directed termination. Cau- Inner face almo st flat. Sulcus median, its o stium mo derately dal co lliculum with distinct, crest-like ventral margin. Do rsal widened and much shorter than cauda and deepened. depressio n wide, mo derately distinct; no ventral furro w. Cauda straight, slightly upward directed, terminating close O uter face co nvex with distinct po stcentral umbo , usually to distinct angle o f po sterio r rim. with so me radial furro ws. Discussion. O toliths o f Kressenbergichthys differ fro m tho se Discussion. Centroberyx integer is amongst the most com- o f Centroberyx in the compressed outline, the high dorsal mo n species in Kressenberg. For co mpariso n with C. fragilis field and the deepened o stium. Similar o toliths are also and distributio n patterns see abo ve. Centroberyx integer also fo und in the related family Trachichthyidae, which are often clo sely resembles C. te ume ri (VO IGT 1926) from the Maas- characterized by a rather pronounced rostrum (except fo r trichtian o f Bavaria, which appears to be differing mainly in Trachichthys, see KOTLYAR 1996, SCHWARZHANS 2010a a more compressed appearance in adults (O L : O H = 1.05- and below). 1.15 vs 1.1-1.25) and a smooth dorsal rim. Ano ther rather similar fo rm is fo und in the enigmatic otolith-based genus Traunichthys SCHWARZHANS 2010 fro m the Maastrichtian o f Bavaria, which was interpreted Genus Kressenbergichthys n. gen. to be related to the Melamphaidae. These otoliths have a more triangular outline with a high, median pronounced Type species: Kre sse nbe rgichthys kuhni n. sp. do rsal rim and shallo w ventral rim. Name: Referring to the Bavarian lo catio n Kressenberg Species: Kressenbergichthys is a monotypic genus with (Danian), which has yielded the majo rity o f the o to liths K. kuhni known from the Paleocene of Bavaria and Aus- described herein. tria.

Palaeo Ichthyologica 12 38

154

153

157

ba 155 156

Fig. 153-157. Kressenbergichthys kuhni n. gen. n. sp. Kressenberg, Danian (P1c). 154, Holotype, BSPG 1984 X 1515. 153, 155-157, Paratypes; 153, BSPG 1984 X 1516; 155, BSPG 1984 X 1517; 156, BSPG 1984 X 1518; 157, BSPG 1984 X 1519. – 20 × .

Kressenbergichthys kuhni n. sp. Posterio r rim high, with marked o btuse angle at level of (Figs. 153-157) caudal tip. Do rsal and ventral rims irregularly undulating; po sterio r rim rather smo o th. Holotype: Fig. 154a-c, BSPG 1984 X 1515. Inner face almo st flat with median sulcus. Sulcus anterio rly o pened, po sterio rly reaching clo se to po sterio r tip Type location: Kressenberg, Bavaria, lo catio n A – 2.5 m. of otolith. O stium short and not much wider than cauda, Type formation: O iching Formation, Danian (P1c), trinidad- somewhat deepened; cauda less deepened, slightly up- ensis zone. turned, wide with straight, tapering tip. CaL : O sL about 1.5. Caudal co lliculum with distinct, crest-like ventral margin. Paratypes: Danian (P1c), trinidade nsis zone: Kressenberg, 4 specimens: loc. A – 2.5 m (same data as holotype) (Fig. 153 – BSPG Do rsal depressio n wide, large; faint ventral furro w no t very 1984 X 1516; Fig. 155 – BSPG 1984 X 1517; Fig. 156a-b – clo se to ventral rim o f o tolith. BSPG 1984 X 1518; Fig. 157 – BSPG 1984 X 1519). O uter face flat to slightly co nvex, smo o th o r with few, sho rt radial furro ws. Further material: 24 specimens. Danian (P1c), trinidade nsis zone: Kressenberg, 22 specimens: 19 spec. loc. A – 2.5 m (same data as holotype) (BSPG 1984 X 1520); 1 spec. loc. A – 6.0 m (BSPG 1984 X 1521); 2 Family Trachichthyidae spec. loc. A – 12.7 m (BSPG 1984 X 1522); Thanetian (P4), upper pseudomenardii zone: Kroisbach, 2 speci- Genus Trachichthys SHAW 1799 mens: loc. Kch 1 (BSPG 1984 X 1523). Trachichthys anomalopsoides n. sp. Name: In honor of Winfried Kuhn and his contribution (Figs. 158-164) to the stratigraphy and paleo ntolo gy o f the Paleo cene of Bavaria and Austria. Holotype: Fig. 158a-c, BSPG 1984 X 1524. Diagnosis. See diagnosis of genus. Type location: Kro isbach, Austria, lo catio n Kch 13. Description. Moderately large, thin, high bodied otoliths Type formation: O iching Formatio n, Thanetian (P5), ve las- reaching about 3 mm length. O L : OH about 1.1 (no coensis zone. specimen with co mpletely preserved ro strum, but so me with o bvio usly small po rtio ns o nly missing). O H : OT = 4.0-4.5. Paratypes: 6 specimens. Danian (P1c), trinidade nsis zone: Kressenberg, 1 specimen: loc. A Dorsal rim high, short, with rounded predorsal and obtuse – 9.6 m (Fig. 159 – BSPG 1984 X 1525); postdorsal angles. Ventral rim deep, with rounded pre- Thanetian (P4), upper pseudomenardii zone: Kroisbach, 5 speci- and po stventral angles sitting far apart. Rostrum o bvio usly mens: loc. Kch 1 (Fig. 160 – BSPG 1984 X 1526; Fig. 161 – very thin and no t fully preserved in any o f the specimens, BSPG 1984 X 1527; Fig. 162 – BSPG 1984 X 1528; Fig. 163 but o bvio usly blunt, massive. No antiro strum o r excisura. – BSPG 1984 X 1529; Fig. 164 – BSPG 1984 X 1530).

Werner SCHWARZHANS 39

ba 159 158

161 160

162 163 164

Fig. 158-164. Trachichthys anomalopsoides n. sp. 158, Kro isbach, Thanetian (P5), Ho lo type, BSPG 1984 X 1524. 159-164, Paratypes; 159, Kressenberg, Danian (P1c), BSPG 1984 X 1525. 160-164, Kro isbach, Thanetian (P4); 160, (BSPG 1984 X 1526); 161, (BSPG 1984 X 1527); 162, (BSPG 1984 X 1528); 163, (BSPG 1984 X 1529); 164, (BSPG 1984 X 1530). – 20 × .

Further material: 14 specimens. Comparison. Trachichthys anomalopsoides differs in its Danian (P1c), trinidade nsis zone: Kressenberg, 5 specimens: 2 spec. more elongate shape (O L : OH = 1.3-1.4 vs < 1.15) from lo c. A – 2.5 m (BSPG 1984 X 1531; BSPG 1984 X 1532); the recent T. australis and the second new species of the 2 spec. loc. A – 9.6 m (BSPG 1984 X 1533); 1 spec. lo c. A – genus from Kressenberg and Kroisbach – Trachichthys 12.7 m (BSPG 1984 X 1534); impavidus n. sp. (see belo w). In this respect, T. anomalopso - Selandian (P3a), angulata zone: Kroisbach, 2 specimens: loc. 1.2 m N of Kch 4 (BSPG 1943 II 712; ex 504); and O ichinger Graben ides resembles certain ano malo pid o toliths such as fro m 1 specimen: loc. N 4 (BSPG 1943 II 519); Kryptophaneron alfredi SYLVESTER & FO WLER 1954 (fo r Thanetian (P4), upper pseudomenardii zone: Kroisbach, 6 speci- o tolith figure see SCHWARZHANS 1980). Ano malo psid mens: loc. Kch 1 (BSPG 1943 II 703; ex 430). otoliths, however, show a more reduced sulcus opening and the ostium is longer than the cauda. Name: Referring to the berycifo rm genus Anomalops of Together with Trachichthys impavidus (see belo w), the family Ano malo psidae, to which the o toliths resemble T. anomalopsoides represents the first fo ssil reco rd o f in their general appearance. the genus, which in the Recent is confined to the waters Diagnosis. Mo derately elo ngate, thin o toliths, with typical aro und Australia. Trachichthys o toliths stand o ut fro m the pentagonal outline; O L : O H = 1.3-1.4. O H : OT about 4.5. other genera placed in the family Trachichthyidae, which O stium wide, slightly sho rter than cauda; CaL : O sL = 1.1- led SCHWARZHANS (2010a) to reco gnize the family as a 1.3. Caudal tip straight, with rounded tip. monogeneric assembly and place all other genera in the no minal family Korso gasteridae. This interpretatio n now Description. Moderately large, thin, moderately elongate receives suppo rt with the o tolith finds in the Paleocene, o toliths up to slightly abo ve 3 mm length. Do rsal rim shallo w, which pro ve the separatio n o f the respective lineages since with rounded pre- and postdorsal angles. Rostrum rather the beginning o f the Tertiary at least and also suggest some sho rt, angular. Posterio r tip angular, do rsally shifted. Ventral relatio nship o f Trachichthys with the Anomalopsidae. rim deep, with preventral angle marking its deepest point. The two do rsal angles, the ro strum, po sterio r tip and the preventral angle co nstitute the pentago nal o utline of the Trachichthys impavidus n. sp. o tolith. Rims smo o th o r slightly undulating. (Figs. 165-169) Inner face lightly co nvex with distinctly supramedian sulcus. Sulcus lo ng, slightly deepened, anterio rly opened Holotype: Fig. 167, BSPG 1984 X 1535. tho ugh reducing due to its o stial margin ventrally turning upwards, po sterio rly reaching clo se to po sterio r tip o f Type location: Kressenberg, Bavaria, location A – 2.5 m. o tolith. O stium slightly sho rter than cauda and distinctly Type formation: O iching Formation, Danian (P1c), trini- wider; cauda with upturned, straight, rounded tip. Do rsal dadensis zone. depressio n narro w; ventral furro w very faint o r absent. O uter face flat and mo stly smo o th.

Palaeo Ichthyologica 12 40

b 166 a

167

165

168 169 Fig. 165-169. Trachichthys impavidus n. sp. Kressenberg, Danian (P1c). 167, Ho lo type, BSPG 1984 X 1535. 165-166, 168-169, Paratypes; 165, BSPG 1984 X 1536; 166, BSPG 1984 X 1537; 168, BSPG 1984 X 1538; 169, BSPG 1984 X 1539. – 20 × .

Paratypes: Danian (P1c), trinidade nsis zone: Kressenberg, 4 soides. The o toliths o f the recent T. australis are even more specimens: 3 spec. loc. A – 2.5 m (same data as holo type) co mpressed (O L : O H < 1.0) and sho w a ventrally expanded - (Fig. 165 – 1984 X 1536; Fig. 166a b – 1984 X 1537; Fig. 168 po sterio r po rtio n o f the o stium. – 1984 X 1538); 1 spec. loc. A – 9.6 m (Fig. 169 – BSPG 1984 X 1539). Further material: 17 specimens. Family Diretmidae Danian (P1c), trinidade nsis zone: Kressenberg, 7 specimens: loc. A – 2.5 m (same data as ho lo type) (BSPG 1984 X 1540); Genus Diretmus JOHNSON 1863 Selandian (P3a), angulata zone: Kroisbach, 3 specimens: loc. 1.2 m N of Kch 4 (BSPG 1943 II 713; ex 508); Diretmus serrativenter n. sp. Thanetian (P4), upper pseudomenardii zone: Kroisbach, 7 speci- (Figs. 170-171) mens: loc. Kch 1 (BSPG 1984 X 1541).

Name: Fro m impavidus (Latin) = impavid, reflecting the Holotype: Fig. 170a-d, BSPG 1984 X 1542. clo se resemblance to o toliths o f the recent Trachichthys aus- Type location: Kro isbach, Austria, lo catio n Kch 1. tralis despite the large time gap between the two species. Type formation: O iching Formation, Thanetian (P4), upper Diagnosis. Co mpressed, thin o toliths, with typical penta- pseudomenardii zone. go nal o utline; O L : OH = 1.05-1.1. O H : OT about 4.5. O stium wide, slightly sho rter than cauda; CaL : O sL = 1.1- Paratypes: Thanetian (P4), upper pseudomenardii zone: Kroisbach, - 1.3. Caudal tip straight, with rounded tip. 2 specimens: loc. Kch 1 (same data as holotype) (Fig. 171a b – BSPG 1984 X 1543; not fig., fragmentary BSPG 1984 X 1544). Description. Moderately large, thin, high bodied otoliths Name: Fro m serratus (Latin) = serrated and venter (Latin) up to nearly 3 mm length. Do rsal rim shallo w, with rounded = venter, belly, lo be, referring to the serrated ventral rim. pre- and po stdo rsal angles. Rostrum sho rt, blunt. Po sterio r tip angular, do rsally shifted. Ventral rim deep, with preventral Diagnosis. Extremely high bo died o toliths, with diago nally angle marking its deepest point. The two dorsal angles, the inclined shape; O L : OH = 0.7-0.9. Anterio r and po sterio r ro strum, po sterio r tip and the preventral angle co nstitute the rims parallel inclined towards rear/ do rsal. Ventral rim very pentagonal outline of the otolith. Rims smooth, but do rsal deep, with distinct denticles all alo ng. O stium very wide, commonly undulating. very sho rt; O sH : CaH = 1.7-1.9. Cauda short, straight, Inner face lightly co nvex with distinctly supramedian with rounded tip. sulcus. Sulcus lo ng, slightly deepened, anterio rly opened, Description. Small, extremely compressed high bodied po sterio rly reaching clo se to po sterio r tip o f o tolith. O stium otoliths up to a height of about 2 mm. Anterior and pos- slightly sho rter than cauda and distinctly wider; cauda with terio r rims lo ng, rather straight, parallel inclined by about upturned, straight, ro unded tip. Do rsal depressio n narrow; 30° towards rear/ do rsal. Do rsal rim sho rt, po stdo rsally no ventral furro w. pro no unced, slightly and irregularly o rnamented. Ventral O uter face flat and mo stly smo o th. rim very deep, preventrally pronounced, with many distinct Comparison. The compressed appearance distinguishes denticles fro m anterio r ventral tip to belo w caudal tip (absent T. impavidus best from the contemporaneous T. anomalop- in paratype, probably due to erosion).

Werner SCHWARZHANS 41

bc 171 a 170 a b d

Fig. 170-171. Diretmus serrativenter n. sp. Kro isbach, Thanetian (P4). 170, Ho lo type, BSPG 1984 X 1542. 171, Paratype, BSPG 1984 X 1543. – 30 × .

Inner face markedly co nvex in ho rizo ntal directio n, with rims. A third, no t figured paratype lacks mo st o f the ventral supramedian sulcus. Sulcus sho rt lo ng, slightly deepened, po rtio n o f the o tolith. anterio rly indistinctly o pened due to fading o ut o f o stium margins, po sterio rly reaching clo se to po sterio r rim of otolith. O stium slightly sho rter than cauda and nearly twice as Suborder Stephanoberycoidei wide; cauda with straight, rounded tip. Dorsal depressio n Family Melamphaidae wide, indistinct; no ventral furro w. Outer face flat and smooth. Genus indet.

Comparison. These small o toliths resemble recent Dire tmus Melamphaes? protoforma n. sp. o toliths in many respects (see SCHWARZHANS 2010a) (Figs. 172-173) except fo r the inclined nature o f its vertical axis and its rather small size. Since the ho lo type particularly exhibits Holotype: Fig. 172a-c, BSPG 1984 X 1545. such advanced morphological features it is presumed that Kro isbach, Austria, lo catio n Kch 1. D. se rrative nte r represents a small species. The slightly Type location: smaller figured paratype differs in the less ventrally ex- Type formation: O iching Formation, Thanetian (P4), upper panded ostium and the lack of the serration of the ventral pseudomenardii zone.

ab 172

173

Fig. 172-173. Melamphaes? protoforma n. sp. Kro isbach, Thanetian (P4). 17, Ho lo type, BSPG 1984 X 1545. 173, Paratype, BSPG 1984 X 1546. – 30 × .

Palaeo Ichthyologica 12 42

Paratype: Thanetian (P4), upper pseudomenardii zone: thus pro bably represent a basal, fo ssil melamphaid genus. Kroisbach, 1 specimen: loc. Kch 1 (same data as holo type) However, some aspects of the generalized appearance of the (Fig. 173 – BSPG 1984 X 1546). o utline may be attributable to the small size o f the o toliths, which may no t represent fully adult specimens. Name: Fro m pro tos (G reek) = first in rank and fo rma (Latin) = fo rm, referring to the “pro totype” appearance of these o toliths within Melamphaidae. Suborder Holocentroidei Diagnosis. Elo ngate, o val o toliths; O L : O H = 1.7. O stium anterio rly clo sed, mo derately wide, co nsiderably lo nger Family Holocentridae than cauda; O sL : CaL = 1.4-1.6; OsH : CaH = 1.2-1.35. Genus Holocentronotus SCHWARZHANS 2010 Caudal tip straight, with ro unded tip.

Description. Small, thin, elongate otoliths up to about Holocentronotus blandus n. sp. - 2.5 mm length. O H : OT about 3. Dorsal rim moderately (Figs. 174 176) shallo w, gently curving, witho ut pro minent angles. Anterio r - and po sterio r rims ro unded; anterio r rim witho ut distinct Holotype: Fig. 174a c, BSPG 1984 X 1547. ro strum. Ventral rim shallo w, gently curving, witho ut pro mi- Type location: Kroisbach, Austria, location 1.2 m N of nent angles. Rims smooth or undulating. Kch 4. Inner face almo st flat with lo ng, central sulcus. Sulcus slightly deepened, anterio rly reaching clo se to anterio r rim Type formation: O iching Formation, Selandian (P3a), o f o tolith, but no t o pening, po sterio rly reaching clo se to angulata zone. po sterio r rim o f o tolith. O stium co nsiderably lo nger than Paratypes: 4 specimens. cauda and somewhat wider, with oval outline and distinct Danian (P1c), trinidade nsis zone: Kressenberg, 2 specimens: co lliculum typical fo r many extant melamphaid o toliths; loc. A – 2.5 m (Fig. 176 – BSPG 1984 X 1548); loc. A – 9.6 m cauda straight, with ro unded tip. Do rsal depressio n indis- (Fig. 175 – BSPG 1984 X 1549); tinct; ventral furro w faint o r absent. Selandian (P3a), angulata zone: O ichinger Graben 1 specimen: loc. N 4 (BSPG 1943 II 736; ex 519); O uter face flat and smo o th o r with few sho rt radial Thanetian (P5), velascoensis zone: Kroisbach, 1 specimen: loc. Kch furro ws and margins. 12 (BSPG 1984 X 1550).

Comparison. Melamphaes? protoforma represents the Name: Fro m blandus (Latin) = nice, go o d lo o king, refer- earliest certain fo ssil melamphaid reco rd, similar in o utline ring to appearance of the otolith. and sulcus o rganizatio n to o toliths o f the living genera Melamphaes, but also Scopeloberyx and Sio . It differs fro m Diagnosis. Elo ngate, fusifo rm o toliths; O L : OH = 1.8. all o f them in the rather lo ng o stium, which is lo nger than Anterio r and po sterio r tips symmetrically po inted. O stium the cauda and anteriorly not as much reduced as this is the widened, sho rter than narro w cauda; CaL : OsL = 1.3. case with o toliths o f recent fishes o f the family. These o toliths Caudal tip o nly mildly bent. No ventral furro w.

b 174 a c

175 176

Fig. 174-176. Holocentronotus blandus n. sp. 174, Kro isbach, Selandian (P3a), Ho lo type, BSPG 1984 X 1547. 175-176, Paratypes, Kressenberg, Danian (P1c); 175, BSPG 1984 X 1549, 176 1984 X 1548. – 20 × .

Werner SCHWARZHANS 43

ba 177 c

178

180 182

179 181

Fig. 177-182. Polymixia polita n. sp. Kressenberg, Danian (P1c). 177, Holotype, BSPG 1984 X 1551. 178-182, Paratypes; 178, BSPG 1984 X 1552; 179, BSPG 1984 X 1553; 180, BSPG 1984 X 1554; 181, BSPG 1984 X 1555; 182, BSPG 1984 X 1556. – 10 × .

Description. Elo ngate, mo derately thin o toliths reaching plus (SCHWARZHANS 1980) also from New Zealand (as about 4 mm length. O H : OT about 2.5. Dorsal rim shal- Adioryx). A true representative o f Ho lo ce ntrus is H. shep- low, gently curved, with broad, obtuse middorsal angle. peyensis (FRO ST 1934) from the Eocene of England and Ventral rim shallo w, gently and regularly curved, deepest Germany. at its middle. Rostrum massive, lo ng, with po inted tip. No The slender, symmetrical shape o f the o tolith and its pro - o r very feeble antiro strum o r excisura. Posterio r tip pointed po rtio ns o f the o utline and the sulcus distinguish H. blandus at level o f caudal tip, almo st symmetrical to ro strum. All from the other species mentioned. rims smo o th. Inner face markedly co nvex with clearly supramedian sulcus. Sulcus anterio rly o pened, po sterio rly reaching mod- Suborder Polymixioidei erately clo se to po sterio r tip o f o tolith. O stium much wider than cauda, particularly ventrally widened, slightly deep- Family Polymixiidae ened; cauda deeper, very narrow and slightly longer than Genus Polymixia LOWE 1836 o stium, its tip mo derately bent and terminating mo derately clo se to po sterio r tip o f o tolith. Do rsal depressio n very feeble; Polymixia polita n. sp. ventral field smo o th witho ut ventral furro w. (Figs. 177-182) O uter face flat to slightly co ncave, smo o th. - Discussion. Ho lo ce ntro no tus appears to be a fairly common Holotype: Fig. 177a c, BSPG 1984 X 1551. specious holocentrid during Late Cretaceous, Paleocene Type location: Kressenberg, Bavaria, lo catio n A – 2.5 m. and o ccasio nally Eocene. It differs fro m the recent genera except Ho lo ce ntrus in the less stro ngly bent caudal tip and Type formation: O iching Formation, Danian (P1c), trini- the po inted o r ro unded po sterio r rim (vs blunt o r concave), dadensis zone. bo th characters perceived to represent a plesio mo rphic Paratypes: 5 specimens. status. Ho lo ce ntro no tus shares these two characters with Danian (P1c), trinidade nsis zone: Kressenberg: 4 spec. loc. A Ho lo ce ntrus which ho wever differs in the shallo w o stium, – 2.5 m (same data as holotype) (Fig. 178 – 1984 X 1552; which also is as long as or longer than the cauda and Fig. 179 – 1984 X 1553; Fig. 180 – 1984 X 1554; Fig. 181 usually curved. – 1984 X 1555); 1 spec. loc. A – 12.7 m (Fig. 182 – BSPG 1984 X 1556). The following species may be accounted for in Ho lo - ce ntro no tus: Maastrichtian o f Bavaria: H. percomorphus Further material: 13 specimens. SCHWARZHANS 2010a (type species); Paleo cene: H. ryab- Danian (P1c), trinidade nsis zone: Kressenberg, 11 specimens: chuni SCHWARZHANS & BRATISHKO 2011 fro m Ukraine 9 spec. loc. A – 2.5 m (same data as holotype) (BSPG 1984 X 1557); 1 spec. loc. A – 9.6 m (BSPG 1984 X 1558; 1 (as genus Holocentridarum) and H. blandus fro m Bavaria spec. loc. A – 12.7 m (BSPG 1984 X 1559); and Austria; Eocene: H. palasulcatus SCHWARZHANS Thanetian (P4), upper pseudomenardii zone: Kroisbach, 2 speci- 1980, H. ve ntrico sus SCHWARZHANS 1980 bo th fro m mens: loc. Kch 1 (BSPG 1984 X 1560). New Zealand (both as genus Holocentridarum) and H. am-

Palaeo Ichthyologica 12 44

Name: Fro m po litus (Latin) = po lite, friendly, referring to with relatively smo o th transitio n to cauda, slightly deepened; the easy reco gnitio n o f these o toliths as representatives o f cauda narrow and markedly longer than ostium, its tip the genus Polymixia. mo derately bent, with ro unded tip. Do rsal depressio n feeble, narro w; ventral field smo o th witho ut ventral furro w. Diagnosis. Co mpressed, high bo died o toliths; O L : OH O uter face flat to slightly co ncave, smo o th o r with few - = 1.1 1.2. Rostrum blunt; po sterio r tip do rsally po inted. sho rt radial furro ws do rsally. Ventral rim deep, deepest anterior of the middle. Sulcus distinctly supramedian; caudal tip mo derately bent. No Discussion. Polymixia po lita exhibits the typical o tolith mo r- ventral furro w. phology of recent species of the genus Polymixia, particularly of the more compressed morphologies such as are found Description. High bo died, mo derately thin o toliths reaching in P. lo we i G ÜNTHER 1859 (see SCHWARZHANS 2010a), about 5 mm length. O H : OT about 4. Dorsal rim shallo w, P. be rndti G ILBERT 1905 (see SMALE et al. 1995) o r P. lo ngi- with bro ad predo rsal and indistinct po stdo rsal angles, spina DENG, XIO NG & ZHAN 1983 (see O HE 1985), smo o th o r slightly crenulated. Ventral rim deep, regularly while tho se o f P. no bilis LOWE 1836 (see SCHWARZHANS curved, deepest anterio r o f its middle, belo w o stium, smooth. 2010a) or P. japonica G ÜNTHER 1877 (see RIVATON & Rostrum high, sho rt, with blunt tip. Antiro strum and excisura BO URET 1999) are considerably more elongate. Polymixia indistinct. Posterio r tip angular, distinct, at level above caudal po lita o toliths are mo re co mpressed still than tho se o f any tip, more pointed than rostrum. of the known recent species. They show a less steeply bent Inner face markedly co nvex with clearly suprame- caudal tip than P. lo we i and P. be rndti, but a more strongly dian sulcus. Sulcus anterio rly o pened, po sterio rly reaching bent tip than P. longispina. clo se to po stventral margin o f o tolith belo w po sterio r tip. Polymixia po lita represents the first certain fo ssil reco rd CaL : OsL = 1.4-1.6. O stium markedly wider than cauda, o f a po lymixiid o tolith-based species.

3.11 Order Zeiformes

Family indet. near Parazenidae moderately deep, with rounded medioventral and preventral angles, deepest far anterio rly belo w ro strum, mo derately Genus Isozen SCHWARZHANS 2010 undulating. Anterio r rim with high, blunt ro strum, deep, sharp excisura and pro minent antiro strum. Posterio r rim Isozen janni (SCHWARZHANS 2003) with rounded tip above termination of cauda. Inner face flat, with slightly supramedian, slightly deep- 2003 genus Zeifo rmo rum janni SCHWARZHANS – fig. 31D-F ened sulcus. Sulcus anterio rly o pen, po sterio rly closed. Material: 1 specimen. O stium and cauda of about equal length. O stial and Thanetian (P4), upper pseudomenardii zone: Kroisbach, location caudal co lliculi very small, deepened; O CL : CCL = 1.4. Kch 1 (BSPG 1984 X 1561). Co llum wide, ventrally co nvex, with lo ng, no t very distinct Remarks. The single repo rted specimen fro m Kro isbach is pseudocolliculum extending across entire collum. Do rsal inco mpletely preserved with the do rsal part o f the o tolith field flat, witho ut marked depressio n; ventral field elevated missing. The thin appearance, the regularly curved ventral belo w co llum, ventrally marked by a bro ad, sho rt ventral rim and the caudal colliculum reaching close to the po sterio r furro w at so me distance fro m ventral rim o f o tolith. rim o f the o tolith all characterize this specimen as I. janni O uter face markedly convex, smooth except for few (SCHWARZHANS 2003). Isozen janni so far has only been radial furro ws ventrally. reported from the Selandian of Denmark. Discussion. Isozen mareikeae resembles mo st I. beateae SCHWARZHANS 2010a fro m the Maastrichtian o f Ba- varia, differing mainly in the deepened co lliculi (vs el- Isozen mareikeae n. sp. evated) and the less deeply curved ventral rim. Isozen janni (Fig. 183)

Holotype: Fig. 183a-c, BSPG 1984 X 1562. Type location: Kro isbach, Austria, lo catio n Kch 1. Type formation: O iching Formation, Thanetian (P4), upper pseudomenardii zone.

Name: After my daughter Anna Mareike. ba Diagnosis. Mo derately high bo died o toliths; O L : OH 183 = 1.15. Sulcus anteriorly open with deep excisura, po sterio rly clo sed. Co lliculi small, particularly caudal colliculum, c deepened, widely separated. Long pseudocolliculum in between co lliculi. O uter face distinctly co nvex.

Description. Small, mo derately thick o tolith o f slightly mo re than 1 mm length. O H : OT about 2.8. Dorsal rim high, with Fig. 183. Isozen mareikeae n. sp. Kro isbach, Thanetian (P4), o btuse medio do rsal angle, slightly undulating. Ventral rim Holotype, BSPG 1984 X 1562. – 30 × .

Werner SCHWARZHANS 45

(SCHWARZHANS 2003) fro m the Paleo cene o f Denmark assessments in terms o f certain co nclusio ns o f po larity and likewise sho ws deepened co lliculi, but they are larger and the zeifo rm clado gram o f interrelatio nships, it is fair to state the caudal colliculum reaches close to the posterio r rim o f that bo th studies regard the Late Cretaceo us / Early Tertiary the o tolith. Also its sulcus lacks a pseudo co lliculum in the otoliths mentioned above as plesiomorphic morphologies co llum and the o tolith is thinner. O ther po ssibly related o f rather basal relatio nship in the evo lutio n o f zeifo rms. In species have been described from the Late Cretaceous of the case of zeiforms, however, paleo-ichthyological data Spain – Iso ze n tyle ri NO LF 2003 (as genus Zeiformorum) – fro m the Cretaceo us and Early Tertiary are far superio r to and the Early Eocene of England – Iso ze n sulcife r (STINTON otolith based data in wealth and complexity and therefo re 1966) (as Amanses sulcifer). it appears inappro priate to expand into further discussio n The distinct specializatio n effects seen in o toliths of the o n o toliths. Acco rding to TYLER & SANTINI (2005) at least Zeifo rmes (and Tetrao do ntifo rmes) were subject o f a system- six independent lineages of zeiforms have been present atic review and phylo genetic evaluatio n by NO LF & TYLER during the Late Cretaceous and survived the Cretaceous/ (2006) and were also discussed in SCHWARZHANS (2010a). Tertiary (K/ T) extinctio n to radiate in Ceno zo ic seas. While there is some principle disagreement between the two

3.12 Order Perciformes

Suborder Percoidei Genus Plesiopoma SCHWARZHANS 2010

Family Acropomatidae Plesiopoma elegantissima n. sp. (Figs. 184-189) The family Acropomatidae is used here in a broad sense (sensu lato) tentatively co ntaining the plesio mo rphic fo s- Holotype: Fig. 184a-b, BSPG 1984 X 1563. sil o tolith-based perco id genus Plesiopoma considered Type location: Kressenberg, Bavaria, location A – 12.7 m. to represent the o nly certain pre-Tertiary o tolith record of percifo rms (see SCHWARZHANS 2010a). In their extensive Type formation: O iching Formation, Danian (P1c), trinidad- discussio n o f the descriptio n o f the enigmatic Late Creta- ensis zone. ceo us fish Nardoichthys francisci SO RBINI & BANNIKOV Paratypes: 5 specimens. (1991) po int to a po ssible relatio nship with primitive perco id Danian (P1c), trinidade nsis zone: Kressenberg: 2 spec. loc. A families mentio ning Acro po matidae as a po ssible relative. – 2.5 m (Fig. 185 – BSPG 1984 X 1564; Fig. 186 – BSPG PATTERSO N (1993) ho wever regarded Nardoichthys as of 1984 X 1565); 3 spec. loc. A – 12.7 m (same data as ho lo type) uncertain relatio nship. In their review o f the Late Cretaceous (Fig. 187 – BSPG 1984 X 1566; Fig. 188 – BSPG 1984 X 1567; to Paleo cene perco mo rphs ARRATIA et al. (2004) regard Fig. 189 – BSPG 1984 X 1568). Nardoichthys as the o ldest kno wn percifo rm skeleton find Further material: 9 specimens. from about 71 Ma, further mentioning Salde nio ichthys Danian (P1c), trinidade nsis zone: Kressenberg: 5 spec. loc. A – LOPEZ-ARBARELLO, ARRATIA & TUNIK 2003, Eose rranus 2.5 m (BSPG 1984 X 1569); 1 spec. loc. A – 9.6 m (BSPG WOODWARD 1908 and Indiaichthys ARRATIA et al. 2004 1984 X 1570); 3 spec. loc. A – 12.7 m (same data as ho lo type) as percifo rms (the latter two as perco ids) o f unkno wn (BSPG 1984 X 1571). relationship from the Late Cretaceous-Paleocene interval Name: Fro m superlative o f elegantus (Latin) = elegant, between 67 and 62 Ma. delicate, referring to the slender o utline o f these o toliths.

Diagnosis. Elo ngate, thin o toliths; O L : OH = 1.7-1.8; O H : OT abo ut 4. Small o tolith-species seemingly no t

186 a 185 184 b

187 188 189 Fig. 184-189. Plesiopoma elegantissima n. sp. Kressenberg, Danian (P1c). 184, Holotype, BSPG 1984 X 1563. 185-189, Paratypes; 185, BSPG 1984 X 1564; 186, BSPG 1984 X 1565; 187, BSPG 1984 X 1566; 188, BSPG 1984 X 1567; 189, BSPG 1984 X 1568. – 20 × .

Palaeo Ichthyologica 12 46

191

190

192

a 193 194 b

195

196 Fig. 190-196. Plesiopoma traubi n. sp. 193, Kressenberg, Danian (P1c), Ho lo type, BSPG 1984 X 1572. 190-192, 194-196, Paratypes; 195-196, Kressenberg, Danian (P1b); 195, BSPG 1984 X 1573; 196, BSPG 1984 X 1574. 191-192, Kressenberg, Danian (P1c); 191, BSPG 1984 X 1575; 192, BSPG 1984 X 1576; 194, BSPG 1984 X 1577. 190, Kroisbach, Thanetian (P4), BSPG 1984 X 1578. – 20 ×.

exceeding 2.5 mm length. Rostrum moderately long, 25- always remains well distinguishable fro m small specimens of 30 % of O L, pointed. No or very feeble excisura. Do rsal the latter species. Hence it is assumed that P. elegantissima rim shallo w. represents a small “dwarf” species.

Description. Small, elo ngate, thin o toliths no t exceeding 2.5 mm length. Do rsal rim shallo w, gently curved, witho ut Plesiopoma traubi n. sp. pro minent angles. Ventral rim shallo w, regularly curved, (Figs. 190-196) deepest slightly anterio r o f its middle. Rostrum mo derately lo ng, po inted. Excisura and antiro strum minute o r absent. Holotype: Fig. 193a-b, BSPG 1984 X 1572. Posterio r rim ro unded. All rims smo o th. Inner face flat, with lo ng, rather wide, slightly deep- Type location: Kressenberg, Bavaria, lo catio n A – 2.5 m. ened, median sulcus. Sulcus anteriorly opened, posteri- Type formation: O iching Formation, Danian (P1c), trinidad- o rly terminating clo se to po sterio r rim o f o tolith. O stium ensis zone. slightly longer and wider than cauda; O sL : CaL = 1.1-1.4; OsH : CaH = 1.2-1.3. Cauda straight, not bent. Colliculi Paratypes: 6 specimens. well marked. Do rsal depressio n small, indistinct; ventral Danian (P1b), pse udo bullo ide s zone: Kressenberg, 2 specimens: furro w mo stly feeble, clo se to ventral rim o f o tolith. loc. B3 (Fig. 195 – BSPG 1984 X 1573; Fig. 196 – BSPG 1984 X 1574); Outer face flat, smooth. Danian (P1c), trinidade nsis zone: Kressenberg, 3 specimens: Discussion. Plesiopoma elegantissima is a co nstantly small 1 spec. lo c. A – 2.5 m (same data as ho lo type) (Fig. 191 – and thin otolith which does not attain the size of the paral- BSPG 1984 X 1575); 2 spec. loc. A – 12.7 m (Fig. 192 – BSPG 1984 X 1576; Fig. 194 – BSPG 1984 X 1577); lel o ccurring P. traubi n. sp., but due to its elongate shape

Werner SCHWARZHANS 47

Thanetian (P4), upper pseudomenardii zone: Kroisbach, 1 specimen: mens: loc. Kch 1 (Fig. 202 – BSPG 1984 X 1588; Fig. 203a-b loc. Kch 1 (Fig. 190 – BSPG 1984 X 1578). – BSPG 1984 X 1589).

Further material: 31 specimens; Further material: 343 specimens. Danian (P1b), pse udo bullo ide s zone: Kressenberg, 12 specimens: Danian (P1b), pse udo bullo ide s zone: Kressenberg, 4 specimens: 2 spec. loc. B1 (BSPG 1984 X 1579); 10 spec. loc. B3 (BSPG 2 spec. loc. A – 1.0 m (BSPG 1984 X 1590); 2 spec. loc. B3 1984 X 1580); (BSPG 1984 X 1591); Danian (P1c), trinidade nsis zone: Kressenberg, 18 specimens: loc. A Danian (P1c), trinidade nsis zone: Kressenberg, 303 specimens: – 2.5 m (same data as ho lo type) (BSPG 1984 X 1581); 208 spec. loc. A – 2.5 m (same data as holotype) (BSPG Selandian (P3a), angulata zone: Kroisbach, 1 specimen: loc. 1.2 m 1984 X 1592); 8 spec. loc. A – 2.5 m (same data as holo- N of Kch 4 (BSPG 1943 II 714; ex 508). type) (BSPG 1984 X 1593); 13 spec. loc. A – 6.0 m (BSPG 1984 X 1594); 1 spec. loc. A – 8.3 m (BSPG 1984 X 1595); Name: In memory of Franz Traub (München) and his most 23 spec. loc. A – 9.6 m (BSPG 1984 X 1596); 2 spec. loc. A – valuable contribution to the understanding of the Bavarian 11.6 m (BSPG 1984 X 1597); 48 spec. loc. A – 12.7 m (BSPG Tertiary strata and their fauna. 1984 X 1598); and O ichinger Graben, 2 specimens: lo c. N 2 - (BSPG 1943 II 739; ex 520); Diagnosis. O val o toliths; O L : OH = 1.3 1.45; OH : OT Danian (P2), uncinata zone: Kressenberg, 11 specimens: loc. A – - about 4. Rostrum not very long, 18 25 % of O L, moderately 2.0 m (BSPG 1984 X 1599); po inted. Excisura sharp. Do rsal rim high, regularly rounded. Selandian (P3a), angulata zone: Kroisbach, 8 specimens: 3 spec. loc. 1.2 m N of Kch 4 (BSPG 1943 II 715; ex 508); 5 spec. Description. O val, mo derately thin o toliths up to abo ut loc. “close to” Kch 4 (BSPG 1943 II 721; ex 512); 8 mm length (according to a single incomplete large speci- Thanetian (P4), upper pseudomenardii zone: Kroisbach, 15 specimen). Do rsal rim high, regularly curved, witho ut prominent mens: 10 spec. loc. Kch 1 (BSPG 1943 II 704; ex 430); 4 spec. angles. Ventral rim mo derately deep, very regularly curved, loc. Kch 1 (BSPG 1943 II 705; ex 444); 1 spec. loc. 8 m S of deepest slightly anterio r o f its middle. Rostrum mo derately Kch 1 (BSPG 1943 II 708; ex 445). long 20-30 % o f O L, mo derately po inted. Excisura sharp, Name: Fro m simplex (Latin) = simple, referring to the no t very deep; antiro strum sho rt, po inted. Posterio r rim plesio mo rphic mo rpho lo gy o f the o toliths. bro adly ro unded. All rims smo o th. Inner face almo st flat, with lo ng, rather wide, slightly Diagnosis. High bo died, ro unded, thick o toliths; O L : OH deepened, median sulcus. Sulcus anteriorly opened, po steri- = 1.2-1.3; OH : OT = 2.5-3.0. Rostrum short, about 15 % o rly terminating very clo se to po sterio r rim o f o to lith. O stium of O L, blunt. Dorsal rim high, with obtuse pre- and po st- slightly longer and wider than cauda; O sL : CaL = 1.1-1.3; do rsal angles. O stium widened, slightly upward turned; OsH : CaH = 1.1-1.4. Cauda straight, not bent. Colliculi well cauda with slightly do wnturned tip terminating clo se to marked. Do rsal depressio n wide, indistinct; ventral furro w po sterio r rim o f o tolith. Single ventral furro w clo se to ventral mo stly well marked, mo derately clo se to ventral rim o f o tolith. rim o f o tolith. O uter face almo st flat, smo o th. Description. High bodied otoliths up to about 4 mm length Discussion. Ple sio po ma traubi is very similar to P. o tio sa with well rounded outline. Dorsal rim high, with obtuse pre- SCHWARZHANS 2010 fro m the Maastrichtian o f Bavaria, and po stdo rsal angles. Ventral rim deep, regularly curved differing mainly in the somewhat more compressed out- and deepest at its middle in large specimens, occasio nally line (OL : OH = 1.3-1.45 vs 1.45-1.65) and the clearly with bro ad po stventral angle in specimens smaller 2 mm develo ped excisura and antiro strum (vs absent o r very in length. Rostrum sho rt, blunt, rarely so mewhat po inted. feeble). Both species are clearly related and P. traubi likely Excisura weak, no t very deep; antiro strum sho rt, mo stly in- represents the descendant o f the earlier P. o tio sa, thereby distinct. Posterio r rim bro adly ro unded. Do rsal and po sterio r establishing o ne mo re lineage o f teleo sts having survived the rims o ften irregularly undulating, but never very intensely. Cretaceo us/ Tertiary bo undary in Bavaria. Ple sio po ma traubi Inner face stro ngly bent alo ng ho rizo ntal axis, much less occurs in parallel with the small and more elongate species in vertical axis. Sulcus lo ng, slightly deepened and slightly P. elegantissima (see abo ve fo r further discussio n). supramedian. Sulcus anterio rly o pened, po sterio rly terminating very clo se to po sterio r rim o f o tolith. O stium sho rter but considerably wider than cauda; CaL : OsL = 1.35-1.6; Family Lactariidae OsH : CaH = 1.6-2.0. O stium slightly turned upwards and deepened; cauda narrower, straight until the slightly down- Genus indet. turned tip. Co lliculi po o rly marked. Do rsal depression wide, Lactarius? simplex n. sp. indistinct; ventral furro w mo derately well marked, clo se to (Figs. 197-204) ventral rim o f o tolith. O uter face flat, with postcentral umbo, with some faint Holotype: Fig. 199a-c, BSPG 1984 X 1582. radial furro ws near rims o f o tolith. Type location: Kressenberg, Bavaria, lo catio n A – 2.5 m. Discussion. Lactarius? simple x is the most common species at Kressenberg. It is an o tolith with a very generalized per- Type formation: O iching Formation, Danian (P1c), trinidad- co id mo rpho lo gy that resembles mo dern and fo ssil lactariid ensis zone. otoliths known since Eocene times from Europe and New Paratypes: 7 specimens. Zealand. It differs fro m mo dern representatives o f the genus Danian (P1b), pse udo bullo ide s zone: Kressenberg, 1 specimen: Lactarius in the absence of a second ventral furrow located lo c. B1 (Fig. 197 – BSPG 1984 X 1583); midway on the ventral field of the inner face. Therefore, Danian (P1c), trinidade nsis zone: Kressenberg, 4 specimens: 3 its allo catio n to the genus Lactarius remains tentative. The spec. loc. A – 2.5 m (same data as holotype) (Fig. 198 – BSPG earliest pro ven lactariid o tolith reco rd so far stems fro m 1984 X 1584; Fig. 200 – BSPG 1984 X 1585; Fig. 201 – BSPG the Ypresian of the London Basin – Lactarius curvido rsalis 1984 X 1586); 1 spec. loc. A – 12.7 m (Fig. 204 – BSPG 1984 X 1587); STINTON 1965 – which is easily distinguished by its much Thanetian (P4), upper pseudomenardii zone: Kroisbach, 2 speci- more elongate shape.

Palaeo Ichthyologica 12 48

198 197

200 b a 199 c

201 a 203 b

202 204

Fig. 197-204. Lactarius? simplex n. sp. 199, Kressenberg, Danian (P1c), Ho lo type, BSPG 1984 X 1582. 197-198, 200-204, Paratypes; 197, Kressenberg, Danian (P1b), BSPG 1984 X 1583; 198, 200-201, 204, Kressenberg, Danian (P1c); 198, BSPG 1984 X 1584; 200, BSPG 1984 X 1585; 201, BSPG 1984 X 1586; 204, BSPG 1984 X 1587; 202-203, Kro isbach, Thanetian (P4); 202, BSPG 1984 X 1588; 203, BSPG 1984 X 1589. – 20 × .

Family Serranidae Paratypes: 3 specimens. Danian (P1c), trinidade nsis zone: Kressenberg, loc. A – 2.5 m (same Genus Polyperca STINTON 1965 data as ho lo type) (Fig. 205 – BSPG 1984 X 1601; Fig. 207 – BSPG 1984 X 1602; Fig. 208 – BSPG 1984 X 1603).

Polyperca exserta n. sp. Further material: 27 specimens. - (Figs. 205 208) Danian (P1c), trinidade nsis zone: Kressenberg, 26 specimens: 19 spec. loc. A – 2.5 m (same data as holotype) (BSPG Holotype: Fig. 206a-c, BSPG 1984 X 1600. 1984 X 1604); 2 spec. loc. A – 2.5 m (same data as holotype) (BSPG 1984 X 1605); 1 spec. loc. A – 11.6 m (BSPG Type location: Kressenberg, Bavaria, lo catio n A – 2.5 m. 1984 X 1606); 4 spec. loc. A – 12.7 m (BSPG 1984 X 1607); Type formation: O iching Formation, Danian (P1c), trinidad- Danian (P2), uncinata zone: Kressenberg, 1 specimen: loc. A – 2.0 m (BSPG 1984 X 1608). ensis zone.

Werner SCHWARZHANS 49

205 206 b c

207 208

Fig. 205-208. Polyperca? exserta n. sp. Kressenberg, Danian (P1c). 206, Ho lo type, BSPG 1984 X 1600. 205, 207-208, Paratypes; 205, BSPG 1984 X 1601; 207, BSPG 1984 X 1607); 208, BSPG 1984 X 1608). – 20 × .

Name: Fro m exsertus (Latin) = pro jecting, referring to the Selandian o f West-G reenland has similar pro po rtio ns, but pronounced rostrum. a shorter rostrum, a pronounced postdorsal angle and is also thinner. Diagnosis. Elo ngate, rather thick o toliths; O L : OH = 1.6- 1.7; O H : OT about 2.5. Rostrum long, with rounded tip, about 20-25 % o f O L. Do rsal rim shallo w, regularly curved. O stium moderately widened; cauda with slightly downturned Family Sparidae tip terminating mo derately clo se to po sterio r rim o f o tolith. Genus Platysepta STINTON 1965 No o r weak ventral furro w. Platysepta kressenbergensis n. sp. Description. Elo ngate, thick o toliths up to abo ut 4 mm (Figs. 209-215) length. Do rsal rim shallo w, regularly curved, witho ut pro minent angles. Ventral rim shallo w too , regularly curved, 2003 “genus Sparidarum” sp. – SCHWARZHANS: fig. 37F-N deepest at its middle. Rostrum lo ng, its do rsal rim usually ho rizo ntal, with ro unded tip. Excisura and antiro strum feeble Holotype: Fig. 209a-b, BSPG 1984 X 1609. o r absent. Posterio r rim with angular tip at abo ut level o f Type location: Kroisbach, Austria, location 1.2 m N of Kch 4. caudal tip. All rims smo o th. Inner face stro ngly bent alo ng ho rizo ntal axis, less in Type formation: O iching Formation, Selandian (P3a), vertical axis. Sulcus lo ng, slightly deepened and slightly su- angulata zone. pramedian. Sulcus anterio rly o pened, po sterio rly terminating Paratypes: 6 specimens. mo derately clo se to po sterio r rim o f o tolith. O stium slightly Danian (P1c), trinidade nsis zone: Kressenberg, 5 specimens: 4 spec. shorter and moderately wider than cauda; CaL : O sL = 1.1- loc. A – 2.5 m (Fig. 210 – BSPG 1984 X 1610; Fig. 212 – BSPG 1.3; OsH : CaH = 1.4-1.7. O stium slightly deepened; 1984 X 1611; Fig. 214 – BSPG 1984 X 1612; Fig. 215a-c – cauda straight until the slightly do wnturned ro unded tip. BSPG 1984 X 1613); 1 spec. loc. A – 12.7 m (Fig. 213 – BSPG Co lliculi po o rly marked. Do rsal depressio n small, indistinct; 1984 X 1614); ventral furro w feeble o r absent. Thanetian (P5), velascoensis zone: Kroisbach, 1 specimen: loc. Kch - Outer face flat, smooth. 12 (Fig. 211a b – BSPG 1984 X 1615). Further material: 30 specimens. Discussion. These plesio mo rphic o toliths are tentatively Danian (P1c), trinidade nsis zone: Kressenberg: 23 spec. loc. A placed to the fo ssil o tolith-based genus Polype rca in the – 2.5 m (BSPG 1984 X 1616); 2 spec. loc. A – 6.0 m (BSPG family Serranidae, mainly because o f their elo ngate shape, 1984 X 1617); 2 spec. loc. A – 8.3 m (BSPG 1984 X 1618); 5 the long rostrum and the moderately widened ostium. Simi- spec. loc. A – 12.7 m (BSPG 1984 X 1619). lar o toliths have been described fro m the Early Ypresian Name: After the lo catio n Kressenberg, Bavaria. (Sparnacian) of the London Basin by STINTON 1965 as Polyperca serranoides. It differs in the mo re slender shape Diagnosis. Elo ngate, mo derately thick o toliths; O L : OH of the otolith as well as the cauda not reaching as far = 1.6-1.7; O H : OT about 3.5. Rostrum short, about 15- backwards. Ano ther similar serranid has been described 18 % of O L. Dorsal rim high, broadly undulating in adults. as Serranidae indet. fro m the Selandian o f Belgium by O stium widened; cauda with markedly downturned tip NO LF 1978, which again is more slender and also thinner. terminating clo se to po sterio r rim o f o tolith. Acropoma? rosenkrantzi SCHWARZHANS 2004 fro m the

Palaeo Ichthyologica 12 50

210

209 a b 211 b 212

ba215

213 c 214

Fig. 209-215. Platysepta kressenbergensis n. sp. 209, Kro isbach, Selandian (P3a), Ho lo type, BSPG 1984 X 1609. 210-215, Para- types; 210, 212-215, Kressenberg, Danian (P1c); 210, BSPG 1984 X 1610; 212, BSPG 1984 X 1611; 213, BSPG 1984 X 1614; 214, BSPG 1984 X 1612; 215, BSPG 1984 X 1613. 211, Kroisbach, Thanetian (P5), BSPG 1984 X 1615. – 10 ×.

Description. Elo ngate, mo derately thick o toliths up to abo ut Juvenile specimens of the same species have been re- 7.5 mm length. Dorsal rim high, with poorly developed corded as Sparidae indet. from the Selandian of Denmark angles, undulating, coarsely undulating in large specimens. by SCHWARZHANS (2003). They differ slightly fro m the Ventral rim mo derately deep, regularly curved, deepest at larger specimens at Kro isbach and Kressenberg in the more its middle, smo o th. Rostrum sho rt, its do rsal rim inclined, po inted po sterio r tip o f the o tolith, which is co nsidered to with po inted tip. Excisura and antiro strum feeble o r absent. represent an ontogenetic effect. Posterio r rim irregular, bro adly ro unded o r with o btuse angle at level o f caudal tip. Inner face stro ngly bent alo ng ho rizo ntal axis, less in Genus Dentex CUVIER 1814 vertical axis. Sulcus lo ng, slightly deepened and slightly supramedian. Sulcus anterio rly o pened, po sterio rly terminat- Dentex? solidus n. sp. ing clo se to po sterio r rim o f o tolith. O stium slightly sho rter (Figs. 216-220) but considerably wider than cauda; CaL : OsL = 1.1-1.3; OsH : CaH = 1.6-2.0. O stium slightly deepened; cauda with Holotype: Fig. 216a-c, BSPG 1984 X 1620. markedly do wnturned tip, increasingly in larger specimens. Type location: Kressenberg, Bavaria, lo catio n A – 2.5 m. Co lliculi po o rly marked. Do rsal depressio n small, indistinct; ventral furro w mo stly present, clo se to ventral rim o f o tolith. Type formation: O iching Formation, Danian (P1c), trinidad- O uter face concave to almost flat, smooth or with few ensis zone. radial furro ws in juveniles. Paratypes: 4 specimens. Discussion. O toliths o f Platysepta kressenbergensis resem- Danian (P1c), trinidade nsis zone: Kressenberg, 2 specimens: ble tho se o f the sympatric Polyperca exserta differing mainly 1 spec. lo c. A – 2.5 m (same data as ho lo type) (Fig. 217 – BSPG 1984 X 1621); 1 spec. loc. A – 3.0 m (Fig. 220 – BSPG in the thinner appearance (O H : OT = 3.5 vs 2.5), the sho rter - - 1984 X 1622); ro strum (15 18 % of O L vs 20 25 %), the more strongly bent Thanetian (P4), upper pseudomenardii zone: Kroisbach, 2 speci- caudal tip and details in the outline of the dorsal rim and mens: loc. Kch 1 (Fig. 218a-b – BSPG 1984 X 1623; Fig. 219 the ro strum. Fro m Platyse pta carribaea (NO LF & DO CK- – BSPG 1984 X 1624). ERY 1993) as Ne mipte rus carribae us it differs in the mo re Further material: 15 specimens. compressed appearance (O L : O H = 1.6-1.7 vs 1.75-1.95) Danian (P1b), pse udo bullo ide s zone: Kressenberg, 1 specimen: and the shape of the dorsal and ventral rims. From the lo c. B3 (BSPG 1984 X 1625); type-species Platyse pta prima STINTON 1965 it differs in Danian (P1c), trinidade nsis zone: Kressenberg, 11 specimens: the more compressed shape, the less pronounced posterio r 10 spec. loc. A – 2.5 m (same data as holotype) (BSPG tip, the wider ostium and the more strongly bent caudal tip. 1984 X 1626); 1 spec. loc. A – 2.5 m (BSPG 1984 X 1627); Platyse pta is mo ved fro m the Serranidae, where STINTON Thanetian (P4), upper pseudomenardii zone: Kroisbach, 1 specimen: placed it, to be understoo d as a primitive Sparidae. spec. loc. Kch 1 (BSPG 1984 X 1628).

Werner SCHWARZHANS 51

b a 216 c 217

218 a b

219 220

Fig. 216-220. Dentex? solidus n. sp. 216, Kressenberg, Danian (P1c), Ho lo type, BSPG 1984 X 1620. 217-220, Paratypes; 217, 220, Kressenberg, Danian (P1c); 217, BSPG 1984 X 1621; 220, BSPG 1984 X 1622; 218-219, Kro isbach, Thanetian (P4); 218, BSPG 1984 X 1623; 219, BSPG 1984 X 1624. – 10 × .

Thanetian (P5), velascoensis zone: Kroisbach, 2 specimens: 1 spec. Family Haemulidae loc. Kch 11b (BSPG 1943 II 725; ex 515); 1 spec. lo c. Kch 12 (BSPG 1943 II 730; ex 516). Genus indet.

Name: Fro m so lidus (Latin) = massive, referring to the high Haemulon? conjugator n. sp. bo died nature o f the o toliths. (Figs. 221-223) Diagnosis. Compact, high bodied, massive otoliths; OL : OH = 1.2-1.3; O H : OT about 3. Rostrum short, about Holotype: Fig. 221a-b, BSPG 1984 X 1629. - 13 16 % o f O L. Do rsal rim high, very irregularly and Type location: Kressenberg, Bavaria, lo catio n A – 2.5 m. coarsely ornamented. O stium wide; cauda with slightly do wnturned tip terminating clo se to po sterio r rim o f o tolith. Type formation: O iching Formation, Danian (P1c), trinidad- No o r weak ventral furro w. ensis zone.

Description. High bodied otoliths up to about 6 mm Paratypes: 4 specimens. length. Do rsal rim high, usually witho ut pro minent angles, Danian (P1c), trinidade nsis zone: Kressenberg, 3 specimens: loc. A – 2.5 m (same data as holotype) (Fig. 222 – BSPG 1984 X 1630; very variable ranging fro m almo st smo o th and regularly Fig. 223 – BSPG 1984 X 1631; not fig. – BSPG 1984 X 1632); curved to deeply and coarsely undulating. Ventral rim deep Selandian (P3a), angulata zone: Kroisbach, 1 specimen, fragmen- too , regularly curved, smo o th, deepest slightly anterio r o f ted: loc. 1.2 m N of Kch 4 (not fig. – BSPG 1984 X 1633). its middle. Rostrum sho rt, with ro unded tip. Excisura and antiro strum feeble o r absent. Posterio r rim with angular tip Name: Fro m co niugo (Latin) = basally related, referring to at about level of caudal tip. the basal relatio nship o f the species to Haemulidae. Inner face stro ngly bent alo ng ho rizo ntal axis, less in Diagnosis. Compact, high bodied, moderately thick oto- vertical axis. Sulcus lo ng, slightly deepened and slightly liths; OL : OH = 1.3-1.4; O H : OT about 3. Rostrum short, supramedian. Sulcus anterio rly o pened, po sterio rly terminat- blunt; no antiro strum o r excisura. Do rsal rim shallo w, ventral ing very clo se to po sterio r rim o f o tolith. O stium co nsider- rim deep. O stium wide, shallow; cauda long, with mark- ably shorter and wider than cauda; CaL : OsL = 1.4-1.7; edly do wnturned tip terminating clo se to po stventral rim o f OsH : CaH = 1.6-1.8. O stium slightly deepened; cauda o tolith. No ventral furro w. with slightly do wnturned mo derately po inted tip. Co lliculi po o rly marked. Do rsal depressio n large, mo derately distinct; Description. High bodied otoliths up to about 3.5 mm ventral furro w very feeble o r absent. length. Do rsal rim shallo w, usually with o btuse medio- and O uter face flat, smo o th o r with few radial furro ws. po stdo rsal angles, rarely smo o th and witho ut angles (possibly due to slight ero sio n). Ventral rim very deep, regularly Discussion. Dentex? solidus is easily reco gnized by its co m- curved, smo o th, deepest anterio r o f its middle. Rostrum pact, high bodied appearance. It resembles modern o toliths sho rt, blunt; no excisura o r antiro strum. Posterio r rim nar- of the subgenus Polyste ganus KLUNZINGER 1870 of the rowed with blunt termination above level of caudal tip. genus Dentex CUVIER 1814 (see SCHWARZHANS 2010b), Inner face strongly convex, smooth. Sulcus long, slightly where this species is tentatively placed and represents the deepened, less in ostium than cauda, and markedly supra- earliest reco rd o f such sparid mo rpho lo gy. median. Sulcus anterio rly o pened, po sterio rly terminating

Palaeo Ichthyologica 12 52

222 a

221

223 Fig. 221-223. Haemulon? conjugator n. sp. Kressenberg, Danian (P1c). 221, Ho lo type, BSPG 1984 X 1629. 222-223, Paratypes; 222, BSPG 1984 X 1630; 223, BSPG 1984 X 1631. – 20 × .

very clo se to po stventral rim o f o tolith. O stium co nsider- haemulid otoliths continue to be a common sight in Euro - ably shorter and wider than cauda; CaL : OsL = 1.8-2.0; pean warm inshore sediments. OsH : CaH = 1.5-1.8. O stium shallow; cauda with markedly do wnturned mo derately po inted tip. Co lliculi po o rly marked. Do rsal depressio n narro w, indistinct; no ventral Suborder Stromatoidei furro w. Family Stromateidae O uter face flat to slightly co ncave, smo o th. Genus indet. Discussion. These are typical, albeit very plesio mo rphic haemulid o toliths with their widened and rather shallo w Stromateidae indet. ostium, the markedly curved cauda and the smooth and (Fig. 224) distinctly co nvex inner face as well as the general appearance of the outline of the otolith. The widening of the Material: 1 specimen, Kroisbach, Austria, location Kch 1, O iching ostium and the bending of the cauda, however are much Formatio n, Thanetian (P4), upper pseudomenardii zo ne (BSPG less than in younger species of haemulids and therefo re 1984 X 1634). H.? co njugato r is interpreted as a basal, generalized rep- Discussion. A single, small o tolith with bro ken ro strum is resentative o f the family. considered to represent a Stromateidae because of its thin Hae mulo n? co njugato r represents ano ther early hae- appearance with the low curvature of the inner face, the mulid reco rd, easily distinguished by its co mpressed ap- absence of a clear downturn of the caudal tip, the lack of a pearance from such slender morphologies as H.? gulle nto psi ventral furro w and the delicate crenulatio n o f all rims. O ther NO LF 1978 from Belgium and Ukraine, H.? makare nko i plesio mo rphic stro mateo id o toliths have been repo rted from SCHWARZHANS & BRATISHKO 2011 fro m Ukraine and the Danish Selandian – Mupus sinuo sus (STINTON 1965) an indeterminated haemulid from the Paleocene of West- and a similar, but more compressed form as another inde- Greenland recorded by SCHWARZHANS (2004). In Eocene, terminated stro mateid in SCHWARZHANS (2003).

224 ac

d b

Fig. 224. Stromateidae indet. Kroisbach, Thanetian (P4), BSPG 1984 X 1634. – 20 ×.

Werner SCHWARZHANS 53

4. Faunal Reconstruction

4.1 Comparison of the Kressenberg and the Kroisbach faunal composition

O toliths fro m the Kressenberg lo catio n are all o f Danian bathypelagic representatives are the five species o f the o rder age. With 32 identified species (one species in open nomen- Stomiiformes, two of which account for the most common clature) it co ntains the richest early Paleo cene o to lith-based at Kro isbach (Argyripnus kroisbachensis and Progonostomia fish fauna kno wn to date. Kro isbach has mainly yielded primo rdialis), Nettastoma davejohnsoni (Nettastomatidae), o toliths fro m Thanetian strata (41 species, thereo f 5 in Dire tmus se rrative nte r (Diretmidae), Ogcocephalus? semen open nomenclature), and much fewer from the Selandian and Melamphaes? protoforma bo th the earliest reco rds o f (17 species, one in open nomenclature) (and the much less their families. Except fo r these elements, many o f the species sampled O ichinger Graben) (fig. 225). o f Kressenberg are also present at Kro isbach, but are usually less common. Congridae for instance are species rich, The o tolith-based fish fauna fro m Kressenberg sho ws a but not as common as in Kressenberg. The same is true for mo derately high diversificatio n index o f 13 (measured as mo st Berycifo rmes and Percifo rmes. In O phidiifo rmes, Bide n- the number of the most common species up to a percentage ichthys lapie rre i is much more common than Amphe ristus o f 90 o f the o tolith asso ciatio n). The do minant gro ups are ne o bavaricus. The ptero thrissid Pte ralbula co nchae fo rmis is primitive Perco idei, particularly Lactarius? simple x, which more common in Kroisbach than in Kressenberg, while Arius is the mo st co mmo n species at Kressenberg; O phidii- danicus is more common in Kressenberg than in Kroisbach. fo rmes with Amphe ristus ne o bavaricus and, less common, Another main difference is the presence of a moderately Bide nichthys lapie rre i; Anguillifo rmes with Rhynchoconger common merlucciid at Kroisbach (Palaeogadus? bratishkoi) intercedens and, less common, Gnathophis probus; and versus a co mplete lack o f G adifo rmes at Kressenberg Berycifo rmes with Centroberyx integer. Berycifo rmes o f (fig. 225). O nly a few o f the species fo und at Kressenberg the families Berycidae, Trachichthyidae, Ho lo centridae are missing fro m the Thanetian o f Kro isbach. Amo ngst the and Polymixidae and primitive Percifo rmes o f the families more common species at Kressenberg are Arius subtilis, Acropomatidae, Lactariidae, Serranidae, Sparidae and Bavarisco pe lus parvinavis, Plesiopoma elegantissima and Haemulidae and the family Co ngridae o f Anguillifo rmes Polyperca exserta. In a nutshell, this faunal asso ciatio n is also represent the mo st species-rich gro ups (fig. 225). All clearly indicative of an open marine environment with an o f these fishes, including the berycifo rm representatives, are unusually high diversificatio n index o f 20. In fact, it rep- interpreted as shallo w water neritic fishes. resents the earliest o pen marine o tolith-based fish fauna The fish fauna fro m the Thanetian o f Kro isbach differs known from the Paleocene, together with the two iso lated significantly fro m Kressenberg in the abundance o f o toliths species recorded from the late Paleocene of South Australia of fishes supposed to relate to open oceanic, mesopelagic (SCHWARZHANS 1985a). to po ssibly even bathypelagic enviro nments. Such meso- to

4.2 Regional comparison and paleoecological interpretation

Figure 225 depicts that the Paleo cene o tolith asso ciatio ns Paleo cene o tolith assemblages are the result o f inadequate of Kressenberg with 32 and Kroisbach with 41 species, sample sizes and would change when adequate collections totaling 54 species, are amongst the most species-rich become available. O n the other hand, the available data assemblages known from the Paleocene to date and co m- exhibit a degree of regionalization of the Paleocene o tolith pares to Denmark with 44 species (SCHWARZHANS 2003), asso ciatio ns fo r the vario us Euro pean basins which is rather Ukraine with 26 species (SCHWARZHANS & BRATISHKO high, higher than observed in Eocene faunas (fig. 225; see 2011), West G reenland with 24 (SCHWARZHANS 2004), also chapters 4.3 and 5.1). U.S. G ulf Co ast with 19 (NO LF & DO CKERY 1993) and The differences in the faunal co mpo sitio ns likely result fro m a Belgium with 14 species (NO LF 1978). combination of paleoecological factors that are bathymetric The number of species shared between Kressenberg and and climatic in nature and reflect geographic differences. Kroisbach with any of the mentioned other locations is Paleo eco lo gical interpretatio n o f o tolith findings depends mo derate, o ne co uld even say surprisingly lo w, at levels either o n strict co rrelatio n with habitats o f the nearest living between 25 % and 35 % of the respective faunas. Similar taxa, which gets increasingly pro blematical with geo lo gical o bservatio ns have been made by SCHWARZHANS & BRA- age because taxonomic positions become more uncertain TISHKO (2011) abo ut the degree o f similarity o f the fauna and because of possible changes in ecological habitats of fro m Luzanivka, Ukraine with tho se o f the o ther European fish gro ups during evo lutio n, o r relies o n paleo eco lo gi- Paleocene locations. None of these Paleocene faunas, cal assessments o f o ther presumably better kno wn fo ssil however, are as yet represented by the number of speci- gro ups asso ciated with the o toliths. In this light the fo llo wing mens as collected from the much more intensely studied paleoecological interpretation and evaluation should be Eocene, O ligocene or Miocene strata. Therefore it is still understoo d with a fair po rtio n o f cautio n. po ssible that part o f the o bserved differences between the

Palaeo Ichthyologica 12 54

Danian Danian Danian P1b P1c P2

uncinata- pseudobulloides-Z. trinidadensis-Z. Z.

Fig. 225. Co mpariso n table o f Paleo cene o toliths fro m Kressenberg and Kro isbach with o ther lo catio ns. 1.0 m 2.5 m 3.0 m 5.5 m 6.0 m 8.3 m 9.6 m 11.6 m 12.7 m 2.0 m ------Kressenberg A Kressenberg B 1 Kressenberg B 2 Kressenberg B 3 Kressenberg B 1 northern N Oiching Graben, 441) m ESE loc. (630 Kressenberg A Kressenberg A Kressenberg A Kressenberg A Kressenberg A Kressenberg A Kressenberg A Kressenberg A northern Oiching Graben, N 2 northern N Oiching Graben, 441) m ESE loc. (550 Kressenberg A Elopiformes Pterothrissidae Pteralbula conchaeformis 6211 family indet. Genartina hauniensis Albulidae Albula sp. Anguilliformes Anguillidae Anguilla pfeili n. sp. 1 Protanguillidae Bavariconger sp. 1 Congridae Conger illaesus 2 Gnathophis probus n. sp. 1 37 3 4 Paraconger vetustus n. sp. 21 Rhynchoconger angulosus 5 Rhynchoconger intercedens n. sp. 1 1 10 126 6 1 9 3 17 5 Gorgasia? turgidus n. sp. Congridae indet. 1 Congridae indet. 2 Nettastomatidae Nettastoma davejohnsoni n. sp. Siluriiformes Ariidae Arius danicus 84 3 Arius subtilis 24 11 Stomiiformes Gonostomatidae Progonostoma primordialis n. gen. n. sp. Progonostoma hagni n. gen. n. sp. Cyclogonostoma disciformis n. gen. n. sp. Sternoptychidae Argyripnus kroisbachensis n. sp. Valenciennellus kennetti n. sp. Aulopiformes Aulopidae Aulopus praeteritus n. sp. Chlorophthalmidae Paraulopus novellus n. sp. 4 2 Paraulopus postangulatus 24 3 4 5 Myctophiformes family indet. Bavariscopelus bispinosus 4126 1 112 Bavariscopelus parvinavis n. sp. 7 1 29 1 Danoscopelus schnetleri Gadiformes Merlucciidae Palaeogadus? bratishkoi n. sp. Gadidae Gadidae indet. Ophidiiformes Ophidiidae Ampheristus neobavaricus n. sp. 4 229 1 8 6 14 14 4 Palaeomorrhua sp. 1 Bythitidae Bidenichthys lapierrei 60 2 5 11 Ogilbia luzanensis 1 Lophiiformes Ogcocephalidae Ogcocephalus? semen n. sp. Beryciformes Berycidae Centroberyx apogoniformis n. sp. 1 32 3 Centroberyx fragilis Centroberyx integer 3 176 1 10 1 6 3 1 Kressenbergichthys kuhni n. gen. n. sp. 24 1 2 Trachichthyidae Trachichthys anomalopsoides n. sp. 2 3 1 Trachichthys impavidus n. sp. 11 1 Diretmidae Diretmus serrativenter n. sp. Melamphaidae Melamphaes? protoforma n. sp. Holocentridae Holocentronotus blandus n. sp. 11 Polymixiidae Polymixia polita n. sp. 14 1 2 Zeiformes family indet. Isozen janni Isozen mareikeae n. sp. Perciformes Acropomatidae Plesiopoma elegantissima n. sp. 7 1 7 Plesiopoma traubi n. sp. 2 12 20 2 Lactariidae Lactarius? simplex n. sp. 2 1 2 220 13 1 23 2 49 2 11 Serranidae Polyperca exserta n. sp. 25 1 4 1 Sparidae Platysepta kressenbergensis n. sp. 27 2 2 6 Dentex? solidus n. sp. 1131 Haemulidae Haemulon? conjugator n. sp. 4 Stromateidae Stromateidae indet. identified specimens 3 4 48 1 1106 2 1 55 13 78 7 174 4 23 Lapilli unidentified fragments and juveniles 1 2 37 1151 2 35 64 3 159 3 7 identified + unidentified specimens 4 4 2 85 1 2257 4 1 90 13 142 10 333 7 30

Werner SCHWARZHANS 55

West Selandian Thanetian Thanetian Den- GB-FR- Bavaria Ukraine US Gulf Green Belgium England P3a P4 P5 mark B land pseudo- Cret- angulata-Z. velascoensis-Z. Paleocene Eocene menardii-Z. aceous Thanetian - Selandian Selandian - - bei Kch 4 1,2 m N Kch 4 4 northern N Oiching Graben, (56 m SE “Wasserbehälter”) Kch 1 bed m south of sandstone marker 8 of Kch 1 10 m N Kch 11 Kch 11 Kch 11b Kch 12 Kch 12a Kch 13 Kch 14 (near 13) Danian total Selandian total Thanetian total of specimens number Maastrichtian Danian Selandian Danian Thanetian Ypresian Selandian Selandian 4 1 15 1 1 10 5 17 32 111 111 11 2123 22 113 451450 33 55 5 1 5 179 6 5 190 222 111 111 1 10 1 10 11 1 2 1 1 15 1 4 20 1819 41 41 41 444 555 61 61 61 23 23 23 444 46410 7 1 36 8 44 20 45 20 65 38 38 666 13 5 3 1 22 22 222 1 1 280 1 1 282 11 7 1 32 1 1 78 8 34 120 11 222 536541 1122 11 1 201 12 213 227229 2 1 11 1 6 3 12 21 3 7 12 3 7 22 333 222 1 1 1 221 5 217219 111 111 15 15 1 1 36 1 1 38 5 3 16 1 326 8 17 351 31 31 1 1 37 1 1 39 31115520 1415 111 6 33 6 331 2 3 1 8 8 1 1 3 1519 45 358 1922 number of identical species (dark blue) and related species (light blue) 22 22 22 3 13 6 2 6 2 0 0 6 12 338 5 9 5 1457 18 357 1839 7 1 2 2 0 1 0 13 12 45 6 691 2 8 1 17 13 1 1 3 2976 63 737 3783 number of valid species and cumulative with species in open nomenclature (in brackets) 33 (44) 37 (44) 22 (26) 8 (19) 21 (24) 8 (14) 5 (5) ~133 percentage of identical species (dark blue) and related species (light blue) 9 3527252825 0 0 Palaeo Ichthyologica 12 30 38 36 50 28 37 0 10 56

100 glimpses o f a pelagic o r bathyal Paleo cene fish fauna to date. 90 Percoidei The Selandian paleobathymetry in locations near Copenha- 80 Ophidiiformes gen was probably about 50 to 100 m according to analysis 70 shallow-water o f the mo llusc faunal asso ciatio n (SCHNETLER 2001) and 60 Beryciformes Gadiformes o f the benthic fo raminifera (LARSEN & JØ RGENSEN 1977). 50 As discussed in SCHWARZHANS (2003) the co mpo sitio n 40 + Myctophiformes of the fish fauna in the Danish Paleocene as reconstructed Stomiiformes 30 fro m the o toliths do es no t reflect a typical shallo w to middle Argentinidae 20 Aulopiformes shelf enviro nment, particularly because o f the abundance of Ariidae 10 macro urid o toliths. Recent macro urids are typically found on Elopiformes Congroidei 0 the deeper shelf, the continental slopes and bentho pelagic US Gulf W-Greenland Denmark Kroisbach Kressenberg Ukraine in the abyssal sea. Such elements in the Danish Paleocene

35 were interpreted as, either fishes that had migrated from deeper water in the North Sea Basin some 300 to 400 km 30 shallow-water away fro m the sampled lo catio ns, o r as an impressio n of Beryciformes Percoidei 25 ‘early’ Macrouridae that were adapted to shallower shelf enviro nments than their living co unterparts. 20 Gadiformes + 15 Aulopiformes The Paleo cene o tolith asso ciatio ns described fro m the Ophidiiformes yctophiformes Midway Formatio n o f Alabama by NO LF & DO CKERY M Stomiiformes 10 (1993), Belgium (NO LF 1978) and West Greenland dae Ariidae 5 Argentini (SCHWARZHANS 2004) all represent inner neritic faunas Congroidei Elopiformes witho ut any, o r at least any significant influence of pelagic US Gulf W-Greenland Denmark Kroisbach Kressenberg Ukraine o r bathyal fishes. The mo st distinctive shallo w water fauna has been sampled fro m the Selandian Tashlik Formation of Fig. 226. Abundance plot of main faunal elements of Paleocene o tolith asso ciatio ns. Upper graph: percent o f total o tolith assem- central Ukraine (SCHWARZHANS & BRATISHKO 2011) fro m blage; lower graph: number of species. fine grained shallow marine near shore sands with co ral patches. These differences in paleoenvironment between the Bavarian and Austrian lo catio ns and the Ukrainian location, which are about 1500 km apart, may help to In terms o f paleo bathymetry, Kro isbach pro bably represents explain the remarkable faunal difference between the two the faunal association with the highest degree of o pen areas expressed in the o tolith asso ciatio ns with o nly 27 % marine-pelagic influence known from any Paleocene o tolith o f the Ukrainian species being shared with the Bavarian/ co mmunity. KUHN (1992) interpreted the dark, silty clay- Austrian asso ciatio n. stones and clayey siltstones o f the O iching Formation as When considering paleoclimatic conditions, reference is depo sited o n a middle to o uter shelf po sitio n fro m the the made to SCO TESE’s (2001, www.sco tese.co m) terrestrial co mpo sitio n o f the Foraminifera. He relates the fo raminiferal climate history reco nstructio ns. The terrestrial Paleocene asso ciatio n with the Midway-type fauna o f BERGGREN & AU- climate in the vicinity o f the lo catio ns fro m which o toliths BERT (1975) co ncluding that the faunal co mpo sitio n of the have been obtained range from ‘warm temperate’ in the O iching Formatio n exhibits a mixture o f paleo enviro nments no rth to ‘ paratro pical’ o r ‘ subtro pical’ at the US G ulf Co ast from about 50 to 150 m water depth that have originated and the Bavarian, Austrian and Ukrainian lo catio ns. The fro m shallo w water co nditio ns o f a nearby island arc and co mpo sitio n o f the fish fauna as reco nstructed fro m the admixture fro m the o uter shelf. A similar paleo bathymetric o toliths suppo rts the general climatic belts as sho wn in po sitio n is sho wn in EGG ER et al. (2009) in their palinspastic SCO TESE. Indicative temperate faunal elements are Ar- restoratio n o f the Penninic Basin. The sedimentary sequence gentinidae and Gadiformes, both the main components in reco rds co nditio ns shallo wing upwards with o nly nearsho re the Paleocene of Denmark, West Greenland and Ellesmere sedimentary enviro nments in the o verlying Eocene. In the Island. In the mo re so utherly lo catio ns at Kressenberg and Paleocene, however, inshore shallow water species o r Kroisbach as well as in the Ukraine and Alabama, Argen- true deepwater fo raminifera o f the Velasco -type fauna of tinidae are missing entirely and G adifo rmes are either, very BERGG REN & AUBERT are missing. Neither KUHN (1992) rare and represented by only few species, or are missing. no r RASSER & PILLER (1999) seem to have co nsidered any Instead, shallo w water berycifo rms and perco ids co nstitute significant differences in the depo sitio nal enviro nments of the dominant faunal element. O phidiiforms, which fo rm the Paleo cene sediments at the lo catio ns Kro isbach and the major faunal element in the shallow water of the Kressenberg in their new litho stratigraphic classification of warm seas of the Eocene, are much less common during Paleogene rocks of Bavaria and Austria. Now such differ- the Paleocene and mostly not very species-rich, except at ences are o bvio us fro m the o tolith asso ciatio n, with abun- Luzanivka in Ukraine. dant Stomiifo rmes in Kro isbach and do minant Perco idei, Anguillifo rmes, O phidiidae and Berycidae in Kressenberg. Figure 226 summarizes the distributio n o f the main elements Kroisbach exhibits a pelagic open marine influence that is o f the fish fauna as reco nstructed fro m the o toliths both missing at Kressenberg. In additio n, mo st assumed shallow in terms of abundance (Fig. 226a) and species-richness water elements at Kressenberg are no t entirely missing at (Fig. 226b). The graphs demonstrate well that the Kro isbach Kroisbach, but are much more rare. In the absence o f kno wl- assemblage readily differs fro m all o thers in the abundance edge of a distinct deep water bathyal Paleocene oto lith-based o f stomiifo rm o toliths, asso ciated with a species-richness fish fauna o f the Velasco -type, Kro isbach and po ssibly the again o f stomiifo rms but also aulo pifo rms and primitive Danish Selandian locations of Copenhagen with their rather myctophiforms, all indicating open marine pelagic influ- common macrourid otoliths probably represent the only ence. Shallow water beryciforms, percoids and congroids

Werner SCHWARZHANS 57 all occur with many species but at low abundance. The ate faunal associations are characterized by a dominance subtro pical shelf asso ciatio ns o f Kressenberg, Ukraine and in abundance and species-richness of argentinids and Alabama are rich in percoids and shallow water berycifo rms. gadiforms, best expressed in the Danish Paleocene, while Kressenberg stands out because of the abundance and in West Greenland the species-richness, but more so the species-richness of congroids and Luzanivka in the Ukraine abundance of the two groups, is reduced at the expense in the species-richness o f o phidiifo rms. The warm temper- o f perco ids and shallo w water berycifo rms.

4.3 Paleobiogeographic interpretation of Paleocene otolith associations

O ur kno wledge o f the spatial distributio n o f Paleo cene fish is a common species in the Selandian of Denmark, much faunas reco nstructed fro m o toliths is still very much in its rarer in the Selandian of West Greenland and the Paleocene infancy, the only areas allowing some degree of interpreta- o f the Helveticum in Bavaria and Austria, tho ugh co nsider- tion being the North Atlantic, Europe and US Gulf Co ast. ably more common at Kroisbach than at Kressenberg. It Figure 227 depicts the faunal co mpariso n listing o f figure pro bably represents a wide-spread No rth Atlantic species 225 and a previo us analysis o f SCHWARZHANS (2004) indicative fo r o uter shelf enviro nments o f intermediate depth. o verlain as a stick co rrelatio n matrix o n a paleo geographic In Alabama and Ukraine rare juvenile, indistinctive o toliths reco nstructio n (60 Ma) co mpo sed fro m SMITH et al. (1994), have been found of a species of the genus Pte ro thrissus. KAZMIN & NATAPO V (1998), SCO TESE (2001), AKHMETIEV The main elo pifo rm representative in Alabama is Albula aff. (2010) and BLACKEY (2011). It depicts the co mparatively bashiana (FRIZZEL 1965); in Ellesmere it is Elo ps ramae ke rsi high amount of faunal regionalization expressed in the SCHWARZHANS 1985b. otolith associations of Paleocene age, even though some o f it may be o verprinted by differences in paleo enviro n- Congridae (Fig. 228b): Co ngridae are generally indicative ment (see above). o f clastic do minated shelf areas o f the tro pical and subtropical seas. In the Paleocene they are most species-rich A set o f specific distributio n maps o f systematic groups and at Kressenberg and moderately common at Kroisbach and key taxa is sho wn o n figures 228a-k and summarized below. in Denmark. There are two ubiquitous species – Conger Elopiformes (Fig. 228a): Ptero thrissidae are generally illae sus and Rhynchoconger angulosus – with Kressenberg wide-spread in Paleogene sediments, but rarely common yielding a number of more species not yet found outside the and never particularly species-rich. Pte ralbula co nchae fo rmis Helveticum. Rhynchoconger sp. recorded from the Paleocene

Fig. 227. ‘ Stick co rrelatio n matrix’ o f Paleo cene o tolith assemblages on the 60 Ma paleogeographic reconstructio n map. Number of species at a given lo catio n in bo ld (in brackets including species in open nomenclature), along 'sticks' refer to common species at any two locations. The paleogeographic reconstruction o n this and the following graphs of fig. 228 is composed and modified from SMITH et al. (1994), KAZMIN & NATAPO V (1998), SCO TESE (2001), AKHMETIEV (2010) and BLACKLEY (2011).

Palaeo Ichthyologica 12 58

1 3 2

1 Pte ralbula co nchae fo rmis Pte ro thrissus sp. *2 Albula aff. bashiana 3 Albula sp. 4 Elo ps ramae ke rsi

Fig. 228a-k. Distributio n maps o f Paleo cene key taxa o f o toliths on the 60 Ma paleogeographic reconstruction map. a. Elo pifo rmes.

1 5 3 1 2 4 5 6

1 Co nge r illae sus 2 Gnathopsis probus 3 He te ro co nge r astro ble maticus 4 Paraco nge r ve tissimus 5 Rhynchoconger angulosus 6 Rhynchoconger intercedens * Rhynchoconger sp. Fig. 228b. Congridae.

o f Alabama by NO LF & DO CKERY (1993) represents a dif- Argentinidae (Fig. 228c): Argentinidae are fishes indica- ferent species fro m R. angulo sus and R. intercedens observed tive of cool to temperate deeper shelf to continental slo pe in Euro pe. The lo catio n Luzanivka in Ukraine is remarkable enviro nments. It is po ssible tho ugh that during Paleocene fo r the first find o f a fo ssil garden eel – Heteroconger astro- they (also ) inhabited shallo wer temperate waters. The wide ble maticus SCHWARZHANS & BRATISHKO 2011 – indicative distribution and abundance of Arge ntina e rratica (RO EDEL of the warm shallow water environment. 1930) in Denmark, England, West Greenland and Ellesmere Island is pro bably o ne o f the best indicators fo r a continu-

Werner SCHWARZHANS 59

1 2 3 4

1 Arge ntina e rratica 2 Arge ntina lo ngiro stris 3 Pro targe ntino lithus balticus 4 Pro targe ntino lithus pro ce rus

Fig. 228c. Argentinidae.

2 5 3 1 4 5

1 Aulo pus prae te ritus 2 Aulo pus to rtus 3 Chlo ro phthalmus udo viche nko i 4 Paraulo pus no ve llus 5 Paraulo pus po stangulatus

Fig. 228d. Aulo pifo rmes.

ous marine connection between all those locations, and the Aulopiformes (Fig. 228d): Aulo pifo rm o toliths belo ng to the lack o f this species in the warmer, mo re so utherly locations more common ones in the Paleocene and are quite species- may also serve as a temperate pro xy and indicator o f the rich when compared to recent fish faunas. There was one asso ciated temperate no rthern Atlantic bio pro vince during widely distributed species fro m the US G ulf co ast to West the Paleocene and Early Eocene. The Paleocene of Denmark Greenland, Denmark, Bavaria and Austria – Paraulo pus has yielded a number o f further argentinid species. po stangulatus – and a number of other species restricted to the Danish, Ukrainian and Bavarian/ Austrian lo catio ns.

Palaeo Ichthyologica 12 60

2 5 2 3 1 2 5 4

1 Archaemacruroides ornatus 2 Raniceps hermani 3 Pale o gadus? antiquus 4 Pale o gadus? bratishko i 5 Palae o gadus sinangulatus

Fig. 228e. G adifo rmes (1).

1 3

1 2 3 4

1 Gadomorpholithus ponderosus 2 Molva palaeomorpha 3 Pro to co llio lus amo rphus 4 Mao rigadus ukrainicus 5 Melanonus ellesmerensis * three species o f Macro uridae Fig. 228f. G adifo rmes (2).

Gadiformes (Figs. 228e-f): G adifo rm o tolith distributio n G adidae (Fig. 228f) in West G reenland, Ellesmere Island, o ffer o ne o f the mo st co nclusive distributio n patterns fo r Denmark and England. These forms are again indicato rs o f paleoecological as well as paleobiogeographical purposes. a temperate no rthern Atlantic bio pro vince. Certain merluc- They are most abundant in temperate to cool environ- ciids of uncertain generic assignment have been found in ments (except fo r the glo bally distributed bentho pelagic Kro isbach and Ukraine, albeit two different species indicat- Macrouridae) as can be shown by the distribution pattern ing a certain degree of difference of the bioprovince of the o f Archaemacruroides, Palaeogadus (Fig. 228e) and the two lo catio ns.

Werner SCHWARZHANS 61

2 4

3 5 7 2 6 1

1 Amphe ristus ne o bavaricus Amphe ristus sp. *2 Ampheristus? seelandicus 3 Fie rasfe ro ide s buccule ntus 4 Gadophycus thulei 5 Gadophycus serratus 6 Palae o mo rrhua bulbus 7 Preophidion convexus

Fig. 228g. O phidiifo rmes (1).

2 3 4 2 3 1

1 Bidenichthys midwayensis 2 Bide nichthys lapie rre i 3 O gilbia luzane nsis 4 Bythite s? ro ze nbe rgi

Fig. 228h. O phidiifo rmes (2).

Ophidiiformes (Figs. 228g-h): The O phidiifo rmes repre- Ampheristus? seelandicus (KOKEN 1885) in Denmark and sent another species-rich group in the Paleogene of Europe. West Greenland, again supporting the concept of both areas During the Eocene, they form one of the most dominant forming part of an interconnected bioprovince, and Amphe- groups both in terms of abundance and species-richness, ristus ne o bavaricus fro m Kressenberg (rare at Kro isbach). while during the Paleocene their abundance is on a much Remarkable are the many, though mostly rare ophidi- lo wer level. O phidiids are represented by a large number ids fro m Luzanivka, Ukraine: Fie rasfe ro ide s buccule ntus of species (Fig. 228g), but only few of them are co mmon: SCHWARZHANS & BRATISHKO 2011, Gadophycis serratus

Palaeo Ichthyologica 12 62

2 3 5

3 4

4 1 2 3 4 2 3 4 9 6 7 8

1 Centroberyx anguincauda 2 Centroberyx apogoniformis 3 Ce ntro be ryx fragilis 4 Centroberyx integer 5 Centroberyx? tener 6 Kre sse nbe rgichthys kuhni 7 Trachichthys anomalopsoides 8 Trachichthys impavidus 9 Ce ntro be ryx? stringe ri

Fig. 228i. Shallo w water Berycifo rmes.

SCHWARZHANS & BRATISHKO 2011, Preophidion convexus Africa. This may in fact represent a ‘refugium’ distributio n (STINTON 1977) and Ho plo bro tula sp. (not shown), all spe- pattern in the Recent. Bavaria and Austria are particularly cies unknown from any of the other Paleocene locations, rich in shallo w water berycifo rm species, so me o f which but clo sely related to the rich o phidiifo rm asso ciation of are not recorded outside this area – Kressenbergichthys the Eocene of the southern North Sea Basin. Presumably, kuhni, Trachichthys anomalopsoides and T. impavidus. In this o phidiid asso ciatio n represents the western extend of a Alabama, Ce ntro be ryx? stringe ri NO LF & DO CKERY 1993 is shallow water Tethyan bioprovince of the Paleocene, which the most common species and there the only shallow water after opening of the Polish Trough spread along the southern berycifo rm (plus a single unidentifiable juvenile Centroberyx sho res o f the No rth Sea Basin during early Eocene establish- sp.) again supporting biogeographical differences to the ing a unifo rm bio pro vince characteristic fo r the warm clastic European fauna. shallo w seas o f the time and stretching fro m the Paris Basin to the Crimea at least (Fig. 229c). Bythitid o toliths are found Percoidei (Figs. 228j-k): Perco id o tolith-based species are in a similarly wide-spread distributio n (Fig. 228h) and are already very diverse in the Paleo cene sediments kno wn so often more common but less species-rich. The genus Bide n- far, altho ugh mo stly with a generalized, ‘ primitive’ mor- ichthys is represented by two species, one from the US Gulf pho lo gy. The Bavarian/ Austrian lo catio ns and Luzanivka in Coast – B. midwayensis (NO LF & DO CKERY 1993) – one Ukraine are particularly rich in perco ids with many species fro m Euro pe (Belgium and Helveticum) – B. lapie rre i – pos- that have not been found at other locations. Amongst the sibly indicative o f certain bio pro vincial differences between more wide-spread species are Acropoma? rosenkrantzi the two areas. O gilbia luzane nsis appears to be distributed SCHWARZHANS 2004 in Denmark and West Greenland throughout the European Paleocene. (Fig. 228j), Platysepta caribbaea (NO LF & DO CKERY 1993) in Alabama and West Greenland, Platysepta kressen- Shallo w water Beryciformes (Fig. 228i): Shallo w water bergensis in Bavaria, Austria and Denmark and Haemulon? berycifo rm o toliths o f the two families Berycidae and gulle nto psi (NO LF 1978) in Belgium and Ukraine. Trachichthyidae are particularly wide-spread and still very species-rich in the Paleocene deposits known to date, ex- Paleogeography (Fig. 229): The paleogeography and cept Ellesmere Island. The two mo st wide-spread species paleo -co astline reco nstructio ns o f the Late Cretaceous to in Europe are Ce ntro be ryx fragilis and C. integer which Eocene depicted in figure 229 are composed using ZIEGLER commonly occur together (except only C. fragilis in West (1990), SMITH et al. (1994), KAZMIN & NATAPO V (1998), Greenland and only C. integer in Belgium), but in varying FRENZEL et al. (1998), TORSVIK et al. (2000), SCO TESE abundance. The pattern of their changing proportions of (2001), AKHMETIEV (2010) and BLACKEY (2011). Naturally, abundance is not yet fully understood, but is probably reco nsiderable uncertainty exists co ncerning paleo -co astline lated to environmental rather than biogeographical reasons reco nstructio ns and several alternative analyses are found (see descriptive part). The so utherly, warmer lo catio ns in in literature. The land/ sea distributio n depicted here at- Europe are more species-rich than the northerly, temperate tempts to make best use o f the available published data, o nes, which is in co ntrast to the recent distribution of the but nevertheless reflects a generalized status and inevitably genera Centroberyx and Trachichthys mainly in the so uthern will co ntain inaccuracies and mistakes. subtro pical to warm temperate seas o f Australia and South

Werner SCHWARZHANS 63

3 4 1 2 5 6

1 Plesiopoma elegantissima 2 Ple sio po ma traubi 3 Acropoma? rosenkrantzi 4 Epigonus? tyassminensis 5 Lactarius? simple x 6 Apogon? rostrosus

Fig. 228j. Perco idei (1).

7 5 7 8 1 3 4 6 2

1 Polyperca exserta 2 Platysepta caribbaea 3 Platysepta kressenbergensis 4 Dentex? solidus 5 Platyse pta spatiata 6 Hae mulo n? co njugato r 7 Hae mulo n? gulle nto psi 8 Hae mulo n? makare nko i

Fig. 228k. Perco idei (2).

During Late Cretaceous (Fig. 229 upper panel), Europe 2003 and SCHWARZHANS 2010a) with data restricted to and North America were widely flooded by epicontinental a few locations in North America and Europe along the seas with relatively few and small to mo derately large paleo -subtro pical belt. The terminal Cretaceo us o to lith as- land masses in between, which would not pose effective sociations of the Maastrichtian and Campanian depict a barriers fo r marine live dispersal in either a lo ngitudinal clear distinctio n o f a paleo - No rth American/ West Atlantic o r a latitudinal directio n. Relatively little is known of the bioprovince and a paleo-European bioprovince (FRENZEL o tolith asso ciatio ns fro m the Late Cretaceo us (VO IGT 1926, et al. 1998) with only few species in common on both sides NO LF & DO CKERY 1990, NO LF & STRING ER 1996, NO LF o f the Atlantic (SCHWARZHANS 2010a).

Palaeo Ichthyologica 12 64

Fig. 229. Reco nstructio n o f fish-bio pro vinces during Maastrichtian, Paleocene and Early to Middle Eocene in Euro pe as reconstructed fro m o tolith asso ciatio ns. The paleo geo graphic reco nstructio ns are co mpo sed and mo dified fro m ZIEGLER (1990), SMITH et al. (1994), KAZMIN & NATAPO V (1998), FRENZEL et al. (1998), TORSVIK et al. (2000), SCO TESE (2001), BRINKHUIS et al. (2006), AKHMETIEV (2010) and BLACKLEY (2011).

Werner SCHWARZHANS 65

The Paleocene (Fig. 229 middle panel) topography of A. A temperate to co o l water bio pro vince was established Europe has been severely influenced by compression and throughout the North Sea Basin and extended into the inversio n during the ‘ Larimide’ phase o f the Early Alpine Labrador Sea to Western Greenland and the isolated record O rogeny. Western and Central Europe formed an un- in the Arctic Basin fro m Ellesmere Island. Judging fro m the interrupted land bridge separating a northern Temperate paleogeographic conditions and paleo-current systems Atlantic bio pro vince fro m subtro pical bio pro vinces estab- (see abo ve) this interco nnected bio pro vince was at least lished in the Helveticum and the Ukrainian Basin. The key active until the end of the Selandian and probably into the land bridge and marine gate areas were: Thanetian. It is depicted as a ‘ Temperate No rth Atlantic’ paleo bio pro vince o n fig. 229 (middle panel). It is charac- 1. The Thule Bridge caused by ejecting sheet basalt flo ws terized by abundance and richness of gadiform species from the Icelandic plume in the Faeroe / Shetland Basin (Fig. 228e,f) and co mmo n argentinid species (Fig. 228c), during the initial spreading of the sea between eastern bo th gro ups characteristic fo r temperate seas until nowa- Greenland and northern Europe. Some paleogeographic days and almo st entirely missing fro m the mo re so utherly reconstructions show a narrow marine connection persisting subtro pical lo catio ns. Ano ther aspect is the abundance of into Early to Middle Paleo cene times (KAZMIN & NATAPO V macro urid o toliths in the Selandian ro cks o f Denmark de- 1998) with flood basalt emplacement commencing during posited at moderate water depth of less than 100 m. The the Selandian (SØ RENSEN 2003), whereas o thers sho w Macro uridae are a family that in the Recent is typical fo r a complete obstruction of a marine gateway by the Thule a bentho-pelagic habitat on the continental slope and the vo lcanism (SMITH et al. 1994, TORSVIK et al. 2000). The 60 abyssal plains (see discussio n in chapter 5.4). Ma paleo geo graphic reco nstructio n o f KAZMIN & NATAPO V The warm subtro pical seas o f Euro pe have so far yielded (1998) depicts a co unterclo ckwise water current system o nly two faunas, the first fro m Kressenberg (Bavaria) and established in the Norwegian Sea and North Sea with a Kro isbach (Austria) in the ro cks o f the ‘ Helveticum’ alloch- branch leading through a gap in the Thule Bridge around the thonous and the second from Luzanivka in Central Ukraine. southern reaches of Greenland back North into the Labrador They represent rather different enviro nmental co nditio ns, Sea and Baffin Basin o f western G reenland. This co ncept but o n top they seem to also represent different paleobio- would explain best the great resemblance of the Selandian pro vinces: o tolith asso ciatio ns o f Denmark and western G reenland (SCHWARZHANS 2004) and is therefo re cho sen fo r the B. The faunal asso ciatio ns o f Bavarian and Austrian loca- Paleocene paleo-coastline reconstruction shown in fig. 229. tio ns are here termed the Helveticum paleo bio pro vince. Their composition show some semblance to the Maas- 2. The Channel/ Hampshire Basins area between southern trichtian o tolith-based fish fauna o f the same area and England and continental Europe is assumed to have become from northern Germany, more so than any of the other emergent during the Selandian and remained an effective Paleo cene o tolith asso ciatio ns o f Euro pe. The Helveticum land barrier (ZIEGLER 1990) until the Early Eocene sea level paleobioprovince thus could be interpreted as the succes- rise asso ciated with the PETM event (SLUIJS 2006). The area so r o f the mo re unifo rmly distributed Euro pean paleobio- again acted as an effective land barrier through mo st o f the pro vince o f Maastrichtian times. It do es no t differ from the O ligo cene to Plio cene times (ZIEGLER 1990, PO PO V et al. Paleo cene o tolith asso ciatio n o f Luzanivka in its principle 2006, see also discussio n in SCHWARZHANS 2010b). composition, and both are dominated by shallow water 3. The co mpressio n and inversio n o f the Polish Tro ugh Berycifo rmes, Perco idei, O phidiifo rmes and Aulo pifo rmes closed the marine connection of the North Sea Basin with the with Berycifo rmes and Perco idei as the mo st abundant. O n Ukrainian po rtio n o f the no rthwestern Tethys, which persisted the species level, however, only few species are co mmon into Danian times, during the Selandian (ZIEGLER 1990). to bo th areas except fo r the mo re unifo rmly distributed During a phase of subsequent ‘relaxation’ movements shallo w water Berycifo rmes (Figs. 226, 228d,g,i,j,k). The (KOCKEL 2003) shallo w marine co nditio ns re-co nnected the faunal asso ciatio n o f Bavaria and Austria further differs No rth Sea Basin with the Tethys / Paratethys intermittently fro m the Ukrainian o ne in the richness o f Anguilliformes. during Eocene and Early O ligo cene times (ZIEGLER 1990, The Helveticum paleobioprovince does not seem to have BRINKHUIS et al. 2006, STOREY et al. 2007). been succeeded by any paleobioprovince in Europe, but there are a few species related to subsequent species fro m 4. The Western Siberian Basin was connected southwards the European Eocene (first Paleocene, then Eocene): Anguilla to the Tethyan realms through the Turgay Gateway during pfe ili – A. re ctangularis, Gnathophis probus – G. ro se nblatti the Maastrichtian (KAZMIN & NATAPO V 1998) but became and G. schepdaalensis, Paraulo pus po stangulatus – P. davisi, briefly interrupted at the terminal Cretaceo us and the Early Arius subtilis – A. subre ctangularis, Ampheristus neoba- Paleo cene (BENIAMO VSKI 2007). Re-established in the Se- varicus – A. to liapicus. KAZMIN & NATAPO V (1998) sho w landian o r Late Danian (BENIAMO VSKI 2007, AKHMETIEV diverging currents in so uthern Euro pe, co unterclo ckwise & BENIAMO VSKI 2009), it is tho ught to have represented north to south in the Ukrainian Basin and counterclo ckwise a marine gateway until its final destructio n during Middle so uth to no rth in the Penninic Basin (including the rocks Eocene (Lutetian) times (AKHMETIEV 2010, RADIO NO VA now incorporated in the allochthonous ‘Helveticum’). Such et al. 2003). counteracting current systems could have been respo nsible to so me extent fo r the discrepancies o f the fish fauna ob- In terms o f ichthyo lo gical bio pro vinces as reco nstructed from served in the two areas. fossil otoliths, the Paleocene epoch shows a much larger degree of regionalization in Europe than the preceding Late C. The faunal asso ciatio n o f Luzanivka, Ukraine (SCHWARZ- Cretaceous period, mainly as a result of the geographic HANS & BRATISHKO 2011) is particularly rich in shallow wa- separation of basins and marine areas by emerged land ter Berycifo rmes, Perco idei and O phidiifo rmes, which for the ridges as described above: mo st part represent species different fro m the o ther known Euro pean Paleo cene realms (except fo r the Berycifo rmes). The most remarkable aspect in terms of biogeography of

Palaeo Ichthyologica 12 66 the Ukrainian o tolith asso ciatio n ho wever is its resemblance European Tethyan paleobioprovince; 2. the paleoecological with the well known Eocene fish fauna of the warm shal- differences between Luzanivka and Kressenberg were larger lo w seas o f Euro pe (Paris Basin, Belgian Basin, England, than assessed in this study, masking the presence o f a more G ermany and Denmark). There are relatively few species westerly reaching near shore Tethyan fauna during the Pale- common to both strata (Preophidion convexus), but several ocene; or 3. a marine gateway existed further to the east o f the Paleo cene species o f Luzanivka are directly related (thro ugh the Polish Tro ugh, see belo w) allo wing o f Tethyan to the common Eocene ones, specifically in O phidiiformes migratio n into the No rth Sea Basin bypassing the Aquitaine and Perco idei. Typical ‘ pairs’ are (first Paleo cene, then Basin in the west. In the complete absence of Eocene Eocene): Chlo ro phthalmus udo viche nko i – C. deflecticauda, o toliths fro m the ‘ Helveticum’ o r co mparable intermediate Arius subtilis – A. subre ctangularis, Palae o gadus? antiquus geographic locations no further assessment can be made. – P.? papillo sus, Fie rasfe ro ide s buccule ntus – F. subre gularis, Another marine gateway (re-)opened further in the east O nuxodon sp. – O . fimbriatus, Gadophycis serratus – G. ova- acro ss the Polish Tro ugh as a result o f tectonic ‘relaxatio n’ lis and G. bramscheensis, Ho plo bro tula sp. – H. le riche i and sagging (KOCKEL 2003) following the Paleocene compres- H. ro busta, Epigonus? tyassminensis – E.? selsiensis, Hae- sion, inversion and exhumation. When exactly a marine con- mulo n? makare nko i – H.? sylve stris. The Paleocene otolith nection became re-established between the south-eastern asso ciatio n o f Luzanivka is therefo re interpreted here as North Sea Basin and the Tethys is debated, ranging fro m representing a European Tethyan paleobioprovince. Early Eocene (BRINKHUIS et al. 2006, STOREY et al. 2007) The subtro pical No rth American o tolith-reco nstructed to Late Eocene (ZIEGLER 1990). The few o toliths recently de- fish fauna (not shown on fig. 229) remains quite different scribed fro m the Lutetian o f the Crimea (BRATISHKO 2009) fro m its Euro pean co unterparts, like in the Late Cretaceous, show total congruence with contemporary faunas from and also into the Eocene. Belgium, France and Germany indicating that a unifo rm Euro pean Tethyan palaeo bio pro vince was firmly established The Eocene (Fig. 229 lower panel) sees a renewed pattern by that time, supporting the concept of a marine gateway o f marine co nnectivity in Central Euro pe, mo st o f which thro ugh the Polish Tro ugh latest during Lutetian. The much were short lived and became abandoned later in the Eocene better known Late Eocene otolith-based fish fauna from the o r during the O ligo cene. First the western Tethyan prob- Mandriko vka suite o f the Crimea (MÜLLER & RO ZENBERG ably became connected with the western North Sea Basin 2003, RO ZENBERG 2003, BRATISHKO 2010) sho ws a through a marine gateway via the Aquitaine and Channel continued close correlation with Middle and Late Eo cene Basins, pro bably as early as the Ypresian and pro bably as a asso ciatio ns o f the No rth Sea Basin. result o f eustatic sea level rise during the Paleo cene-Eocene The Ypresian o tolith asso ciatio ns fro m the Londo n Basin Thermal Maximum (PETM) (BRINKHUIS et al. 2006, SLUIJS sho w so me specific elements (STINTON 1965) no t kno wn 2006, SLUIJS et al. 2008, STOREY et al. 2007). fro m the mo re so utherly lo catio ns in the No rth Sea Basin. A rich otolith-based fish fauna has been recorded fro m So me o f them represent typical temperate No rth Atlantic the Aquitaine Basin (NO LF 1988) and Belgium (STEURBAUT elements of argentinids and gadiforms of the same species & NO LF 1990). The Belgian fauna includes many of the as in the Paleocene of Denmark (Argentina erratica, Archae - Tethyan elements listed abo ve as successo rs o f the Paleocene macruro ide s o rnatus) or related to them (Pro to co llio lus Luzanivka fauna, but the fauna of the Aquitaine Basin is eocenicus, Palaeogadus pinguis). This indicates that certain quite different fro m the Belgian fauna. Acco rding to STEUR- lineages o f temperate fish gro ups persisted acro ss the PETM BAUT & NO LF (1990) this difference is mainly due to the event in the No rth Sea away fro m its warm so uthern sho res. Aquitaine Basin having been well exposed to the oceanic realm with pelagic species co nstituting an impo rtant po rtio n Conclusion. The otolith-based Paleocene fish fauna of o f the fauna. [In this respect the many stomiifo rm o toliths Bavaria and Austria, interpreted as the successo r o f the in the Aquitaine Basin co rrelate well with tho se from the Euro pean paleo bio pro vince o f Maastrichtian times as known Paleocene of Kroisbach, while the abundance of myctophids from Bavaria and northern Germany, contracted however of the genus Diaphus in the Aquitaine Basin is no t shared to the Helveticum paleo bio pro vince (Fig. 229 upper and with the mo re primitive myctophifo rms fro m Kro isbach.] middle panel). The difference o f the o tolith asso ciation of The Belgian o tolith-based fish fauna fro m the Ypresian Bavaria and Austria to the o ne fro m Luzanivka, Ukraine, has little resemblance with the Paleocene fauna of the North which is thought to represent a European Tethyan paleobio- Sea Basin and o nly mo derate similarity with the Helveticum province, is both of biogeographic and paleoenvironmental based o n relatively few species (see abo ve). The fact that nature and it is difficult with the present kno wledge to assess the Luzanivka fauna has mo re similarity with the Ypresian which differences are due to which effect. There are no fauna of Belgium than the geographically intermediate data available o n Eocene o toliths fro m Bavaria o r Austria. fauna fro m the Helveticum o f Kressenberg can o nly be Elsewhere in the well explored European Eocene, no o tolith explained by three alternatives: 1. the fo rmer Helveticum asso ciatio n is kno wn that co uld qualify as a faunal succes- paleo bio pro vince o f the Paleo cene lo st its distinction in the so r to the Helveticum paleo bio pro vince o f the Paleo cene Early Eocene and became incorporated in an advancing (Fig. 229 lo wer panel).

Werner SCHWARZHANS 67

4.4 Paleocene otolith biostratigraphy

O ur knowledge of Paleocene otolith-based fish faunas is still Ma number of species sparse, as can be seen from the stratigraphic species distri- 40 bution plot of figure 230. As stated above, the Paleocene fauna represents a lo gical successo r o f the Maastrichtian fauna of the same area, despite of the fact of the many Lutetian 45 ~185 extinctio ns acro ss the K-T bo undary. Since a successo r in the Eocene paleogeography is not known, a stratigraphic analysis o nly makes sense fo r the Maastrichtian to Paleocene 50 Middle Eocene interval o f Bavaria and Austria, while co rrelatio n with Eocene o toliths is merely o f general value. Ypresian ~133 Early Figure 231 summarizes the stratigraphic o ccurrences of a 55 number o f fish gro ups which are particularly well represented in the Paleocene and Late Cretaceous of Bavaria and Austria. Thanetian 49 In additio n to three species that have survived acro ss the K-T 60 extinctio n event – Arius danicus, Paraulo pus po stangulatus Selandian 57 91 and Bavarisco pe lus bispino sus – a number of lineages have

as well (first Maastrichtian, then Paleo cene): Pte ralbula fo - Paleocene Danian 65 35 re yi – P. conchaeformis, Bavarico nge r po lle rspo e cki – B. sp., Amphe ristus bavaricus – A. neobavaricus (and A. to liapicus in

Early Eocene), Bide nichthys cre pidatus – B. lapie rre i, Centro- ) Maastrichtian 70 39 be ryx te ume ri – C. integer (and C. eocenicus in Early Eocene) part ( and Plesiopoma otiosa – P. traubi. The species of the genera Amphe ristus, Bide nichthys, Centroberyx and Plesiopoma 51 are particularly common and can be used to distinguish Campanian 75 bio stratigraphically between Maastrichtian and Paleocene. (part)

The Paleo cene sequence fro m Bavaria and Austria is ex- Late Creatceous ceptio nally well do cumented by o toliths, while mo st other 80 lo catio ns studied in the past yielded o toliths fro m sho rter Fig. 230. Develo pment o f the o tolith species richness fro m Maas- chro no stratigraphic intervals. It is readily visible fro m figure trichtian to Eocene. Eocene data are estimated due to lack of 231 that there is little bio stratigraphic useful variatio n o f co mprehensive revisio n. o tolith diversity within the Paleo cene o f Bavaria and Austria. For tho se few effects that may be reco gnized it remains unclear whether they are o f bio stratigraphic value or are The Early Eocene (Ypresian) o f many o f the No rth Sea Basin related to lo cal enviro nmental differences between the two locations in Belgium, England and France has yielded some lo catio ns Kressenberg (Danian) and Kro isbach (Selandian species related to fo rms fro m the Paleo cene o f Bavaria and and Thanetian) (see chapter 4.1). So far, a few species Austria and pro bably descended fro m them (first Paleocene, are exclusively kno wn fro m the Danian (Bavarisco pe lus then Eocene): Rhynchoconger intercedens – R. eocenicus, parvinavis and Plesiopoma elegantissima), one from the Arius subtilis – A. subre ctangularis, Paraulo pus po stangu- Danian and Selandian (Rhynchoconger anglosus) and latus – P. davisi, Amphe ristus ne o bavaricus – A. to liapicus, one from the Selandian (Genartina hauniensis), the latter Centroberyx integer – C. eocenicus, C. fragilis – C. ingens. also co rro bo rated superregio nal with finds in Denmark Apart fro m these deemed persistent lineages there are a (SCHWARZHANS 2003). Due to the regio nal differences few more linking the Paleocene fauna of Luzanivka with the o bserved in the Paleo cene o tolith asso ciatio ns so far studied Eocene of the North Sea Basin, and many new groups in the (see chapters 4.2 and 4.3) no superregional biostratigraphic Eocene that o verall result in a quite different appearance of co rrelatio n is attempted. the Eocene o tolith asso ciatio ns fro m any o f the Paleocene asso ciatio ns so far kno wn.

Palaeo Ichthyologica 12 68

Haemulon? pulcher Haemulon? gullentopsi* Haemulon? kokeni Haemulon? ovalis Haemulon? gibbosus Haemulon? sylvestris Haemulon? makarenkoi* Haemulon? conjugator rs rs reflect inferred o ccurrences bridg- Plesiopoma elegantissima Plesiopoma traubi Plesiopoma otiosa Centroberyx anguinicauda* Centroberyx apogoniformis

Centroberyx ingens * species not recorded from Bavarian/Austrian Paleocene Centroberyx fragilis Centroberyx eocenicus Centroberyx integer Centroberyx teumeri Bidenichthys lapierrei Bidenichthys crepidatus Ampheristus traunensis Ampheristus brevicauda and selected species from European Eocene. Grey ba Ampheristus toliapicus Ampheristus neobavaricus Ampheristus bavaricus Danoscopelus schnetleri Bavarisopelus parvinavis Bavariscopelus bispinosus

Paraulopus davisi Paraulopus novellus Paraulopus postangulatus Arius subrectangularis Arius subtilis Arius danicus

Rhynchoconger eocenicus

Rhynchoconger intercedens Maastrichtian and Paleo cene o f Bavaria and Austria Rhynchoconger angulosus Rhynchoconger piger Bavariconger sp. Bavariconger pollerspoecki Pterothrissus angulatus Pteralbula conchaeformis Pteralbula foreyi Ma 55.8 58.7 61.7 70.6 65.5 40.4 48.6 P 2 P 5 P 4 P 8 P 7 P 6 P 3 P 9 P 1 a Late P 12 P 11 P 10 Early P 1 c P 1 P 1 b P 1 Danian Lutetian Ypresian Selandian Thanetian Stratigraphic range o f key o species tolith fro m the

Eocene Paleocene Maastrichtian Fig. 231. ing n distributio gaps.

Werner SCHWARZHANS 69

5 Phylogenetic Analyses and Evolutionary Interpretation

5.1 The Paleocene fish fauna: Linking the Late Cretaceous with the modern bloom

Maas- Paleocene Paleo- Paleo- Eocene Oligocene 5.1.1 Evolutionary interpretation trichtian Bavaria cene cene Germany Germany Bavaria and Ukraine Denmark (example) (example) % Austria 100 – O toliths, unlike skeleton findings, do no t allo w a direct interpretatio n o f the nature o f the fish itself. Instead, interpretatio n o f fo ssil o toliths stro ngly depends o n co rrelatio n with Recent material. This dependence of course puts Category 1 certain limitatio ns o n the systematic use o f o toliths. While 90 – 18 % co rrelatio n with Recent fishes o f fo ssil o toliths from Neogene strata is usually straight fo rward, this beco mes increasingly difficult the o lder the fo ssils are. As a rule o f thumb one may expect predo minantly differences at the species level to 80 – about O ligocene times and increasingly at the genus level Category 1 Category 1 during Eocene times. Beyo nd, particularly prio r to the K-T 45 % 46 % Category 1 extinctio n event, o ne must always co nsider the po ssibility to Category 1 54 % find o toliths o f extinct higher taxa as evidenced by skeleton 50 % based teleo sts fro m which o toliths are co mpletely unknown. 70 – It is therefo re mandatory to exercise great care when at- Category 2 tempting to co rrelate Late Cretaceo us o toliths with those of 32 % recent fishes, particularly if the results seem to co ntradict Category 1 skeleton findings. O toliths fro m the Early Cretaceo us o r 77 % the Jurassic mo stly sho w so little resemblance to living taxa 60 – that they can only be interpreted in a very general manner (SCHWARZHANS 1996, 2010a).

SCHWARZHANS (1996) presented a scheme o f fo ur mo rpho lo gic-evo lutio nary catego ries designed to allow 50 – Category 2 Category 2 14 % a better handling and interpretatio n o f o tolith data. The 11 % fo ur mo rpho lo gic-evo lutio nary catego ries were defined Category as fo llo ws: 2a 10 % Catego ry 1 – persistent taxa: Mo rpho lo gies that have Category 2 40 – not altered significantly when compared to those of Recent 12 % o toliths. Category 3 Catego ry 2 – extinct specialized taxa: Extinct specialized 15 % Category 2 9 % mo rpho lo gies witho ut apparent affinities to living taxa. Category 3 – Category 3 Catego ry 3 – extinct plesio mo rphic taxa: Plesio mo rphic 30 Category 3 21 % 8 % mo rpho lo gies which are o ften attributable to living families, 41 % but are of problematic allocation due to their generalized appearance. Catego ry 4 – ‘ missing links’ : A so mewhat info rmal 20 – heading attempting to combine certain plesiomorphic mor- Category 3 19 % phologies thought to be situated near major dichoto mies in Category 2a Category 4 the phylo geny o f ‘ mo dern’ teleo st gro ups. The distinction 13 % Category 4 towards catego ry 3 (extinct plesio mo rphic taxa) is fluent. 28 % 26 % 10 – Category 4 Category 1 co ntains all tho se species that can be attributed 21 % Cat. 2 2 % to extant genera with sufficient certainty, i. e. some Elopi- fo rmes, many Anguillifo rmes, Arius, Aulo pifo rmes, Bide nich- Category 4 Category 4 Category 3 9 % 10 % 5 % thys and O gilbia o f the Bythitidae, Centroberyx, Trachichthys, Cat. 4 3 % Dire tmus and Polymixia o f the Berycifo rmes here totaling 0 – 46 %. Interestingly, it co mprises mainly fishes that are either Fig. 232. Co mpo sitio n o f o tolith asso ciatio ns in the light o f evolu- characterized as ‘ o ld’ gro ups (Albulidae, Ptero thrissidae, tio nary catego ries fo r key lo catio ns o f the Euro pean Early Tertiary Anguillifo rmes and Aulo pifo rmes) o r ‘ survivo rs’ o f the K-T and Maastrichtian. extinctio n event which have transfo rmed into ‘ living fo ssils’ (Centroberyx, Trachichthys and Polymixia).

Category 2 contains less species summing up to about 11 %. Kressenbergichthys and Ho lo ce ntro no tus, others are newly So me are ‘ survivo rs’ o f the K-T extinctio n event fo r o nly a evolved specialized groups that apparently have become sho rt while (o r suspect o f it) like Bavarico nge r, Amphe ristus, extinct later such as G e nartina o r Danoscopelus.

Palaeo Ichthyologica 12 70

Category 3 is similarly large with abo ut 15 % co ntaining a the Paleocene of the North Atlantic Temperate bioprovince few survivo rs fro m the Cretaceo us such as Bavarisco pe lus (see chapter 4.3). Their lacking in o lder strata and relative and Plesiopoma, but mo stly difficult to assess co ngrids and scarcity elsewhere in Paleocene and Eocene strata is more Cyclogonostoma. likely due to lack of knowledge of adequate temperate faunas than o f phylo genetic significance. Berycifo rmes are Category 4, the ‘ missing links’ represent a fairly large gro up a dominant group during Late Cretaceous with many now o f abo ut 28 % in this assessment, mo stly due to the large extinct lineages, but as o toliths do cument, they are still number of highly plesiomorphic, generalized morphologies abundant during the Paleocene (Fig. 233), albeit with taxa found in the many species of the Percoidei, but also other o f gro ups which are mo stly still represented no wadays. fo rms such as Progonostoma, Palae o gadus? bratishko i, Ogcocephalus? semen, Melamphaes? protoforma and Isozen of an undetermined family of the Zeiformes. These are representatives o f higher systematic units which have 5.1.2 Influence of the K-T boundary expanded in speciation during post-Paleocene times. extinction event for teleost evolution When comparing the findings from Kressenberg and Kro isbach with similarly well kno wn o ther Paleo cene (based on otolith analysis) otolith-based fish faunas from the Ukraine and Denmark (Fig. 232) (SCHWARZHANS 2003, SCHWARZHANS & BRA- The Maastrichtian-Paleocene sequence in Bavaria and TISHKO 2011) it beco mes o bvio us that the relatio n o f the Austria is unique in Euro pe so far to have yielded a nearly fo ur catego ries are rather similar despite the fact that the uninterrupted sequence of otolith assemblages across faunal co mpo sitio ns themselves are rather different at all the K-T boundary extinction event and unique overall in three lo calities. In co ntrast, the Maastrichtian fauna shows the wealth o f o tolith related data fro m that time interval a distinctly different co mpo sitio n o f the fo ur respective cat- (Fig. 234a-b). ego ries (Fig. 232) (SCHWARZHANS 2010a), naturally with As no ted in chapter 4.4 three species survived the a low percentage of persistent taxa and a high percentage K-T boundary extinction event and a number of genera o f specialized taxa that went extinct at the K-T bo undary (including genera o f uncertain relatio nship) did as well. In event, but also with a comparatively low amount of taxa total 14 genera became extinct out of 31 observed in the asso ciated with ‘ missing links’ . Maastrichtian o tolith assemblage o f Bavaria, co rresponding Two columns have been added in figure 232 as examples to an extinction rate of about 45 % (Fig. 234a,b). This is fo r the Eocene (fro m SCHWARZHANS 2007) and O ligo cene higher than any extinctio n rate so far o bserved at any time (fro m SCHWARZHANS 1994) which sho w an ever increasing during the Tertiary and it do es indeed suppo rt the prime amount of category 1 and a decrease of category 4 (and o rder o f the event also fo r the Teleo stei. category 3 since the O ligocene) as would be expected. The extinctio n events o ccur acro ss the entire o tolith-based Interestingly, the catego ry 2 o f extinct specialized morpholo- teleost fauna containing many groups of the evolutio nary gies still co ntains so me 15 % to 19 %. Ho wever, mo st of category 2, which often can only be tentatively associated these extinct specialized Eocene and younger forms can be with living taxa and thus indicate extinctio n o f a considerable co nveniently placed in extant families and o ften close to number of suprageneric taxa as well, for instance in O steo- extant genera. Therefore a Category 2a is being introduced glo ssifo rmes, Albulo idei, Stomiifo rmes, Myctophifo rmes and fo r o tolith o f these taxa in yo unger strata to allo w a more Beryciformes. O f the 14 genera that became extinct at the meaningful comparison. The proper category 2 diminishes K-T bo undary, seven are fro m the Berycifo rmes. These extinct rapidly during Eocene and becomes nearly absent during supposed beryciform taxa pose a certain problem to inter- O ligocene times. Most faunal elements placed in catego ry pretatio n because o f their highly derived o tolith morphology 2a are from the many extinct ophidiiform taxa adapted to (SCHWARZHANS 2010a), leaving their systematic po sition shallow warm clastic seas during Eocene and in O ligocene in a preliminary status until such o toliths have been found are composed of several now extinct gadiform lineages in ‘ in situ’ . the No rth Sea Basin. O f the 17 genera which supposedly survived the K-T boundary event, not less than five become extinct subse- Conclusion. The statistic pattern o f the co mpo sitio n o f quently during Paleocene and three more during the Eocene. the evo lutio nary catego ries (Fig. 232) sho ws that the Late This means that only nine ‘genera’ (including tentatively Cretaceo us time is do minated by extinct taxa bo th o f the assignments and genera in open nomenclature) of 31 ob- categories 2 and 3, while the Paleocene is particularly rich served in the Maastrichtian o f Bavaria carry thro ugh until in mo rpho lo gies co nsidered in catego ry 4 (‘ missing links’ ). today (co rrespo nding to the evo lutio nary catego ries 1 and This suggests that the Paleo cene was an impo rtant transient 4 equaling 27 %, see Fig. 232). time in the evo lutio n o f many teleo sts at a higher systematic level linking the Late Cretaceo us largely extinct diverse tel- A total of 61 new records are added during various times eost fauna with the onset of the modern bloom which then of the Paleocene of a total of 78 genera observed in the rapidly evolved with the onset of the Eocene. European Paleocene (78 %). Although many of these new The main shifts o f faunal co mpo sitio n thro ugh this faunal entrants are fro m paleo bio pro vinces unrelated to the time interval invo lves 5 o rders o f teleo sts – Anguillifo rmes, Maastrichtian fauna fro m Bavaria (Fig. 229), their unusual G adifo rmes, O phidiifo rmes, Berycifo rmes and Percifo rmes high amount nevertheless underlines the importance of the (Fig. 233) and is subject o f further detailing and discussio n K-T bo undary event fo r the evo lutio n o f the Teleo stei. It is in the fo llo wing chapters. As an o verall trend Anguillifo rmes, further an indicatio n o f the evo lutio nary dynamic that the O phidiifo rmes and Percifo rmes sho w significant radiatio n Teleostei were undergoing during Paleocene. It is particularly events during Paleocene and Eocene, almost explosive in the large amount of generalized, plesiomorphic morpholo- the latter fo r O phidiifo rmes and Percifo rmes. G adiformes gies o f perco id o toliths that adds an impo rtant new element already exhibit a wealth o f, mo stly persistent, lineages during to the Paleo cene o tolith asso ciatio ns (see chapter 5.5). The

Werner SCHWARZHANS 71

Lutetian Ypresian Thanetian Selandian Danian Maastrichtian Eocene Paleocene Cretaceous

Pleuronectiformes

Scombroidei

Stromateoidei Gobioidei

Trachinoidei Perciformes

Percoidei

Stephanoberycoidei

Polymixioidei Holocentroidei

Berycoidei Beryciformes extinct indet. Berycoidei and Holocentroidei

Bythitidae

Ophidiidae

Carapidae

Macrouridae

Gadidae Merlucciidae Lotidae Ranicipitidae Gadiformes Ophidiiformes Maorigadus and Bregmacerotidae

Myctophidae

Myctophiformes indet.

and Synodontidae Aulopidae and Aulopiformes

Myctophiformes Chlorophthalmidae

Gomostomatoidei

Argentinoidei

Heterenchelyidae

Congridae and Nettastomatidae

Protanguillidae Anguilliformes Stomiiformes

Anguilloidei

Fig. 233. Stratigraphic summary chart depicting numbers o f species o f key teleo st gro ups fro m Maastrichtian to Middle Eocene. Eocene data are estimated due to lack of comprehensive revisio n.

Palaeo Ichthyologica 12 72

Maastrichtian Danian Selandian Thanetian Ypresian (part) Cretaceous Paleocene Eocene Hymenocephalus Macrouridae Coryphaenoides Coelorinchus

Gadidae † Protocolliolus † Molva Lotidae † † Gadomorpholitus

† Palaeogadus Merlucciidae †

Gadiformes Palaeogadus?

Ranicipitidae Raniceps

? Euclichthyidae † Archaemacruroides †

Gadiformes indet. † Maorigadus ?

Percopsiformes indet. Percopsis? ?

† Danoscopelus † Myctophiformes indet. phi- Mycto- formes † Bavariscopelus †

Chlorophthalmus Chlorophthalmidae Paraulopus

† Archaulopus † formes Aulopi- Aulopidae Aulopus

Stomiatoidei indet. † Palaeostomias † Valencienellus Sternoptychidae Argyripnus † Auriculithus †

† Cyclogonostoma † Gonostomatidae Stomiiformes † Progonostoma †

Bathylagidae † Protobathylagus †

† Protargentinolithus † Argentinidae formes

Salmoni- Argentina

Ariidae - Siluriformes Arius

Clupeidae - Clupeiformes Clupea?

Hoplunnis Nettastomatidae Nettastoma Heterenchelyidae Heterenchelys? Conger ? Muraenesox Pseudophichthys Rhynchoconger Congridae Paraconger Heteroconger Gnathophis Anguilliformes Conger † Alaconger

Anguillidae Anguilla?

Protanguillidae † Bavariconger †

Albuloidei indet. † Genartina near Pterothrissidae † Pollerspoeckia †

Albulidae Albula

Pterothrissus Pterothrissidae Elopiformes † Pteralbula †

Osteoglossiformes indet. † Kokenichthys † Extinctions 14 1 3 8 First records 17 38 6 12

Fig. 234. O tolith range chart depicting faunal turn-o ver ratio s acro ss the K-T bo undary extinctio n event and the PETM event in Euro pe (Eocene may be incomplete).

Werner SCHWARZHANS 73

Maastrichtian Danian Selandian Thanetian Ypresian (part) Cretaceous Paleocene Eocene Ostraciidae - Tetraodontiformes Ostracion

Centrolophidae Centrolophus? Scombridae Scomber ? ? Gempylidae Gempylus? ? Leiognathidae Leiognathus? ?

Haemulidae Haemulon?

Sparidae various genera

various genera Serranidae ?

Perciformes Carangidae Caranx? Apogonidae Apogon

Epigonidae Epigonus?

Lactarius Lactariidae Lactarius? ?

Acropomatidae † Plesiopoma †

Scorpaenidae - Scorpaeniformes Scorpaena?

† Zeiformes indet. † Isozen

Melamphaidae Melamphaes? Polymixia Polymixiidae Polymixia? ?

† Traubiella †

Holocentroidei indet. † Sillaginocentrus † † Pfeilichthys † Holocentrus Holocentridae † Holocentronotus †

† Traunichthys † † Berycoidei indet. † Beauryia † Argyroberyx † Antigonia

Beryciformes Antigoniidae Antigonia? † Diretmidae Diretmus Hoplostethus Trachichthyidae Trachichthys

† Kressenbergichthys †

Berycidae Centroberyx

Ogcocephalidae - Lophiiformes Ogcocephalus?

Bythites? Dinematichthys? ? Bythitidae Ogilbia Bidenichthys

† Ampheristus? † † Palaeomorrhua † Gadophycis

† Sirembola Ophidiidae † Praeophidion Hoplobrotula Ophidiiformes † Protobythites †

† Ampheristus

Onuxodon Carapidae † Fierasferoides

Extinctions 14 1 3 8 First records 17 38 0 12 Fig. 234. (co ntinued).

Palaeo Ichthyologica 12 74 shift in berycifo rm o toliths, which are still species rich but O n a larger scale, the otolith record is well enough reduced to rather few genera, is documented by genera established no w to allo w interpretatio n o f the influence of mo stly persisting until today (see chapter 5.2). the PETM event o n teleo st evo lutio n. A total o f 12 genera became extinct during the Paleocene, at least eight of them with the PETM event. This co rrespo nds to an extinction rate of 15 % (Fig. 234), a mo derately elevated but significant ratio . 5.1.3 Influence of the PETM event 39 Paleocene genera can be traced across the PETM event, for teleost evolution and a good proportion of the remaining 26 probably as well, (based on otolith analysis) but have not yet been proven from Eocene strata. O n the species level the turn-o ver fro m Paleo cene to Early Eocene is mo re dramatic, with very few species pro ven befo re and Since the initial reco gnitio n o f the Paleo cene-Eocene Thermal after the PETM. Amo ngst tho se few are Arge ntina e rratica Maximum (PETM) by KENNETT & STOTT (1991) a multi- (RO EDEL 1930) and Preophidion convexus (STINTON 1977). tude o f scientific research wo rk has been perfo rmed and Many other Paleocene species, however, have closely linked published by many earth scientists that has led to a much co unterparts, pro bably descendants, in Early Eocene times. advanced knowledge and understanding of this unique In respect to the Paleo cene o tolith asso ciatio ns o f Bavaria event in the earth’ s yo unger history. The exceptio nal though and Austria these are (Paleo cene first, then Eocene): Genar- short lived heating of the earth atmosphere and the seas tina hauniensis – G. hampshirensis, Anguilla pfeili – A. rec- has led to subtropical temperatures and environments in tangularis, Gnathophis probus – G. schepdaalensis, Arius very high latitudes (BRINKHUIS et al. 2006, SLUIJS et al. subtilis – A. subre ctangularis, Paraulo pus po stangulatus – P. 2009) and effects such as ocean water acidization, deep davisi, Palae o gadus? bratishko i – P.? papillo sus, Ampher- water CaCO disso lutio n and surface water o ligo tro phy 3 istus ne o bavaricus – A. to liapicus, Bide nichthys lapie rre i – are in discussio n (BRALOWER 2002, NG UYEN et al. 2009, B. symme tricus and Centroberyx integer – C. eocenicus. The SLUIJS 2006, UCHIKAWA & ZEEBE 2010, ZEEBE & ZACHO S Paleo cene fauna o f Ukraine (SCHWARZHANS & BRATISHKO 2007). Effects o f the PETM event o n the bio ta has been 2011) adds a few more pairs: Ranice ps he rmani – R. lati- studied fo r planktonic fo raminifera (Q UILLÉVÉRÉ & NO RRIS de ns, Fie rasfe ro ide s buccule ntus – F. subre gularis, Epigonus? 2003, KAIHO et al. 2006, G UASTI & SPEIJER 2007), benthic tyassminensis – E.? selsiensis and Hae mulo n? makare nko i fo raminifera (THO MAS 1998, 2003, 2007; ALEGRET & – H.? sylve stris. A few more are added from the Paleocene O RTIZ 2006; PUJALTE et al. 2003, 2009), nannoplankton of Denmark and Western Greenland: Pro to co llio lus amo r- (AUBRY 1998, BUJAK & BRINKHUIS 1998, TREMO LADA & phus – P. eocenicus and Gadophycis thulei – G. ovalis. BRALOWER 2004, G IBBS et al. 2006), diatoms (O RESHKINA & O BERHÄNSLI 2003), mo llusks (DO CKERY 1998), co ral Thus, the change in the composition of otolith asso ciatio ns reefs (SCHEIBNER & SPEIJER 2008) and terrestrial fauna indicates that the influence o f the PETM event o n teleo st (G ING ERICH 2000, 2003; BLOIS & HADLY 2009; WILF evolution has rather led to an accelerated speciation than & LABANDEIRA 1999) and flo ra (JARAMILLO et al. 2010; to extinctio n. An exceptio n is the apparent extinction of a WING et al. 2003, 2005). The reaction of the biota to the number of beryciform lineages below the genus level which PETM appears to have been diverse, ranging from adaptio n do not seem to have carried over into Eocene times (chapter through migration, temporary adaption by size or chemical 5.2). The interpretation of such an accelerated speciatio n co mpo sitio n, extinctio n o r initiatio n o f evo lutio nary radiatio n. event is also supported by the sudden increase of species The fish fauna during the times o f the PETM is too insuf- and diversificatio n o f mo rpho lo gies in such teleo st groups ficiently kno wn to permit a high reso lutio n study o f the PETM as Co ngro idei, O phidiifo rmes and Percifo rmes (chiefly Per- effects o n the teleo st fauna. An exceptio n is the still po o rly coidei) during early and middle Eocene (Fig. 233; chapters described Mo-clay fish fauna from the Paleocene-Eocene 5.3, 5.4 and 5.5). transition of Denmark (BO NDE 1966, 1979, 2008).

5.2 The Beryciformes: The fate of the survivors from the Cretaceous (deferred extinction, evolutionary stasis leading to ‘living fossils’, adaption to niches)

The Berycifo rmes fo rmed a rich, highly diverse and com- genera o r higher taxa (NO LF & DO CKERY 1990; NO LF mon group of teleosts during the Late Cretaceous. Skeletal & STRING ER 1996; NO LF 2003; SCHWARZHANS 1996, findings have revealed a multitude o f extinct berycifo rm 2010a). These have been interpreted as mostly represent- genera and families (PATTERSO N 1964, 1993). Many of ing extinct berycifo rm gro ups o f beryco id and ho lo centro id these extinct berycifo rms o f the Late Cretaceo us were highly affinities by SCHWARZHANS (2010a), but many similar adapted to specific enviro nments, similar to tho se occupied morphologies have been associated with perciforms, mo stly by percifo rms after the K-T bo undary. A go o d summary of perco ids by NO LF (in NO LF & STRING ER 1996). Except fo ssil berycifo rm taxa is listed in KOTLYAR (1996). Nothing for the slightly upward bent caudal tip found in many is kno wn o f the o toliths o f tho se specialized extinct skeleton- Berycoidei and Stephanoberycoidei there is no apomo rphic based berycifo rms (as there are no kno wn o toliths ‘ in situ’ character distinguishing berycifo rm fro m percifo rm o toliths. fro m Cretaceo us times anyway). Nevertheless, several Particularly ho lo centro id and po lymixio id o toliths are quite apparent acanthomorph otolith morphologies are known similar to many percifo rm o toliths because o f the down- fro m the Late Cretaceo us which o bvio usly represent extinct turned caudal tip (see chapter 5.5 fo r further discussio n)

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Fig. 235. Phylo gram o f the shallo w water Berycifo rmes fo r Late Cretaceous and Paleogene times in Europe depicting the effects o f the K-T bo undary extinctio n event and the seco ndary blo om of shallow water Berycoidei during Paleocene. Shaded areas meant to highlight specific evo lutio nary events. – This and the fo llo wing phylo grams are all exclusively based o n o toliths and are not intended to depict a cladistic analysis no r exact timing o f phylo genetic dichotomies.

Palaeo Ichthyologica 12 76 again demonstrating the importance of direct compariso n the Paleo cene, but mo stly with species o f living genera such o f fo ssil o toliths with recent material and at the same time as Centroberyx, Ho plo ste thus, Trachichthys and a ‘dwarfed’ highlighting the limitatio ns o f o tolith interpretation of pre- Dire tmus. These are represented by 10 different species Tertiary mo rpho lo gies. during Paleocene, while they become reduced to only two in Eocene. The latest record of Centroberyx in Europe is Since percifo rms are widely held to have derived from bery- C. manens NO LF & BRZO BO HATY 2004 fro m the Middle cifo rms, it has been discussed in the literature whether the Miocene. In the Recent, the genus Centroberyx contains 7 Percifo rmes may no t be o f po lyphyletic o rigin in which ‘the Indo-Pacific species, four of which are endemic to Australia ancestors o f the Percifo rmes must have passed at o ne time and Trachichthys with a single species prevalent to Australia. through a stage which would be classified as berycifo rm’ Amongst the new records from the Paleocene are a few (PATTERSO N 1964: 466). The knowledge of acanthomorph early deep water beryciforms – a ‘dwarfed’ Dire tmus and skeleton findings acro ss the Late Cretaceo us /Paleocene a very plesiomorphic melamphaid – and two species o f interval has increased since (ARRATIA et al. 2004), but the Zeifo rmes, which have derived fro m Berycifo rmes and still the Late Cretaceous has remained the domain o f the which stem from a lineage already known since the Late Berycifo rmes whereas the Percifo rmes o nly became the Cretaceous. The melamphaid and zeiform representatives dominant element of acanthomorph fishes during Early exhibit very plesio mo rphic o tolith mo rpho lo gies truly rep- Tertiary. O toliths have now become much better known from resenting the evo lutio nary catego ry 4. the critical interval aro und the K-T bo undary, but as stated befo re sho uld be used carefully fo r testing o f the hypo thesis Conclusion. Paleo cene berycifo rm o tolith mo rpho lo gies fo rmulated by PATTERSO N (1964, 1993). co ntain three different gro ups (Fig. 235). My interpretatio n o f o toliths presented here adheres to the 1. A gro up o f deferred extinct genera which finally become view that the specialized Late Cretaceous acanthomo rph extinct during Paleocene or Eocene (Ho lo ce ntro no tus, mo rpho lo gies represent berycifo rms (Figs. 233-235; for a Kressenbergichthys and an unspecified po lymixiid). detailed discussio n see SCHWARZHANS, 2010a). At least 2. A group of living genera whose otoliths have remained seven such suppo sed extinct berycifo rm o tolith-based genera mo rpho lo gically stable, essentially in an evo lutio nary are o bserved in the Maastrichtian o f Bavaria (Fig. 234, 235) stasis since the Paleo cene o r Late Cretaceo us – Centro- and more may be contained when considering NO LF’s beryx, Hoplosthetus, Trachichthys, Diretmus. Some of reco rds fro m U.S.A. after revisio n. Two o f the fo ssil genera these genera become less and less species rich over time, are persisting into Paleo cene – Ho lo ce ntro no tus and two but o thers like Ho plo sthe tus have adapted to oceanic unspecified po lymixiid species (Polymixia ? harde ri and enviro nments and are very species-rich today. P.? groenlandica) previo usly tho ught to represent veliferids 3. A gro up o f plesio mo rphic berycifo rms and berycifo rm (SCHWARZHANS 2003, 2004). Ano ther extinct beryco id derivatives which have adapted to specific niches where reco rd so far exclusively kno wn fro m the Paleo cene – they thrive until today – the bathypelagic Stephano be- Kressenbergichthys – is likely to stem fro m pre-Tertiary ro o ts. rycoidei and the pseudoceanic Zeiformes. Remarkably, berycifo rm o toliths are still quite co mmon in

5.3 The Anguilliformes and Aulopiformes early radiations

Anguillifo rm and aulo pifo rm o toliths are well kno wn since the recently established family Pro tanguillidae (JO HNSO N Late Cretaceous times and also occur commonly in the et al. 2011), which is supposed to represent a ‘living fos- Early Tertiary. Anguillifo rms are especially species-rich and sil’ eel family. co nsistent, the do minant o tolith-reco rded families being Aulo pifo rm o toliths are also co mmo n in the Late Cretaceous the Congridae. This is in accordance with observations in and Paleocene of Europe until their disappearance from the yo unger sediments thro ugho ut the Tertiary and also in sub- region after the Middle Eocene. The species are not many, recent sea bottom dredges on the shelf (unpublished data). but they represent three majo r evo lutio nary lineages of Co ngridae are represented by several living genera already aulopiforms since the Paleocene with the genera Aulo pus, in the Paleocene and Early Eocene. Clearly, the anguillifo rm Chlo ro phthalmus and Paraulo pus. Paraulo pus is known radiatio n at the family level must have o ccurred deep in since the Late Cretaceous, together with an extinct aulopi- the Cretaceous, while radiation at the genus level appears form – Archaulo pus SCHWARZHANS 2010. O bvio usly, the to be as o ld as Early Tertiary o r at times Late Cretaceous. Aulo pifo rmes are an early evo lved gro up o f teleo sts with Anguillifo rm o toliths are species rich in the Paleo cene of their main branching events during Late Cretaceous. Their Euro pe with at least 10 species. The fo ssil o tolith-based o tolith mo rpho lo gically sho ws a stable pattern ever since genus Bavarico nge r from the Maastrichtian and Paleocene the K-T bo undary (Fig. 236). o f Bavaria is remarkable fo r its suppo sed relatio nship to

Werner SCHWARZHANS 77

Fig. 236. Phylogram of the Aulopidae and Chlorophthalmidae depicting their relative abundance and early evo lutio nary maturatio n during Late Cretaceous and Early Paleogene. – * Record from North America; ** records from New Zealand.

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Fig. 237. Phylo gram o f the O phidiifo rmes during Late Cretaceo us and Paleogene in Europe depicting the wealth of diversity o f extinct Eocene warm shallo w water taxa fo llo wing the PETM event and the turn-over to deep water migration later in Eocene/ O ligocene. Light shaded areas reflect evo lutio nary clusters o f shallo w water o phidiifo rms; dark shaded areas clusters o f deep water o phidiifo rms. – All taxa Euro pean except fo r Xenosirembo and Joe nie lse nia fro m No rth America.

Werner SCHWARZHANS 79

5.4 The rise of the Ophidiiformes (with an excursion to the Gadiformes)

O toliths o f the O phidiifo rmes fo rm a do minant faunal when describing the Maastrichtian o toliths (SCHWARZHANS element in Eocene sediments of the warm shallow seas of 2010a), clearly assignable species were identified o f Euro pe and mo st likely o ther regio ns as well. They are not the Neo bythitinae (O phidiidae) and the Bro smo phycini o nly abundant but also very species rich during that time (Bythitidae). The Paleo cene then yielded further well defined period with about 18 species of the families Carapidae, groups associated with the Carapidae, several groups of O phidiidae (the mo st co mmo n family) and Bythitidae the Neobythitinae such as Ho plo bro tula and Sirembini during their peak times in the Middle Eocene of Europe (sensu SCHWARZHANS 1981a) and three gro ups o f the (Fig. 233; NO LF 1980, 1988; SCHWARZHANS 1981a). Bythitidae. This already very diverse o phidiifo rm fauna, Judging fro m their o tolith size they must stem fro m mo stly however, was not very abundant at any of the Paleocene small fishes, smaller than the average living o phidiifo rms. lo catio ns studied so far. Also the species richness is quite They also represent many extinct genera, which are o ften variable, ranging from only one neobythitine and one witho ut clear relatio n to any o f the living o nes. Nowadays, bythitid in the Selandian o f Denmark to seven o phidiifo rm shallo w water o phidiifo rms are rare, mo stly co nfined to the species in Luzanivka, Ukraine. The fauna from Luzanivka tribe O phidiini on sandy bottom and the Dinemytichthyini is remarkable in that it sho ws the clo sest relatio nship to o f the Bythitidae in reef and ro cky sho re enviro nments. the subsequent Eocene shallow water faunas of Europe. Representatives o f these gro ups are present since the Early Then, after the PETM event, o phidiifo rm fishes start to really Tertiary too , but are mo stly rare. So me extinct mo rphologies flo urish in Euro pe with many mo re lineages. At least 15 relate to the rare shallow water genera Sire mbo , Petro tyx and species are recorded from the Ypresian and 18 from the Spo tto bro tula o f the o phidiid subfamily Neo bythitinae, which Lutetian (Figs. 233, 237). It seems that the O phidiifo rmes no wadays is mo stly living in bathyal enviro nments. Several were amo ngst the teleo sts benefiting mo st fro m the PETM species of the oceanic neobythitine genus Ho plo bro tula are event. Thereafter, during Bartonian they start to fade off recorded from near shore environments during Eocene as and in parallel with gradual cooling during O ligocene lose well as numerous probably related extinct genera. O ne all their abundance and richness of species. At the same of them – Amphe ristus – is amo ngst the few with o toliths time more deep marine ophidiiforms are picking up in reco rded ‘ in situ’ . abundance (NO LF & STEURBAUT 1987, 2004).

All in all, the O phidiifo rmes represented a highly diverse The suppo sed sister gro up o f the O phidiifo rmes within the gro up o f fishes in the shallo w warm seas during the Early Paracanthopterygii, the Gadiformes, have a much more Tertiary, unlike their mo stly bathyal preferences today, and o bscure fo ssil history during the Early Tertiary. They have they contained an abundance of genuinely extinct higher not yet been identified in the Late Cretaceous, neither by taxa during that perio d altho ugh so me persistent o nes o toliths no r by skeletons. Their first reco rd is suddenly, spe- were already present as well. O phidiifo rm fishes fo r a cies rich and with many diverse mo rpho lo gies well attribut- sho rt perio d o f the teleo st evo lutio n during Eocene must able to living families and even genera at times during the have been competitors to the then evolving perciforms (o r Selandian o f Denmark (SCHWARZHANS 2003). O bvio usly, so me gro ups o f them) and po ssibly also the gadifo rms of they represented a group of fishes already then adapted the temperate seas. to temperate seas. This may explain their lack in the sediments o f the tro pical to subtro pical Euro pean seas fro m For some unknown reason the extraordinary wealth of Early the Late Cretaceous as well as the Eocene. Their diversity Tertiary (and Late Cretaceo us) o phidiifo rm o tolith-based spe- in the Paleocene, however, is a clear indication that main cies is no t mirro red by skeleton findings (SCHWARZHANS radiatio ns at the higher hierarchies o f gadifo rms must 2010a). Altho ugh it is difficult to exactly po int o ut what have o ccurred in pre-Tertiary times. In Euro pe, they o nly defines o phidiifo rm o toliths it is nevertheless well accepted become a common, in fact often the dominating group amo ngst o tolith research wo rkers that they are quite un- in the North Sea Basin since the O ligocene, representing mistakable in mo st instances and thus their identificatio n a clear faunal response to the global cooling of the seas is widely co nsidered valid. and the southward shift of the temperate climate zo ne of The main radiatio n at family and subfamily level must have the northern hemisphere. already occurred in the Late Cretaceous. As I pointed out

5.5 The early radiation of the Perciformes/ Percoidei

The Percifo rmes are the largest and mo st diverse group of Cretaceo us fish skeletal finds are almo st entirely devoid of living marine teleo sts. Mo st o f their subo rders and mo st percifo rms. Berycifo rms then represented the main acan- families o f the co re subo rder Perco idei are well established thomorph group (PATTERSO N 1964, 1993) (see chapter since Eocene times. The uniquely rich and well preserved 5.2). PATTERSO N (1993) no ted, ho wever, that there is a fauna from the Middle Eocene of Monte Bolca (Italy) has gap of fish skeletal reports of about 20 Ma between the Late yielded mo re than 160 species o f teleo st fish including many Campanian and the Late Paleocene. ARRATIA et al. (2004) first reco rds, particularly fro m percifo rms (BLOT 1980, PAT- list o nly a few articulated percifo rm reco rds fro m the Late TERSO N 1993). Monte Bolca already displays a ‘modern’ Cretaceo us/ Early Paleo cene strata wo rld wide. faunal composition of acanthomorphs. In contrast, Late

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Fig. 238. Phylo gram o f the Perco idei during Late Cretaceo us and Paleogene in Europe depicting the two phases of rapid radiatio n during Paleo cene fo llo wing the K-T bo undary extinction event and Early to Middle Eocene following the PETM event. Dark shaded areas indicate Paleo cene percifo rm blo sso ming; light shaded areas Eocene perciform blossoming.

Werner SCHWARZHANS 81

O toliths have no w yielded co mplementary material o f for instance the genus Plesiopoma SCHWARZHANS 2010, acantho mo rph fishes fro m this critical time interval fro m which is also described here from the Paleocene. the Campanian to Maastrichtian (NO LF & DO CKERY 1990, NO LF & STRING ER 1996, SCHWARZHANS 2010a) and In co ntrast to the pro blematic acantho mo rph o tolith mor- the Paleocene (NO LF 1978; NO LF & DO CKERY 1993; phologies from the Late Cretaceous, many of those from the SCHWARZHANS 2003, 2004; SCHWARZHANS & BRA- Paleo cene are clearly attributable to percifo rms, and exhibit TISHKO 2011). The classificatio n o f these finds, ho wever, so me wealth in diversificatio n. The o utstanding message is hampered by the fact that o tolith allo catio n depends tho ugh o f the percifo rm o toliths fro m the Paleo cene is that largely o n co rrelatio n with recent o toliths and that there they are predo minantly small / stemming fro m small fish, have been many teleost groups during the Late Cretaceous and that they are very plesio mo rphic in mo rpho lo gy, mo stly which became extinct at the K-T boundary and of which no t allo wing a detailed taxo no mic allo catio n. This o bserva- otoliths have remained completely unknown. In acantho- tio n was first made and co mmented by SCHWARZHANS mo rphs such extinct gro ups are mo stly o f the Beryciformes & BRATISHKO (2011): ‘ Such findings can be characteristic (chapter 5.2). fo r an early evo lutio nary phase, when early plesio morphic There is no single character or character combination blueprints have evolved from which the subsequent perci- that wo uld distinguish percifo rm fro m berycifo rm o to liths, fo rm radiatio n derived’ (ALFARO et al. 2009; FRIEDMAN altho ugh several autapo mo rphies exist in certain ‘ sub- 2009, 2010). And: ‘It may also serve as an indication that gro ups’ o f the two o rders that allo w co nfident allo cation of not much wealth of fossil Percoidei can be expected prio r to fo ssil o toliths yo unger than the Eocene and in fact mo stly the Tertiary’ . Ho wever with the level o f perco id diversificatio n also in the Paleocene. To mention but a few complexities in o bserved in the Paleo cene it appears likely that a number of berycifo rm/ percifo rm o tolith mo rpho lo gies: 1. mo st percoid percoid lineages have originated in Late Cretaceous times o toliths are distinctly hetero sulco id (SCHWARZHANS 1978) as indicated by the genus Plesiopoma and finds from the meaning that the ostium is strongly widened and the caudal Late Cretaceo us o f No rth America no twithstanding certain tip markedly bent do wnwards, but this is also the case in o ther allo catio ns by NO LF & STRING ER (1996), which I ho lo centrids and po lymixiids; 2. mo st beryco id o toliths regard as doubtful. have a wide o stium co mbined with a straight o r slightly upward turning cauda, but similar morphologies are also The wealth of Paleocene perciforms (as evidenced by o toliths) fo und in a few percifo rm fishes like apo go nids. It has been is thus pro bably an expressio n o f rapidly evo lving percifo rm discussed since PATTERSO N (1964) whether the Perciformes fishes o ccupying space left fo r re-co lo nizatio n fro m teleo sts might be o f po lyphyletic o rigin fro m a berycifo rm root, but that became extinct during the K-T bo undary extinction event at this stage o tolith analysis can no t shed additio nal light (FRIEDMAN 2010). The PETM event appears to have been into this hypo thesis (SCHWARZHANS 2010a). the next majo r event favo ring further radiatio n and evolu- With all the abo ve in mind it is no t straight fo rward tion of the already expanding perciform group leading to to taxo no mically allo cate o toliths o f Late Cretaceo us a ‘modern’ acanthomorph fauna in Middle Eocene times. acanthomorphs. NO LF & DO CKERY (1990) and NO LF & This two phase radiatio n is sho wn in figure 238 fro m the STRING ER (1996) have taken the po sitio n to place many Late o tolith perspective. Cretaceo us o tolith mo rpho lo gies in so me extant percifo rm The fo llo wing families o f the Percifo rmes have been families o r subo rders thereby seemingly extending the fos- identified by o toliths fro m the Paleo cene: Perco idei: Acro - sil perciform record way beyond the knowledge provided pomatidae (Plesiopoma from Campanian to Thanetian, by skeletal finds. PATTERSO N (1993) co mpared the fo ssil Acropoma? since Danian), Lactariidae (since Danian), Ep- skeleton and o tolith evidence fro m the Late Cretaceous and igonidae (since Selandian), Carangidae (since Selandian), Paleocene concluding that in his opinion ‘the only o utstand- Serranidae (since Danian), Sparidae (since Danian), Hae- ing difference is in Cretaceous percoids, where there are no mulidae (since Danian), Leiognathidae (since Selandian); skeletal reco rds and rather diverse o tolith reco rds’. In 2010a, Scombroidei: Gempylidae (since Danian), Scombridae I questio ned the unscrupulo us asso ciatio n o f Late Cretaceous (since Selandian); Stro mateidei: Centro lo phidae (since o tolith mo rpho lo gies to percifo rm taxa by previo us wo rks, Selandian), Stro mateidae (since Thanetian). Sparidae and arguing that many of those supposed perciform otolith Haemulidae o ccur with several lineages in parallel. Cer- mo rpho lo gies sho w o nly superficial resemblance to modern tain higher percifo rm subo rders and percifo rm derivatives perciforms and could well be understood as representing (as well as many mo re perco id families) o ccur first during extinct berycifo rm gro ups. SCHWARZHANS (2010a) left o nly Eocene: Trachino idei (since Lutetian), G o bio idei (since late few o tolith finds tentatively with presumed basal percifo rms, Ypresian), Pleuro nectifo rmes (since late Ypresian).

5.6 The Stomiiformes, Myctophiformes, Macrouridae: Early records of a bathyal fauna

The deep sea is the wo rld’ s largest habitat and it is populated or, in high latitude deep water, the Zoarcidae and finally by teleo st fish with a wide variety o f specializations. The most less co mmo n the vario us primary meso - to bathypelagic common ones are of meso- to bathypelagic groups feeding fish predators. The fo ssil reco rd o f fish fro m this largest on zooplankton and often undertaking daily vertical migra- habitat is inadequate in co mpariso n to its size and to the tio n, fo r instance the G o no stomatoidei o r the Mycto phidae. kno wledge o f shallo w water fo ssil habitats. O ne o f the best Another important group are benthopelagic bottom feed- bathyal skeletal fish faunas kno wn fro m the Paleo gene is ers such as Macrouridae and deep water O phidiiformes from the O ligocene Maikop Formation of southern Russia

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Fig. 239. Phylo gram o f the Myctophifo rmes since Late Cretaceo us depicting early ‘ primitve’ myctophifo rms beco ming extinct at the PETM event and the rise o f myctophid diversity fo llo wing the PETM event and subsequently the glo bal co lling event during late Eocene and Early O ligo cene. Shaded areas reflect three distinct lineages within the genus Diaphus. – * Reco rd fro m no rthwestern G reenland; ** reco rd fro m So uth Australia.

and Ukraine and the time equivalent Mennelite Formatio n STEURBAUT 1984). These o tolith asso ciatio ns o btained from o f Poland (JERZMANSKA 1968, PRO KOFIEV 2006). These the Aquitaine Basin in France, Piemo nt in Italy and Eger in faunas are already relatively mo dern co ntaining several Hungary are all rich in go no stomatoids, myctophids and stomiifo rm and myctophid fishes. macrourids, and in addition contain less common deep water o phidiifo rms, sco pelarchids, diretmids and melamphaids. The reco rd o f o toliths fro m o ceanic fishes is fairly well estab- In their entire co mpo sitio n they are a clear fo reshadowing lished fo r the po st-O ligo cene times (NO LF & BRZO BO HATY of the deep sea teleost fauna of the Recent. Earlier data 1994, 2002, 2004; NO LF & STEURBAUT 1987, 2004; than from the O ligocene are few: from the Eocene of the

Werner SCHWARZHANS 83

Aquitaine Basin (NO LF 1988) and the fauna described here genera o f uncertain affinities – Bavarisco pe lus in Late from the Paleocene of Kroisbach, plus a few more erratic Cretaceous and Paleocene times and Danoscopelus fro m data po ints fro m the Eocene o f Australia (SCHWARZHANS the Paleocene. Well defined myctophid otoliths occur first 1985a) and New Zealand (SCHWARZHANS 1980). during Late Paleo cene with the fo ssil o tolith-based genus The Paleo cene o f Kro isbach co ntains the earliest o tolith Eokre fftia and in the Eocene with several species placed in association known to date with deep water fishes chiefly Diaphus (o r the fo ssil skeletal-based genus Eomyctophum). composed of gonostomatoid components. O ther than the From the O ligocene onwards, a diverse myctophid oto lith so mewhat enigmatic o toliths o f Bavarisco pe lus and Dano- mo rpho lo gy is established similar to the o ne o bserved in scopelus (and so me similarly difficult to interpret presumed the Recent (Fig. 239). [NO LF & BRZO BO HATY (1996) repo rt ‘ primitive’ myctophifo rm skeletal finds; PRO KOFIEV 2006), a myctophid turn-over also from the O ligocene-Miocene distinct myctophid o toliths are first reco rded fro m the boundary.] The Macrouridae finally are found with ‘ mod- Late Paleo cene o f So uth Australia – Eokre fftia pre diaphus ern’ o tolith mo rpho lo gies since the Paleo cene, without any SCHWARZHANS 1985 – and thereafter from the Eocene known predecessors, in environments of the temperate and o f the Aquitaine Basin and So uth Australia asso ciated with rather shallo w No rth Sea Basin. O nly during Eocene (So uth the genus Diaphus. The sister-family Neo sco pelidae is also Australia) and O ligo cene (Italy) are macro urid o toliths fo und recorded since Paleocene – Neoscopelus nuussuaqensis in pelagic sediments. SCHWARZHANS 2004. Figure 239 summarizes the early MILLER et al. (1992) described three main turno vers in Paleogene and Maastrichtian finds of the Neoscopelidae, the composition of deep-sea benthic Foraminifera during Myctophidae and the extinct genera of uncertain relatio n- the Cenozoic. After deep water benthic foraminifera had ship. Three presumably apomorhic characters define the survived the K-T bo undary event with no majo r impact, the o toliths o f the Myctophidae and separates them fro m the first turno ver o ccurred at the PETM event, pro bably as a two other groups: the shallow caudal colliculum, the forma- result o f warm water influx into the deep sea (THO MAS tion of a caudal pseudocolliculum and the shortening of 2006). Re-colonization of the deep sea occurred during the cauda being always shorter than the ostium. Early Eocene with migratio n o f certain neritic benthic fo- Deep water macrourids are reported from the Eocene raminifera into the deep ocean (MILLER et al. 1992). Could o f So uth Australia (SCHWARZHANS 1985a), which already it be po ssible that the Macro uridae too migrated to the bear much resemblance to their extant counterparts, as do deep water during that period as a reaction to the shrinking the many macrourid species recorded from the bathyal sedi- neritic temperate realms and the warming up of the deep ments o f the Early O ligo cene o f Italy (NO LF & STEURBAUT water in the oceans? In this respect it may be mentioned 1987, 2004). In fact also the macrourid otoliths described again that modern myctophids seem to have raised during fro m the Selandian o f Denmark (SCHWARZHANS 2003) Eocene and deep water ophidiiformes are first recorded in sho w a surprisingly clo se resemblance to tho se o f living taxa. abundance from the Early O ligocene. These o toliths ho wever o riginate fro m sediment pro bably Much is still to be learned about timing and mechanism not deposited at greater depth than 100 m and at so me of the colonization of the deep sea by teleosts. In respect to distance from deep water. the three dominant groups discussed here it appears likely that the Gonostomatoidei were already deep water fish The still sparse data o f o toliths fro m deep water fishes prior to the K-T boundary. The ‘modern’ Myctophidae likely of Paleogene times suggest that the Gonostomatoidei adapted to their meso- bathypelagic life during Paleogene fo rmed a well established bathyal gro up already in Pale- and the Macrouridae might have migrated into the deep o cene, and pro bably ro o ted well into pre-Tertiary times. sea during the Paleogene, both possibly after the PETM The Myctophifo rmes are represented by two o tolith-based event.

6. Acknowledgments

The o toliths described herein were co llected by Friedrich Pfeil suppo rted in geo lo gical and stratigraphical info rmatio n relevant (München) in the years 1981 to 1982 with support by Bernhard to the lo calities. Additio nal material was co llected in earlier years Beaury (München) and Jürgen Pollerspöck (München), to whom I by the late Franz Traub (München) and was kindly made available wo uld like to sincerely thank fo r making this diverse and rich col- through the help of Pieter A. M. Gaemers (Leiden). Trevo r H. Wo rthy lectio n available fo r studying and descriptio n. Friedrich Pfeil further (Sydney, Australia) is thanked fo r impro ving the English.

Palaeo Ichthyologica 12 84

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(1978): O tolith-mo rpho lo gy and its usage Belgium and northwestern France. – Bulletin de la Société fo r higher systematical units, with special reference to the belge de Géologie, 97: 321-347. Myctophiformes s.l. – Mededelingen van de Werkgroup STINTON, F. C. (1965): Teleo st o toliths fro m the Lo wer London vo o r Tertiaire en Kwartaire G eo lo gie, 15: 167-185. Tertiaries. – Senckenbergiana lethaea, 46a: 389-425. ——— (1980): Die tertiäre Teleo steer-Fauna Neuseelands, reko n- ——— (1966): Fish o toliths fro m the Londo n Clay. – Pp. 404-478 struiert anhand vo n O tolithen. – Berliner geo wissenschaftli- in: CASIER, E. Faune ichthyologique du London Clay; British che Abhandlungen, A, 26: 1-211. [English translatio n 1984 Museum (Naturural History). in: Report New Zealand Geological Survey, 113: 1-269.] ——— (1975): Fish o toliths fro m the English Eocene. 1. – Palae- ——— (1981a): Vergleichende morphologische Untersuchungen ontographical Society Monographs, 544: 1-56. an rezenten und fossilen O tolithen der O rdnung O phidii- ——— (1977): Fish o toliths fro m the English Eocene. 2. – Palae- fo rmes. – Berliner geo wissenschaftliche Abhandlungen, A, ontographical Society Monographs, 548: 57-126. 32: 63-122. ——— (1978): Fish o toliths fro m the English Eocene. 3. – Palae- ——— (1981b): Die Entwicklung der Familie Ptero thrissidae ontographical Society Monographs, 555: 127-189. (Elo po mo rpha, Pisces) reko nstruiert nach O tolithen. – ——— (1980): Fish o toliths fro m the English Eocene. 4. – Palae- Senckenbergiana lethaea, 62: 77-91; Frankfurt/ M. ontographical Society Monographs, 558: 191-258. ——— (1985a): Tertiäre O tolithen aus So uth Australia und Victoria ——— (1984): Fish o toliths fro m the English Eocene. 5. – Palae- (Australien). – Palaeo Ichthyo lo gica, 3, 60 pp; München. ontographical Society Monographs, 565: 259-320. ——— (1985b): Fish o toliths fro m the lo wer Tertiary o f Ellesmere STINTON, F. C. & D. NO LF (1970): A teleo st o tolith fauna from the Island. – Can. Journ. Earth. Sci., 23: 787-793. sands of Lede, Belgium. – Bulletin de Societé belge de Gèol- ——— (1996): O toliths fro m the Maastrichtian o f Bavaria and o gie, Palèo ntolo gie et Hydro lo gie, 78: 219-234; Bruxelles. their evo lutio nary significance. – Pp. 417-431 in ARRATIA, STOREY, M., R. A. DUNCAN & C. C. SWISHER (2007): Paleocene- G. & G. VIO HL (eds.). Meso zo ic Fishes – Systematics and Eocene Thermal Maximum and the O pening of the North- Paleo eco lo gy; München (Pfeil). east Atlantic. – Science, 316: 587-589. ——— (2003): Fish o toliths fro m the Paleo cene o f Denmark. – THO MAS, E. (1998): Bio geo graphy o f the late Paleo cene benthic G eo lo gical Survey o f Denmark and G reenland Bulletin, fo raminiferal extinctio n. – Pp. 214-243 in: AUBRY, LUCAS 2: 1-94. & BERGG REN (eds.). Late Paleo cene – Early Eocene climatic ——— (2004): Fish o toliths fro m the Paleo cene (Selandian) of and bio tic events in the marine and terrestrial reco rds. West Greenland. – Meddelser om Grønland, 42: 1-32. Co lumbia University Press. ——— (2007): The o toliths fro m the Middle Eocene o f O steroden ——— (2003): Extinctio n and fo o d at the seaflo o r: A high- near Bramsche, north-western Germany. – Neues Jahrbuch reso lutio n benthic fo raminiferal reco rd acro ss the Initial für Geologie und Paläontologie, Abhandlungen, 244: Eocene Thermal Maximum, Southern O cean Site 690. – Pp. 299-369. 319-332 in: WING, G ING ERICH, SCHMITZ & THO MAS ——— (2010a): O tolithen aus den G erhartsreiter Schichten (eds.). Causes and consequences of globally warm climates (O berkreide: Maastricht) des G erhartsreiter G rabens in the Early Paleo gene. The G eo lo gical So ciety o f America, (O berbayern). – Palaeo Ichthyologica, 4: 100 pp. Special Paper 369. ——— (2010b): The otoliths from the Miocene of the North Sea ——— (2007): Cenozoic mass extinctions in the deep sea: What Basin. – 352 pp.; Backhuys Publishers & Margraf Publish- pereturbs the largest habitat on earth? – The Geolo gical ers. 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THO MAS, E., H. BRINKHUIS, M. HUBER & U. RÖ HL (2006): An Anhalt. – Zeitschrift der G eschiebefo rschung, 2: 172- ocean view of the early Cenozoic greenhouse world. – 187; Berlin. O ceanography, 19: 94-103. WEILER, W. (1942): Die O tolithen des rheinischen und nordwest- TORSVIK, T. H., D. CARLOS, J. MO SAR, L. R. M. CO CKS & T. N. deutschen Tertiärs. – Abhandlungen des Reichsamts für MALME (2000): G lo bal reco nstructio ns and No rth Atlantic Bodenforschung, Neue Folge, 206: 1-140. paleogeography 440 Ma to Recent. – Pp. 18-39 in: EIDE, WEITZMAN, S. H. (1974): O steo lo gy and evo lutio nary relatio n- E. A. (co o rd.). BATLAS: Mid No rway plate reco nstructio ns ships o f the Sterno ptychidae, with a new classification of atlas with glo bal and Atlantic perspectives: G eo lo gical stomiatoid families. – Bulletin o f the American Museum Survey o f No rway. o f Natural History, 153: 327-478. TRAUB, F. (1979): Weitere Paleo zän-G astro po den aus dem Helveti- WILF, P. & C. C. LABANDEIRA (1999): Respo nse o f Plant-Insect kum des Haunsberges nördlich von Salzburg. – Mitt. Bayer. Associations to Paleocene-Eocene Warming. – Science, Staatsslg. Paläo nt. hist. G eo l., 19: 93-123; Taf. 12-18. 284: 2153-2156. TREMO LADA, F. & T. J. BRALOWER (2004): Nanno fo ssil assem- WING, S. L., G. J. HARRINGTON, G. J. BO WEN & P. L. KOCH blage fluctuations during the Paleocene-Eocene Thermal (2003): Floral change during the Initial Eocene Thermal Maximum at Sites 213 (Indian O cean) and 401 (North Maximum in the Powder River Basin, Wyoming. – Geolo gi- Atlantic O cean): palaeoceanographic implications. – Ma- cal So ciety o f America, Special Paper, 369: 425-440. rine Micro paleo ntolo gy, 52: 107-116. WING, S. L., HARRINGTON, G. J., F. A. SMITH, J. I. BLOCH, D. M. TYLER, J. C. & F. SANTINI (2005): A phylo geny o f the fo ssil and BO YER & K. H. FREEMAN (2005): Transient Flo ral Change extant zeifo rm-like fishes. Upper Cretaceo us to Recent, with and Rapid Global Warming at the Paleocene-Eocene comments on the putative zeomorph clade (Acanthomor- Boundary. – Science, 310: 993-996. pha). – Zoologica Scripta, 34: 157-175. ZEEBE, R. E. & J. C. ZACHO S (2007): Reversed deep-sea car- UCHIKAWA, J. & R. E. ZEEBE (2010): Examining po ssible effects o f bonate ion basin gradient during Paleocene-Eocene seawater pH decline o n fo raminiferal stable iso topes during thermal maximum. – Paleoceanography, 22, PA3201, the Paleocene/ Eocene Thermal Maximum. – Paleoceano- doi:10.1029/2006PA001395. graphy, 25, PA2216, doi:10.1029/2009PA001864. ZIEGLER, P. (1990): G eo lo gical Atlas o f Western and Central VO IGT, E. (1926): Über ein bemerkenswertes Vorko mmen neuer Euro pe. – Shell Internatio nale Petro leum Maatschappij Fischotolithen in einem Senongeschiebe von Cöthen in B.V.: 239 pp.

8. Addendum

Otolith nomenclature

Ernst Koken (1860-1912) was the pioneer who established cases, where an otolith-based species was assigned to a scientific work based on otolith morphology and he was the genus and it was only kept for those instances where this first to describe o tolith-based fo ssil species in 1884. With was no t po ssible. WEILER (1968) stated that the familial this, he established a no menclatural system in which he name in genitive plural, no t in italics, was o nly meant to placed all fo ssil o tolith-based species in a co llective genus represent a taxonomic indication of the author’s systematic O to lithus followed by the name of the genus, respectively assessment without nomenclatorial meaning and hence was family in genitive, respectively genitive plural, in brackets put between angular brackets, in this example: O to lithus fo llo wed by the species name. His ratio nal was that fo ssil [G adidarum] elegans KOKEN 1884. Nevertheless, WEILER o toliths as iso lated remains o f fishes wo uld o nly rarely be (1968) was of the opinion that the insertion would pro tect assignable to a fish thro ugh finds o f o toliths in situ and against ho mo nymy fo r instance o f O to lithus [Gadidarum] that Recent o toliths were po o rly kno wn so that in many elegans KOKEN 1884, O to lithus [Sparidarum] elegans instances only placement into a family would be possible PRO CHAZKA 1893 and O to lithus [O phidiidarum] elegans fo r tho se fo ssil finds (KOKEN, 1884, p. 502). For instance FRO ST 1934, in co ntrary to the view o f ZILCH (1965). a species that he would feel comfortable in assigning to the Ho wever, this practice is no t co mpliant with article 6.1 of extant genus Gadus LINNAEUS 1758 he would describe as the ICZN, which only allows subgeneric names in brackets O to lithus (G adi) tube rculo sus KOKEN 1884, and a species between genus and species names. he would only be able to associate with the family Gadidae Also the collective genus name O to lithus KOKEN 1884 as O tolithus (Gadidarum) elegans KOKEN 1884. The view, could be confused with O to lithe s O KEN 1817, a genus of as expressed by WEILER (1968) was that it co uld at a later the family Sciaenidae, which has been written O to lithus by stage be replaced by a ‘proper’ genus name whenever ad- CUVIER 1829 and authors. Now, while O to lithus CUVIER equate kno wledge o f Recent o toliths o f the family G adidae 1829 represents a junio r o bjective syno nym o f O to lithe s in this example would have become available. And indeed, O KEN 1817 it also represents a senior homonym to G AEMERS (1972) was able to place this very species in the O to lithus KOKEN 1884. Therefo re, HUDDLESTON (1983) genus Triso pte rus RAFINESQ UE 1814, so that it is repo rted introduced a replacement collective group name O to litho p- since as Triso pte rus e le gans (KOKEN 1884). sis for all those otolith-based taxa of unknown generic and The general view o f o tolith researchers was that KO- familial position, which hitherto had been placed in the KEN’ s ‘ inventio n’ was a fine metho d to reflect exactly and ‘ family’ incertae sedis. adequately the pertinent level o f kno wledge. PO STHUMUS The entire o riginal no menclatural practice o f KOKEN (1924) was the first to mentio n that KOKEN’ s no menclature was finally abando ned so o n after WEILER (1968). G AEMERS was not in compliance with the regulations of the ICZN (1971) and authors dropped the collective group name to then established and therefore the usage of the collective refer to a Gadidarum elegans KOKEN 1884. NO LF (1977), group name O to lithus was so o n after stopped fo r all tho se and explained in more detail in 1985 (page 30), pro posed

Palaeo Ichthyologica 12 88 a co llective gro up system in which the plural genitive names well belong to any other known or unknown genus in the are preceded by ‘genus’ instead of ‘O to lithus’, fo r instance particular family. “genus G adidarum” elegans KOKEN 1884 (to stay with the In respect to my fo rego ing study entitled “The o to liths example), making reference to RICHTER (1948). RICHTER fro m the Paleo cene o f Kressenberg Bavaria and Kro isbach (page 146) indeed discussed the nomenclatural system pre- Austria”, it has beco me o bvio us that it is no t advisable to sented by NO LF, but did not find it optimal, and in respect continue the ‘collective group’ terminology currently in use to o toliths pro po sed the use o f the co llective genus name in otolith research. I have therefore made use of Recent and O to lithus fo r all tho se instances, where a generic affilia- fo ssil (o tolith-based) genera wherever available and appro- tio n is unreso lved. Nevertheless, the no menclatural systems priate and used the methodology proposed by JANSSEN intro duced by G AEMERS and particularly that intro duced by (2012b), wherever maintaining of an unresolved generic NO LF are widely used in contemporaneous otolith research allocation appeared favorable. This nomenclatural system until today. G AEMERS & V. HINSBERGH (1978) stated that also allo ws fo r a simple transfer and safeguarding of pre- they were well aware that this no menclature system “is no t vious species names and authorships recorded under the quite in agreement with the ICZN”. They recommended to previo us no menclatural systems used in o tolith research. The hand in a proposal with the ICZN commission to ask the collective group name O to litho psis HUDDLESTON 1983 is next amendment of the Code “in such a way that the es- available fo r all species o f unkno wn familial relatio nship. tablished practice in o tolith systematics beco mes legitimate”, but they did not take action. The kno wledge o f Recent o toliths has tremendo usly G AEMERS, P. A. M. (1971): Bonefish-o toliths fro m the Anversian increased since the early days o f KOKEN, but o f co urse has (Middle Miocene) of Antwerp. – Leidse Geol. Meded., - not reached a status that could be described as ‘nearing 46: 237 267. ——— (1972): O toliths fro m the type lo cality o f the Sands o f Berg co mpletio n’ . Still in many cases, including the mentioned (Middle O ligocene) at Berg, Belgium. – Meded. Werkgr. family G adidae, o ur kno wledge o f Recent o toliths is well Tert. Kwart. Geol., Leiden, 9: 73-85. adequate to judge, whether a specific fo ssil o tolith-based G AEMERS, P. A. M. & V. HINSBERGH, V. (1978): Rupelian (Middle species represents any o f the extant genera o f the family O ligo cene) fish o toliths fro m the clay pit ‘ De Vlijt’ near or, as the case may be, a fossil otolith-based genus. [O ur Winterswijk, The Netherlands. – Scripta G eo lo gica, Leiden, kno wledge o f in situ o toliths o f fo ssil skeleton-based gen- 46, 77 pp. era is rudimentary at best and it is unlikely that it will ever JANSSEN, A. (2012a): Systematics and bio stratigraphy of ho- become much better due to the different nature of the fo ssil loplanktonic Mollusca from the O ligo-Miocene of the Maltese Archipelago. – Boll. Mus. Regionale Sci. Natur., preservatio n o f o toliths as co mpared to skeletons.] While the Torino , 28: 197-601. increased knowledge base of Recent comparative otoliths ——— (2012b, in press): Validatio n o f ho lo planktonic mo lluscan has lead to a much more reliable recognition of the nature taxa from the O ligo-Miocene of the Maltese Archipelago, and co mpo sitio n o f fo ssil o tolith-based fish faunas, most intro duced in vio latio n with ICZN regulatio ns. – Cainozoic o tolith research wo rkers will agree that there is still a need Research, 9. to maintain a no menclatural system that allo ws describing KOKEN, E. (1884): Über Fisch-O tolithen, insbeso ndere über o f a fo ssil o tolith-based species with unreso lved generic diejenigen der norddeutschen O ligocän-Ablagerungen. – - assignment. Z. dtsch. Geol. Ges., Berlin, 36: 500 565. HUDDLESTON, R. W. (1983): O to litho psis, nom. nov., new name In a recent monograph dealing with mollusks from fo r O to lithus Koken, 1884, non Cuvier, 1829 (Pisces: family the Tertiary o f Malta, JANSSEN (2012a) applied the no- incertae sedis). – Journal o f Paleo ntolo gy, 57: 601-602. menclatural system o f o tolith research fo r the first time fo r NO LF, D. (1977): Les o toliths des téléo stéens de l’ O ligo-Miocène mo lluscan systematics. Sho rtly thereafter, it was brought to Belge. – Ann. Soc. Roy. Zool. Belg., Brussels, 106: his attentio n that the “recently intro duced species applying 3-119. ‘open generic nomenclature’ by using the indication ‘Genus ——— (1985): O tolithi Piscium. – Handbo o k o f Paleo ichthyology, Clionidarum’ instead of a formal genus name are vio lating G ustav Fischer Verlag, 10, 145 pp. ICZN art. 11.9.3’”. In a subsequent paper JANSSEN (2012b) PO STHUMUS, O . (1924): O tolithi Piscium. – Fossilium Catalogus, I: Animalia, W. Junk, Berlin, 24, 42 pp. stated that he followed “the format generally adopted in RICHTER, R. (1948): Einführung in die Zo o lo gische No menklatur o tolith literature fo r taxa that canno t be assigned to known durch Erläuterung der Internationalen Regeln. – Verlag Dr. genera, a system so far never questio ned fo r these fo ssils by Waldemar Kramer, Frankfurt am Main, 252 pp. editorial bo ards and/ o r peer-reviewers o f many prestigio us WEILER, W. (1968): O tolithi Piscium. – Fossilium Catalo gus, I: Ani- perio dicals”. He went o n validating tho se taxa by combin- malia, Dr. W. Junk N. V., ‘s-Gravenhage, 117, 196 pp. ing the new names with an unambiguous genus-name, ZILCH, A. (1965): Die Typen und Typoide des Natur-Museums i.e. the name of the type-genus of the family, followed by Senckenberg, 31: Fossile Fisch-O tolithen. – Senckenber- - a questio n mark, indicating that tho se species might as giana lethaea, 46a: 453 490.

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In der Reihe ISSN 0724-6331 Palaeo ISBN 978-3-89937-155-0 Ichthyologica erscheinen zoologische und paläontologische Arbeiten zur Systematik Morphologie Palökologie Paläogeographie Stratigraphie der Fische.