Cloutier and Forey 1991 Diversity of Extinct and Living Actinistian Fishes (Sarcopterygii)

Environmental Biology of Fishes 32: 59-74, 1991. 0 1991 Kluwer Academic Publishers. Printed in the Netherlands.

Diversity of extinct and living actinistian fishes (Sarcopterygii)

Richard Cloutierl,* & Peter L. ForeyZ I Museum of Natural History, Department of Systematics and Ecology, The University of Kansas, Lawrence, KS 660452454, U. S. A. 2 Department of Palaeontology, British Museum (Natural History), Cromwell Road, London SW7 5BD, England

Received 15.4.1990 Accepted 11.8.1990

Key words: Coelacanth, Fossil record, Species diversity

Synopsis

A total of 121 actinistian species belonging to 47 genera and 17 undetermined actinistians is reported from the literature. There are 69 valid species with fair assessment of their phylogenetic position; 21 valid species with poor assessment of their phylogenetic position; 31 actinistian incertae sedis; and 18 taxa that had been identified incorrectly as actinistians or are nomen nuda. The fossil record of the actinistians covers a history of approximately 380 million years. The greatest diversity occurred during the Scythian (Early Triassic).

Introduction Actinistian systematic overview

The Actinistia (= Coelacanthi, Coelacanthia, Coelacanthiformes, Coelacanthii, Coelacanthina), represented today by a single living species, Lati- meria chalumnae Smith, has a long fossil record beginning with the Middle Devonian Euporosteus eifelianus (Givetian) and ending with the Late Cre- taceous Macropoma lewesiensis (Santonian). Most of us are familiar with the past 65 million year stratigraphic gap (i.e., lack of fossils). Nevertheless the group had been known from fossils prior to the discovery of L. chalumnae for more than a century. The present contribution provides a list of actinistian taxa with our assessments of synonyms. In presenting this taxonomic list, we hope to provide some basis for assessment of species diversity, and also to give some idea of the basis of knowledge since it remains true that many species are known only by very fragmentary remains.

The systematics of actinistians is poorly resolved. Prior to the discovery of Latimeria chalumnae in 1938, actinistians were thought to belong to a single crossopterygian family - the Coelacanthidae. Some authors (Berg 1940, Vorobyeva & Obruchev 1964, Lehman 1966, Romer 1966, Andrews et al. 1967, Thomson 1969, Moy-Thomas & Miles 1971, Andrews 1973, Carroll 1988) have addressed the classification of actinistians, but these attempts have resulted in the recognition of gradal and monotypic groups (Forey 1981, Cloutier 1990). No generally accepted phylogeny or higher taxonomic scheme is used for the Actinistia. However, the most commonly referred classifications are those of Berg (1940), Vorobyeva & Obruchev (1964), An- drews et al. (1967), Lund & Lund (1985), and Car- roll (1988). Moy-Thomas (1937)) Forey (1981)) and Cloutier (1990) have investigated the systematics of Paleozoic actinistians and provided lists of Paleo- zoic taxa. However, the clarification of interrela-

60

tionships of post-Paleozoic actinistians, which corresponds to most of the specific diversity, has only recently been attempted (Cloutier 1991, Forey 1991).

The compiled tabulation (Table 1) provides a list of all genera, species, and undetermined actinistians mentioned in the literature. This list is supple- mented by geological, stratigraphic, and geograph- ic information for each taxon. The primary author for each taxon is provided and included in the bibliography article (Forey & Cloutier 1991); however, authors of subsequent taxonomic changes are not given in the list but are incorporated in the bibliography article. For each genus, the type species is listed first, followed by other referred species alphabetically and the undetermined species.

Assessments about synonymies of taxa have been followed only when it was justified in the literature or following work in progress by one or other of us. Many actinistian species were described originally under different genera; in order to indicate these nomenclatural changes, we follow the convention adopted in the Handbook of Pale- oichthyology (see e.g., Denison 1978) by refering to the original generic assignment as the first entry of synonymy. Spelling of species names is in agree- ment with the International Code of Zoological Nomenclature (Ride et al. 1985). Among the species that have been recognized herein, it remains likely that some are synonyms or not valid, but further systematic studies are needed and beyond the scope of this paper. For example, there are three species in the genus Rhipis, all of which are Late Cretaceous taxa known solely from scales, and two of them come from the same locality; thus it is possible that these forms represent a single species. For this reason a brief statement of parts of the animal known is given with each entry.

Our tabulation is divided into four categories reflecting our systematic interpretation of these taxa. For each category the genera are listed alphabetically. Category ‘A’ deals with the genera and species that can be placed with some confi- dence within at least one of the cladistic classifications given in Forey (1991) and Cloutier (1991). This category includes species that can be posi- tioned in reference to terminal taxa, nodes, re-

stricted clades, or a restricted range of nodes on the cladograms.

Category ‘B’ lists the genera and species that are considered to be valid species but for which little else may be said about their relationships (i.e., phylogenetic position). This category accounts for the Actinistia incertae sedis. These taxa are impor- tant for estimating total diversity but are of little use when estimating rates of clade evolution (morphological and taxonomic).

Category ‘C’ lists genera and species of those remains that can only be identified as actinistians. Their status as valid species is questioned. Finally, category ‘D’ includes the taxa that are undefined (nomen nuda), or those that have been cited as coelacanths but are referable to other fish groups.

Based on our tabulation, there are 69 valid species with fair assessment of their phylogenetic position (Category A); 21 valid species with poor assessment of their phylogenetic position (Category B); 31 actinistian incertae sedis (Category C); and 18 taxa that had been recognized in the literature as actinistians but are not (Category D). There are 47 recognized genera, 29 of which are monotypic and 17 undetermined actinistians (most of which are partial remains). Thus, there is a total of 90 valid species and 31 incertae sedis actinistians distributed from the Givetian (Middle Devonian) to the Re- cent; this total differs from that of Forey (1984) who reported approximately 70 species belonging to 28 actinistian genera.

Fossil actinistians have been found on most con- tinents with the exception of Antarctica and Aus- tralia. They are predominant in Europe (47 valid species and 17 incertae sedis), Africa (17 valid species and 6 incertae sedis), and North America (19 valid species and 5 incertae sedis); this probably reflects the relative bias of collecting rather than corresponding to some real biogeographic patterns.

Less than half (46%) of the actinistian taxa (categories A-C) are known from entire skeletons var- ying in state of preservation. Of the remaining taxa, 36% are known almost exclusively from head or head fragments, 10% from scales solely, and 8% from partial trunk.

61

I = Category A

: = Category B i = Category C

Fig. 1. Diversity of actinistian species through geological time based on the listing of Table 1. Category A is represented by a straight bar, category B by a dotted line, and category C by an undulating line. Each bar corresponds to the number of species for a given biostratigraphic stage.

Patterns of diversity

Histograms of generic diversity through geological time have been provided by Schaeffer (1952a, 1953), Thomson (1977), andForey (1984,198). As pointed out by Smith & Patterson (1988), monotypic and paraphyletic taxa are problematic in generic-level studies because they provide historical- ly uninformative patterns. Among actinistians there is a high proportion of monotypic genera (see above) and others are known to be non-mono- phyletic (e.g., Rhabdoderma and Coelacanthus). However, temporal diversity studied at the specific level can be used to address historical questions (e.g., patterns and trends of species diversity).

Here we use Table 1 to provide some details of species diversity. We plot the number of species of categories A-C (above) against time (Fig. 1). The geological time scale of Palmer (1983) is used for the determination of age-boundaries and duration of geological units (i.e., age, period, and stage); in contrast to Palmer (1983), the Tertiary is divided into five epochs (Pliocene, Miocene, Oligocene,

Eocene, Paleocene; Holmes 1959) rather than 15 stages. The geological stage was used as the basic biostratigraphic unit for the count of taxa. In the diversity histograms, the bars were plotted according to the total diversity (i.e., number of taxa) occuring during a given stage. The bars were set on a proportional time axis (X-axis) at the mid-time between consecutive boundaries of each stage (i.e., [t&,1/2).

The proportional distribution of valid species (categories A and B) and known taxa (categories A-C; provided in parentheses) through geological era is as follows: 28.9% (33.05%) Paleozoic, 70% (66.12%) Mesozoic, 0% (0%) Cenozoic, and 1.1% (0.83%) Quaternary. There are four primary modes in the species diversity: (1) Frasnian with 6 species, (2) Namurian with 7 species (plus 2 incertae sedis), (3) Scythian with 17 species (plus 4 incertae sedis), and (4) Tithonian with 10 species (plus 1 incerta sedis). There are two major geological gaps in the fossil record of the actinistians: (1) Middle Jurassic (gap of ca. 29 My) and (2) Tertiary (gap of ca. 66My).

62

Table 1. List of actinistian species with their respective geological, stratigraphic, geographical, and morphological information. Fm. = Formation, Gr. = Group, Mbr. = Member, Subgr. = Subgroup, Supergr. = Supergroup

‘Category A’ Alcoveria Beltan 1972 A. brevis Betlan 1972 (type species)

Middle Triassic; Ladinian. Muschelkalk. Spain.

Allenypterus Melton 1969 A. montanus Melton 1969 (type species)

Lower Carboniferous; Namurian A (E,B). Bear Gulch Limestone Mbr., Heath Fm., Big Snowy Gr. Montana, USA.

Axelia Stensiii 1921 A. robustu Stensio 1921 (type species)

Lower Triassic; Smitho-Spathian. Sticky Keep Fm., Sassendalen Gr. West Spitsbergen.

A. elegans Stensio 1921 Lower Triassic; Smitho-Spathian. Sticky Keep Fm., Sassendalen Gr. West Spitsbergen.

Axelrodichthys Maisey 1986 A. araripensis Maisey 1986 (type species)

Lower Cretaceous; Apto-Albian. Lower Romualdo Mbr., Santana Fm. Ceara, Brazil.

Caridosuctor Lund & Lund 1984 C. populosus Lund & Lund 1984 (type species)

Lower Carboniferous: Namurian A (E,B). Bear Gulch Limestone Mbr., Heath Fm., Big Snowy Gr. Montana, USA.

Chagri’nia Schaeffer 1962 C. enodis Schaeffer 1962 (type species)

Upper Devonian; Famennian. Chagrin Shale. Ohio, USA.

Changxingia Wang & Liu 1980 C. aspratilis Wang & Liu 1980 (type species)

Upper Permian; Tatarian. Changxing Fm. Zhejiang Province, China.

Chinlea Schaeffer 1967 C. sorenseni Schaeffer 1967 (type species)

Upper Triassic; Carnian - Norian. (1) Chime Fm. and (2) Tecovas Fm., Dockum Gr. (1) Colorado, New Mexico, Utah, (2) Texas, USA.

C. sp. Ash 1978 Upper Triassic. Ciniza Lake Beds; Chinle Fm. New Mexico, USA.

entire skeleton

entire skeleton

entire skeleton

entire skeleton

entire skeleton

entire skeleton

entire skeleton

entire skeleton

entire skeleton

scales

63

Table 1. Continued

Coccoderma Quenstedt 1858 suevicum Quenstedt 1858 (type species) [C. nudum Reis 18881

Upper Jurassic; Tithonian. Solnhofen Lithographic Limestone. Germany.

harlemensis (Winkler 1871) [Coelacanthus, Undina] Upper Jurassic; Tithonian. Solnhofen Lithographic Limestone. Germany.

sp. Fabre et al. 1982 Lower Cretaceous; Barriasian. France.

Coelacanthus Agassiz 1839 C. granulatus Agassiz 1839 (type species) [C. hassiae Miinster 1842, C. gracilis Agassiz 1844,

C. caudalis Egerton 1850, C. macrocephalus Willemoes-Suhm 1869, C. granulosus Agassiz] Upper Permian; Ufimian. Kupferschiefer. Germany and England.

‘C.’ madagascariense Woodward 1910 [Rhabdoderma] Lower Triassic; Dienerian. Middle Sakamena Gr. Madagascar.

Diplocercides Stensiij 1922 D. kayseri (von Koenen 1895) (type species) [Holoptychius, Coelucanthus, Nesides schmidti

Stensio 19371 Upper Devonian; Frasnian. Germany [also Poland according to Gorizdro-Kulczycka (1950)].

D. davisi (Moy-Thomas 1937) [Rhabdoderma, R. (?) abdenense Moy-Thomas 19371 Lower Carboniferous; Visean Pi. Ireland and Scotland.

D. heiligenstockiensis (Jessen 1966) [Nesides] Upper Devonian; Frasnian. Upper Plattenkalk. Germany.

D. jaekeli Stensio 1922 Upper Devonian; Frasnian. Germany.

D. sp. Janvier 1974 Upper Devonian; Frasnian. Iran.

Diplurus Newberry 1878 D. longicaudatcts Newberry 1878 (type species) [Rhabdiolepis, R. gwyneddensis Bock 1959,

R. striata Bock 19591 Lower Jurassic; Hettangian. Newark Supergr. Connecticut, New Jersey, Pennsylvania, Virginia, USA.

entire skeleton

entire skeleton

entire skeleton

entire skeleton

entire skeleton

entire skeleton

head fragments, pectoral girdle

entire skeleton

partial head

lower jaws

entire skeleton D. newarki (Bryant 1934), Schaeffer 1954 [Coelacanthus, Osteopleurus, 0. milleri Shainin 1943,

0. milleri grantonensis Shainin 1943, Pariostegus myops Cope 18681 Upper Triassic; Carnian. Newark Supergr. New Jersey, Pennsylvania, Virginia, USA. entire skeleton

64

Table 1. Continued.

Euporosteus Jaekel1927 E. eifelianus Jaekel 1927 (type species)

Middle Devonian; Givetian. Germany.

Gurnbergia Martin & Wenz 1984 G. ommatu Martin & Wenz 1984 (type species)

Middle Triassic; Ladinian. Upper Muschelkalk. Germany.

partial head

head, partial trunk Hudronector Lund & Lund 1984 H. donbairdi Lund & Lund 1984 (type species)

Lower Carboniferous; Namurian A (E,B). Bear Gulch Limestone Mbr., Heath Fm., Big Snowy Gr. Montana, USA. entire skeleton

Heptunema Bellotti 1857 H. puradoxum Bellotti 1857 (type species)

Middle Triassic; Ladinian. Italy. entire skeleton

Holophugus Egerton 1861 H. gulo Egerton 1861 (type species) [Truchymetopon Ziussicum Hennig 19511

Lower Jurassic; Sinemurian. England. entire skeleton

Zndocoelucunthus Jain 1974 I. robustus Jain 1974 (type species)

Lower Jurassic; Toarcian. Kota Fm. India.

Latimeria Smith 1939 L. chalumnae Smith 1939 (type species) [Malania anjouanae Smith 1953,

Lutimeria anjouanae (Smith) Lenoble & Le Grand 19541 Recent. Comoro Islands.

entire skeleton

entire anatomy Laugiu Stensio 1932 L. groenlandicu Stensio 1932 (type species) [L. ? sp. Stensio 19321

Lower Triassic; Scythian. Wordie Creek Fm., Nordenskiold Subgr. East Greenland. entire skeleton

Libys Miinster 1842 L. polypterus Mtinster 1842 (type species)


L. superbus Zittel 1887 Upper Jurassic; Tithonian. Solnhofen Lithographic Limestone. Germany

head, trunk

entire skeleton Lochmocercus Lund & Lund 1984 L. aciculodontus Lund & Lund 1984 (type species)

Lower Carboniferous; Namurian A (E,B). Bear Gulch Limestone Mbr., Heath Fm., Big Snowy Gr. Montana. USA. entire skeleton

65

Table 1. Continued.

Lualabaea de Saint-Seine 1955 L. levichei de Saint-Seine 1955 (type species)

Upper Jurassic; Kimmeridgian. Lualabaea Series. Zaire.

L. henryi de Saint-Seine 1955 Upper Jurassic; Kimmeridgian. Lualabaea Series. Zaire.

Macropoma Agassiz 1835 M. lewesiensis (Mantel1 1822) (type species) [Amia?; Macropoma mantelli Agassiz 18351

Upper Cretaceous; Cenomanian - Santonian Chalk. England.

M. praecursor Woodward 1909 Lower-Upper Cretaceous; Albian - Cenomanian. Gault and Chalk. England.

M. speciosum Reuss 1857 [M. forte Fritsch 18781 Upper Cretaceous; Turonian. Czechoslovakia.

Macropomoides Woodward 1942 M. orientalis Woodward 1942 (type species) [Coelacanthe ‘B’ and Coelacanthe ‘c’ of Gaudant 19751

Upper Cretaceous; Cenomanian. Lebanon.

Mawsonia Woodward 1907 M. gigas Woodward 1907 (type species) [Mawsonia minor Woodward 19081

Lower Cretaceous; Apto-Albian. Romualdo Mbr., Santana Fm. Ceara, Brazil.

M. lavocati Tabaste 1963 Lower Cretaceous; Albian. Morocco.

M. libyca Weiler 1935 Lower Cretaceous; Albian. Egypt.

M. tegamensiswenz 1973 Lower Cretaceous; Aptian. Niger.

M. ubangiana Casier 1961 Lower Cretaceous; Neocomian. Zaire.

Miguashaia Schultze 1973 M. bureaui Schultze 1973 (type species)

Upper Devonian; Frasnian. Escuminac Fm. Quebec, Canada.

Mybzcanthus Stensio 1921 M. lobatus Stensio 1921 (type species)

Lower Triassic; Smitho-Spathian. Sticky Keep Fm., Sassendalen Gr West Spitsbergen.

entire skeleton

scales, fin rays

entire skeleton

head, partial trunk

entire skeleton

entire skeleton

entire skeleton

head fragments

head fragments

head

head fragments

entire skeleton

head fragments, scales

66

Table 1. Continued.

M. spinosus Stensiii 1921 Lower Triassic; Smitho-Spathian. Sticky Keep Fm., Sassendalen Gr. West Spitsbergen.

Piveteauiu Lehman 1952 head fragments

P. madagascariensis Lehman 1952 (type species) Lower Triassic; Dienerian. Middle Sakamena Gr. Madagascar.

Polyosteorhynchus Lund & Lund 1984 P. simplex Lund & Lund 1984 (type species)

Lower Carboniferous; Namurian A (E2B). Bear Gulch Limestone Mbr., Heath Fm., Big Snowy Gr. Montana, USA.

Rhabdoderma Reis 1888

entire skeleton

entire skeleton

R. elegans (Newberry 1856) (type species) [Coelacanthus; C. lepturus Agassiz 1844, C. robustus Newberry 1856, C. ornatus Newberry 1856, C. elongatus Huxley 1866, C. summiti Wellburn 1903, C. watsoni Aldinger 1931, Conchiopsisfiliferus Cope 1873, C. angul[ferus Cope 1873, C. corrugatus Moy-Thomas 1935, Hoplopygus binneyi Agassiz 18441

Upper Carboniferous; Namurian - Stephanian. Allegheny Gr. Ohio, USA; England, Scotland, Wales, Ireland, France, Belgium, Holland, Germany,

Ukraine. R. aldingeri Moy-Thomas 1937

Upper Carboniferous; Namurian A (EJ . Wales.

R. ardrosseme Moy-Thomas 1937 Lower Carboniferous; Visean P,. Calciferous Sandstone Series. Scotland.

R. exiguum (Eastman 1902) [Coelacanthus] Upper Carboniferous; Westphalian D. Carbondale Fm. Illinois, USA.

R. huxleyi (Traquair 1881) [Coelacanthus, Dumfregia] Lower Carboniferous; Visean CaS,. Upper Border Gr. Scotland.

R. tingleyense (Davis 1884) [Coelacanthus; C. mucronatus Pruvost 1913, C. granulostriatus Moy-Thomas 19351

Upper Carboniferous; Westphalian A-C. England, Wales, France, Belgium, Holland, Germany.

Sassenia Stensiii 1921 S. tuberculata Stensio 1921 (type species)


Scleracanthus Stensiii 1921 S. asper Stensio 1921 (type species)


entire skeleton

head, partial trunk

entire skeleton

entire skeleton

entire skeleton

entire skeleton

partial head


67

Table 1. Continued.

Spermatodus Cope 1894 S. pustulosus Cope 1894 (type species)

Lower Permian. Admiral Fm. Texas, USA.

Synaptotylus Echols 1963 head

S. newelli (Hibbard 1933) (type species) [Coelacanthus, C. arcuatus Hibbard 19331 Upper Carboniferous; Stephanian. Lansing Gr. Kansas, USA.

Ticinepomis Rieppel 1980 7: peyeri Rieppel 1980 (type species)

Middle Triassic; Ladinian.

entire skeleton

Grenzbitumen horizon. Switzerland. entire skeleton

Undina Miinster 1834 U. penicillata Miinster 1834 (type species) [Holophagus, Coelacanthus, Coelacanthus striolaris

Mufister 1842, C. koehleri Mtinster 1842, C. major Wagner 1863, Undina acutidens Reis 18881 Upper Jurassic; Tithonian. Solnhofen Lithographic Limestone. Germany. entire skeleton

U. cirinensis Thiolliere 1854 [Undina minuta Wagner 1863, Coelacanthus minutus Willemoes-Suhm 18691

Upper Jurassic; Tithonian. Solnhofen Lithographic Limestone. France and Germany. entire skeleton

Ii. purbeckensis Woodward 1916 Upper Jurassic; Purbeckian. England. partial head, trunk

Whiteia Moy-Thomas 1935 W. woodwardi Moy-Thomas 1935 (type species) [Coelacanthus evolutus Beltan 19801

Lower Triassic; Dienerian. Middle Sakamena Gr. Madagascar.

W africanus (Broom 1905) [Coelacanthus] Lower Triassic; Scythian. Upper Beaufort Beds; Beaufort Series. Republic of South Africa.

W. tuberculata Moy-Thomas 193.5 Lower Triassic; Dienerian. Middle Sakamena Gr.

entire skeleton

head, partial trunk

Madagascar. entire skeleton W. sp. Gardiner 1966

Lower Triassic; Smithian. Toad, Grayling, and Sulphur Mountain Fms. Alberta, British Columbia, Canada.

Wimania Stensio 1921 W sinuosa Stensio 1921 (type species) [Leioderma sinuata Stensid 19181


entire skeleton


68

Table 1. Continued.

Youngichthys Wang & Liu 1981 I: ninhuainsis Wang & Liu 1981 (type species)

Upper Permian; Tatarian. Changxing Fm. Zhejiang Province, China entire skeleton

‘Category B’ Bogdanovia Obrucheva 1955 B. orientalis Obrucheva 1955 (type species)

Upper Devonian; Frasnian. Central Kazakhstan, USSR.

Coccodema Quenstedt 1858 C. bavaricum Reis 1888


C. gigas Reis 1888 Upper Jurassic; Tithonian. Solnhofen Lithographic Limestone. Germany

C. substriolatum (Huxley 1866), Reis 1888 Upper Jurassic; Kimmeridgian. Kimmeridge Clay. England.

Coelucanthus Agassiz 1839 Coelacanthus banffensis Lambe 1916

Lower Triassic; Smithian. Spray River Fm. Alberta, Canada

C. gracilis Agassiz 1844 ? Middle Triassic. ? Germany.

‘c’. Iunzensis Reis 1900 Upper Triassic; Carnian. Lunz Sandstone. Austria.

C. welleri Eastman 1908 Lower Carboniferous; Tournaisian or Lower Visean. Kinderhook Limestone. Iowa, USA.

Gruphiurichthys White & Moy-Thomas 1937 G. callopterus (Kner 1866) (type species)

Upper Triassic; Carnian. Raibl Beds. Austria.

Hainbergia Schweizer 1966 H. gramdata Schweizer 1966 (type species)

Middle Triassic; Ladinian. Muschelkalk. Germany.

Mawsonia Woodward 1907 M. sp. Wenz 1981

Lower Cretaceous; Neocomian. Niger.

head fragments

head and trunk fragments

lower jaw

partial head

partial trunk

caudal skeleton

entire skeleton

trunk

entire skeleton

entire skeleton

lower jaw

69

Table 1. Continued.

Moe&o& Schaeffer & Gregory 1961 M. wellesi Schaeffer & Gregory 1961 (type species)

Middle Triassic; Anisian. Moenkopi Fm. Arizona, USA.

Rhipis de Saint-Seine 1950 R. moorseli de Saint-Seine 1950 (type species) [R. moorseli forma undulatus Casier 19653

Upper Cretaceous.

head fragments

Kinko beds; Kwango Series. Zaire.

R. tuberculatus Casier 1965 Upper Cretaceous. Kinko beds; Kwango Series. Zaire.

R. sp. Casier 1965 [R. sp. indet. Casier 19651 Upper Cretaceous. Kwango Series. Congo.

Sassenia Stensiii 1921 S. (?) guttata (Woodward 1912) [Coelacanthus]


Sinocoelacanthus Liu 1964 S. fengshanensis Liu 1964 (type species)

Lower Triassic. Lolou Series. Kwangsi Province, China.

Undina Mtinster 1834 U. (?) barroviensis Woodward 1890

Lower Jurassic; Sinemurian. England.

CJ. grandis Eastman 1914 Upper Jurassic; Tithonian. Solnhofen Lithographic Limestone. France.

U. willemoesi (Vetter’ 1881) [Macropoma, Heptanema] Upper Jurassic; Tithonian. Solnhofen Lithographic Limestone. Germany.

Wimania Stensiti 1921 W. (?) multistriata Stensiii 1921


scales

scales

scales

partial head, trunk

caudal skeleton

entire skeleton

trunk

entire skeleton


‘Category C’ Coelacanthopsis curta Traquair 1901 (type species)

Lower Carboniferous; Visean. Scotland.

Coelacanthus dendrites Gardiner Permian. Republic of South Africa.

entire skeleton

scales

70

Table 1. Continued.

C. phillipsii Agassiz 1844 Upper Carboniferous; Westphalian A. Halifax Hard Bed. England.

C. stensioei Aldinger 1931 Upper Carboniferous; Namurian E,. Germany.

C. sp. Aldinger 1931 Upper Carboniferous; Namurian E. Germany.

C. sp. indet. of Chabakov 1927 Upper Carboniferous; Stephanian B. Ukraine, USSR.

C. sp. of Fletcher 1884 Upper Carboniferous. Coal Measures. Nova Scotia, Canada.

Mawsonia sp. Campos & Wenz 1982 Lower Cretaceous; Apto-Albian. Romualdo Mbr., Santana Fm. Ceara, Brazil.

Mylacanthus? sp. or Scleracanthus? sp. Stensio 1921 Lower Triassic; Smitho-Spathian. Sticky Keep Fm., Sassendalen Gr. West Spitsbergen.

cf. Rhabdoderma Forey & Young 1985 Carboniferous; Dinantian. Scotland.

Undina picena Costa 1862 [ Urocomus] Upper Triassic; Norian. Italy.

Undina sp. Andersson 1916 Middle Triassic; Ladinian. Italy.

Wimania? sp. Stensid 1921 Lower Triassic; Smitho-Spathian. Sticky Keep Fm., Sassendalen Gr. West Spitsbergen.

Actinistia gen. et sp. indet. of Lelievre & Janvier 1988 Upper Devonian; Famennian. Morocco.

? Actinistia gen. et sp. indet. of Janvier, Lethiers, Monod & Balkas 1984 Lower Carboniferous; Tournaisian. Koprtilii Shales. Turkey.

Actinistia indet. of Schultze & Moller 1986 Middle Triassic; Anisian. Muschelkalk. Germany.

Coelacanth of Patton & Tailleur 1964 Lower Triassic; Scythian. Shublick Fm. Alaska, USA.

caudal skeleton

head fragments

head fragments

scale

gular plate

head

partial head

partial head

entire skeleton

head and trunk fragments

partial trunk

head fragments, scale

scale

scales

partial trunk

71

Table 1. Continued.

[Coelacanth] identical scales of Gardiner 1973 ? Upper Permian - Lower Triassic. Madumabisa Shales. Zimbabwe.

]Coelacanthe] Basisphenoide 1 of Wenz 1979 Upper Jurassic; Oxfordian France.

[Coelacanthe] Basisphenoide 2 of Wenz 1979 Middle Jurassic; Callovian. France.

[Coelacanthe] Car&entopttrygoide 1 of Wenz 1979 Upper Jurassic; Kimmeridgian. France.

Coelacanthe indttermint of Martin & Wenz 1984 Middle Triassic; Ladinian. Upper Muschelkalk. Germany.

Coelacanthidae cf. Undina of Forey, Monod & Patterson 1985 Upper Jurassic; Tithonian. Akkuyu Fm. Turkey.

Coelacanthidae indet. of Schultze & Chorn 1988 Upper Carboniferous; Stephanian. Bern Limestone Fm. Kansas, USA.

Coelacanthidae gen. et sp. indet. of Martin 1981 Upper Triassic. Morocco.

Coelacanthidae gen. et sp. indet. of Zidek 1975 Upper Carboniferous; Stephanian. Wild Cow Fm. New Mexico, USA.

Coelacanthidae nov. gen. of Dehm 1956 Middle Triassic. France.

Coelacanthoidea of Gall, Grauvogel & Lehman 1974 Lower Triassic; Scythian. Buntsandstein. France.

Forme ‘B’ of Campos & Wenz 1982 Lower Cretaceous; Apto-Albian. Romualdo Mbr., Santana Fm. Ceara, Brazil.

Undescribed actinistian of Cloutier (1990) Upper Devonian; Famennian. New York, USA.

Undet. coelacanth of Berger 1832 Upper Triassic. Keuper. Germany.

scales

basisphenoid

basisphenoid

quadrate, pterygoid fragment

scales

partial head

articular, scales

angular

scales, trunk fragments

partial trunk

entire skeleton

head

entire skeleton

partial head

72

Table 1. Continued.

Bunoderrna baini de Saez 1940 Upper Jurassic. Argentina.

Coelacanthus abdenesis Traquair 1903 Lower Carboniferous; Visean. Scotland.

C. distuns Wellburn 1902 Upper Carboniferous; Namurian. England.

C. gigunteus Winkler 1880 Triassic. Germany.

C. hindi Wellburn 1902 Upper Carboniferous; Namurian. England.

C. minor Agassiz 1844 Middle Triassic; ? Ladinian. LunCville, France.

C. munsteri Agassiz 1844 [ Undina] Carboniferous. Coal Lebach.

C. spinatus Wellburn 1902 Upper Carboniferous; Namurian. England.

C. tuberculatus Wellburn 1920 Upper Carboniferous; Namurian. England.

C. woodwardi Wellburn 1902 Upper Carboniferous; Namurian. England.

Dictyonosteus arcticus StensiB 1918 Middle Devonian. West Spitsbergen.

Rhabdoderma (?) aegyptiaca Heide 1955 Lower Carboniferous. Egypt.

‘Category D’

Celacantideo of Richter 1985 Lower Permian. Irati Fm. Brazil.

Coelacanth bone of Orvig 1986 Palaeocene. Sweden.

Coelacanth remains of Gardiner 1966 Upper Devonian Alberta, Canada.

Coelacanthidae gen. et sp. indet. of Dziewa 1980 Early Triassic. Knocklofty Fm. Tasmania, Australia.

Coelacanthidae genus non det. of Woodward 1895 [‘Coelacanth’ of Schaeffer 19411 Upper Jurassic. Talbragar Beds. New South Wales, Australia.

Coelacanthinien genre indtt. of Casier 1961 Lower Cretaceous. Congo.

(not positively determinable)

(nomen nudum)

(nomen nudum)

(nomen nudum)

(nomen nudum)

(nomen nudum)

(nomen nudum, dipnoan Conchopoma gadiforme Kner 1868) (nomen nudum)

(nomen nudum)

(nomen nudum)


(rhizodont)



(dipnoan)


(actinopterygian)


73

Trends of diversity

Schaeffer (1952b, p. 109) maintained that the orig- ination (speciation) rate of actinistians ‘. . . re- mained consistently low throughout their long history’. Raup et al. (1973) have suggested that the pattern of actinistian species diversity is one which shows an initial burst of speciation followed by a drastic decrease in diversity ending in a single sur- viving species.

The variation of the diversity through the duration of the group can be evaluated using non-parametric Spearman’s rank-correlation coefficient. This coefficient is used to quantify the correlation between the number of taxa and the geological age. This is done in order to describe the trends of diversity through time. Rank-correlations were calculated for the two different units of time: geological period (rs period) and biostatigraphic stage (rs stage ). The rankings were attributed as follows: rank 1 equals Middle Devonian, rank 2 equals Late Devonian, rank 3 equals Early Carboniferous, . . . , and rank 20 equals Quaternary for the geological period; rank 1 equals Givetian, rank 2 equals Fras- nian, rank 3 equals Famennian, . . ., and rank 48 equals Holocene for the biostratigraphic stage. The number of species occurring in any one of the geological periods and biostratigraphic stages are ranked numerically; for instance, the Early Triassic is the geological period and the Scythian is the biostratigraphic stage containing the most species and for the purpose of computation this is given the highest rank. For equal rankings of species num- bers, the computation procedure described by Dagnelie (1977) is invoked.

These rankings were then applied to the non- parametric Spearman’s rank-correlation coefficient algorithm [see Cloutier (1991) for discussion of the method].

The history of the actinistians ranged over 20 (14 with actinistians) geological periods (from Middle Devonian to Quaternary) including 48 (31 with actinistians) biostratigraphic stages (from Givetian to Holocene) for a corresponding duration of 380 million years. The coefficients of rank-correlation calculated for the geological periods and the biostratigraphic stages show a low negative correlation

(for categories A and B: r, perjod = - 0.4459, Y s srage = - 0.2455; for categories A-C: r, pen’od = - 0.4944, r, stage = - 0.3551) between the diversity-rank and time-rank. None of these correlations are significant. Therefore, these results support Schaeffer’s (1952b) conclusion that there has been no significant decreasing or increasing trends in species diversity through time,

Some of the peaks of actinistian diversity are coincident to major geological events. These events are not interpreted as causal effects on the diversity. They might have an influence on actual diversity because of vicariance events or habitat diversification, or an effect on apparent diversity by increasing the likelihood of fossilisation owing to an increase of sedimentation, for example. The Frasnian peak succeeds the Caledonian orogeny which affected Europe and precedes the Acadian orogeny which affected North America; most of the Frasnian species are found in north-east Amer- ica and Europe. A second peak of species diversity, chiefly reflecting North American actinistian fau- na, occurred during the Namurian, stage associated with the beginning of the Sudetian erogenic phase. The greatest diversity occurred during the Scythian which corresponds to a worldwide marine transgression (Schaeffer & Mangus 1976). The greatest actinistian diversity occurring during the Tithonian is documented mostly in Europe, and may be associated with the Late Kimmerian oro- genie phase. Finally, the Al&an peak may be broadly contemporaneous with the Subhercynian/ Austrian orogeny which influenced western Afri- ca; most of the Cretaceous actinistians are found in northern Africa and South America.

References cited

See ‘References cited’ in Forey & Cloutier (1991) for all taxonomic references.

Cloutier, R. 1991. Patterns, trends, and rates of evolution within the Actinistia. Env. Biol. Fish. 32: 23-58. (this volume)

Dagnelie, P. 1977. ThCorie et mkthodes statistiques. Applica- tions agronomiques. Vol. 1, 2nd ed. Presses Agronomiques de Gembloux, Gembloux. 378 pp.

Denison, R.H. 1978. Placodermi. pp. l-62. In: H.-P. Schultze

74

(ed.) Handbook of Paleoichthyology, Vol. 2.) Gustav. Fisch- er Verlag, Stuttgart.

Forey, P.L. 1991. Latimeria chalumnae and its pedigree. Env. Biol. Fish. 32: 75-97. (this volume)

Forey, P.L. & R. Cloutier. 1991. Literature relating to fossil coelacanths. Env. Biol. Fish. 32: 391-401. (this volume)

Holmes, A. 1959. A revised geological time scale. Trans. Edin- burgh Geol. Sot. 17: 183-216.

Lenoble, J. & Y. Le Grand. 1954. Le tapis de l’oeil du coelacanthe (Latimeria alzjouanae [Smith]). Bull. Mus. Hist. nat., Paris 26: 460-463.

Palmer, A.R. 1983. The decade of North American geology. 1983. Geologic time scale. Geology 11: 503-504.

Raup, D.M., S.J. Gould, T.J.M. Schoff & D.S. Simberloff. 1973. Stochastic models of phylogeny and the evolution of diversity. J. Geol. 81: 525-542.

Ride, W.D.L., C.W. Sabrosky, G. Bernardi & R.V. Melville (ed.). 1985. International Code of Zoological Nomenclature.

XX General Assembly of the International Union of Biolog- ical Sciences. International Trust for Zoological Nomencla- ture. H. Charlesworth & Co. Ltd., Huddersfield. 338 pp.

Schaeffer, B. & M. Mangus. 1976. An early Triassic fish as- semblage from British Columbia. Bull. Amer. Mus. Nat. Hist. 156: 517-563.

Smith, A.B. & C. Patterson. 1988. The influence of taxonomic method on the perception of patterns of evolution. Evol. Biol. 23: 127-216.

Thomson, K.S. 1977. The pattern of diversification among fishes. pp. 377-404. In: A. Hallam (ed.) Patterns of Evolution, as Illustrated by the Fossil Record, Development in Palaeontol- ogy and Stratigraphy, Vol. 5, Elsevier, Amsterdam.

Vorobyeva, E.I. & D.V. Obruchev. 1967. Subclass Sarcoptery- gii. pp. 420-509. In: Y.A. Orlov (ed.) Fundamentals of Pale- ontology, Vol. XI Agnatha, Pisces, Israel Program for Scien- tific Translations, Jerusalem. (Russian original 1964).

Cloutier and Forey 1991 Diversity of Extinct and Living Actinistian Fishes (Sarcopterygii)

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