Zemes un vides zinātnes Earth and Environment Sciences - Latvijas ...

Zemes un vides zinātnes Earth and Environment Sciences - Latvijas ... Zemes un vides zinātnes Earth and Environment Sciences - Latvijas ...

19.11.2013 Views

72 ADVANCES IN PALAEOICHTHYOLOGY All actinopterygians are characterised by the presence of ganoine, all sarcopterygians by the presence of a broad based peg, cosmine and true enamel. The course of the lateral line canals in bone is the common character of all osteichthyans. The sister group relationship of Lophosteus and Climatius is the result of the presence of symmetrical and paired fin spines, and of tooth whorl. The presence of a spiny process on the shoulder girdle appears to be a basal character, lost within actinopterygians more derived than Andreolepis and within sarcopterygians in forms more derived than Psarolepis. This agrees with the analysis of Zhu et al. (1999), who placed special importance on the occurrence of spines and spiny processes in early osteichthyans. Tesserae appear as a unique character of Climatius, because Lophosteus is coded with tesserae or plates. All osteichthyans have a closed skull roof (plates), a feature not known from Lophosteus. Still there are plates larger than what is normally considered as tesserae. Fig. 7. Interrelationship of Lophosteus. Node 1: 8 1 , 9 2-1 , 11 1 ; node 2: 17 1 ; node 3: 5 1 ; node 4: 2 1 , 6 1 , 14 1 [bold = unique character change; for explanations see Appendix 2 and text]. Characters of the ornament-like stellate tubercles (also found in Sigaspis) and ridges (also present in some actinopterygians and sarcopterygians) occur in parallel to Lophosteus in other gnathostomes. Nodules, another form of ornamentation, is known in arthrodires (Gross 1973), acanthodians (Gross 1971b) and Lophosteus (Gross 1969). That such a similar ornament can appear again and again, is very well established.

H.-P. Schultze, T. Marss. Revisiting Lophosteus 73 Conclusions For over 100 years, Lophosteus Pander, 1856 was known only from the upper Pridoli of one locality, the Ohesaare Cliff on Saaremaa, Estonia. Märss (1986) described a second species from the lower Pridoli of a second locality on Saaremaa. About 10 years later, Burrow (1995a) and Märss (1997) extended the range of the genus to the Lochkovian of Australia and the Urals. Since the beginning of this century, the genus has been known from the lower Ludlow to the upper Lochkovian and from Australia to the Canadian Arctic and the Urals. The genus falls within the distribution, in time and paleogeography of the oldest actinopterygian scales (Naxilepis, upper Wenlock – upper Ludlow, China [Wang and Dong 1989]; Andreolepis, middle Ludlow – lower Pridoli, Scandinavia [Gross 1968], Baltic region, southern Britain, Asian part of Russia [Märss 2001] and China [Burrow et al. 2000]; Ligulalepis, upper Ludlow to Emsian, Australia [Schultze 1968, Burrow 1994] and China [Wang and Dong 1989]; Terenolepis, upper Lochkovian, Australia [Burrow 1995b]; Dialipina, Lochkovian – Emsian, Canadian Arctic, Siberia [Schultze 1968, 1977, 1992] and Sichuan, China [Burrow et al. 2000]). Psarolepis (upper Pridoli – upper Lochkovian, China [Zhu and Schultze 1997, Yu 1998]) and the sarcopterygians Onychodus (Pragian, China [Wang 1992], Emsian, China [Burrow et al. 2000]), and Achoania (Zhu et al. 2001), Youngolepis (Chang 1982), and Diabolepis (Chang and Yu 1984) appear later (upper Lochkovian, China) as do Powichthys (Jessen 1975, 1980) and the lungfish Uranolophus (Denison 1968) and Speonesydrion (Campbell and Barwick 1982). The 17 characters, which can be collected from Lophosteus, place the genus outside all osteichthyans, whereas these characters separate actinopterygians and sarcopterygians within osteichthyans. Lophosteus is the sister group of the acanthodians as suggested by Otto (1991). One may place it between acanthodians and osteichthyans as a basal osteichthyan. Acknowledgments. - The senior author thanks Mr. P. Czaja for the SEM pictures (Figs 3 D, E, 4, 5 B, 6), Dr. St. Schultka for photographs (Fig. 1 A-C) and Mrs. E. Siebert for the drawings (Figs 2 and 7), all from the Museum für Naturkunde, Berlin. T.M. thanks all following institutions and colleagues. Drs H. Perens and E. Kala, Geological Survey of Estonia, provided the core samples. The samples from the central Urals were collected by expeditions in the years 1973 to 1986 organized by Drs A. Zhivkovich and P. Chekhovich, Moscow. The 1994 Canadian Arctic expedition, in which TM participated, was financed by the Polar Continental Shelf Project, McGill University Northern Studies, the Natural Science and Engineering Research Council, Canada, the Muséum national d’Histoire naturelle Paris, France, and the UNESCO/IUGS (IGCP Project 328); participants were Drs M.V.H. Wilson, P.-Y. Gagnier, D. Goujet, M. Caldwell, and Mr. A. Lindoe. SEM pictures (Figs 3 A-C, 5 A, C) were taken by V. Mikli in the Centre for Material Research at Tallinn University of Technology. This study was supported by the Estonian Science Foundation grant 5726 for T.M. Dr. D. Unwin, Berlin, kindly corrected the English. References Basden A., Burrow C., Hocking M., Parkes R., Young G. 2000. Siluro-Devonian microvertebrates from southeastern Australia. Courier Forschungsinstitut Senckenberg, 223: 201-222. Burrow C. J. 1994. Form and function in scales of Ligulalepis toombsi Schultze, a palaeoniscoid from the Early Devonian of Australia. Record of the South Australian Museum, 27: 175-185.

H.-P. Schultze, T. Marss. Revisiting Lophosteus<br />

73<br />

Conclusions<br />

For over 100 years, Lophosteus P<strong>and</strong>er, 1856 was known only from the upper Pridoli<br />

of one locality, the Ohesaare Cliff on Saaremaa, Estonia. Märss (1986) described a<br />

second species from the lower Pridoli of a second locality on Saaremaa. About 10<br />

years later, Burrow (1995a) <strong>and</strong> Märss (1997) extended the range of the genus to the<br />

Lochkovian of Australia <strong>and</strong> the Urals. Since the beginning of this century, the genus<br />

has been known from the lower Ludlow to the upper Lochkovian <strong>and</strong> from Australia to<br />

the Canadian Arctic <strong>and</strong> the Urals. The genus falls within the distribution, in time <strong>and</strong><br />

paleogeography of the oldest actinopterygian scales (Naxilepis, upper Wenlock – upper<br />

Ludlow, China [Wang <strong>and</strong> Dong 1989]; Andreolepis, middle Ludlow – lower Pridoli,<br />

Sc<strong>and</strong>inavia [Gross 1968], Baltic region, southern Britain, Asian part of Russia [Märss<br />

2001] <strong>and</strong> China [Burrow et al. 2000]; Ligulalepis, upper Ludlow to Emsian, Australia<br />

[Schultze 1968, Burrow 1994] <strong>and</strong> China [Wang <strong>and</strong> Dong 1989]; Terenolepis, upper<br />

Lochkovian, Australia [Burrow 1995b]; Dialipina, Lochkovian – Emsian, Canadian<br />

Arctic, Siberia [Schultze 1968, 1977, 1992] <strong>and</strong> Sichuan, China [Burrow et al. 2000]).<br />

Psarolepis (upper Pridoli – upper Lochkovian, China [Zhu <strong>and</strong> Schultze 1997, Yu 1998])<br />

<strong>and</strong> the sarcopterygians Onychodus (Pragian, China [Wang 1992], Emsian, China<br />

[Burrow et al. 2000]), <strong>and</strong> Achoania (Zhu et al. 2001), Yo<strong>un</strong>golepis (Chang 1982), <strong>and</strong><br />

Diabolepis (Chang <strong>and</strong> Yu 1984) appear later (upper Lochkovian, China) as do<br />

Powichthys (Jessen 1975, 1980) <strong>and</strong> the l<strong>un</strong>gfish Uranolophus (Denison 1968) <strong>and</strong><br />

Speonesydrion (Campbell <strong>and</strong> Barwick 1982).<br />

The 17 characters, which can be collected from Lophosteus, place the genus outside<br />

all osteichthyans, whereas these characters separate actinopterygians <strong>and</strong> sarcopterygians<br />

within osteichthyans. Lophosteus is the sister group of the acanthodians as suggested<br />

by Otto (1991). One may place it between acanthodians <strong>and</strong> osteichthyans as a basal<br />

osteichthyan.<br />

Acknowledgments. - The senior author thanks Mr. P. Czaja for the SEM pictures (Figs 3 D, E, 4,<br />

5 B, 6), Dr. St. Schultka for photographs (Fig. 1 A-C) <strong>and</strong> Mrs. E. Siebert for the drawings (Figs<br />

2 <strong>and</strong> 7), all from the Museum für Naturk<strong>un</strong>de, Berlin. T.M. thanks all following institutions <strong>and</strong><br />

colleagues. Drs H. Perens <strong>and</strong> E. Kala, Geological Survey of Estonia, provided the core samples.<br />

The samples from the central Urals were collected by expeditions in the years 1973 to 1986<br />

organized by Drs A. Zhivkovich <strong>and</strong> P. Chekhovich, Moscow. The 1994 Canadian Arctic<br />

expedition, in which TM participated, was financed by the Polar Continental Shelf Project,<br />

McGill University Northern Studies, the Natural Science <strong>and</strong> Engineering Research Co<strong>un</strong>cil,<br />

Canada, the Muséum national d’Histoire naturelle Paris, France, <strong>and</strong> the UNESCO/IUGS (IGCP<br />

Project 328); participants were Drs M.V.H. Wilson, P.-Y. Gagnier, D. Goujet, M. Caldwell, <strong>and</strong><br />

Mr. A. Lindoe. SEM pictures (Figs 3 A-C, 5 A, C) were taken by V. Mikli in the Centre for<br />

Material Research at Tallinn University of Technology. This study was supported by the Estonian<br />

Science Fo<strong>un</strong>dation grant 5726 for T.M. Dr. D. Unwin, Berlin, kindly corrected the English.<br />

References<br />

Basden A., Burrow C., Hocking M., Parkes R., Yo<strong>un</strong>g G. 2000. Siluro-Devonian microvertebrates<br />

from southeastern Australia. Courier Forsch<strong>un</strong>gsinstitut Senckenberg, 223: 201-222.<br />

Burrow C. J. 1994. Form <strong>and</strong> f<strong>un</strong>ction in scales of Ligulalepis toombsi Schultze, a palaeoniscoid<br />

from the Early Devonian of Australia. Record of the South Australian Museum, 27: 175-185.

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