Morphology, Phylogeny and Paleobiogeography of Fossil Fishes Honoring Meemann Chang
David K. Elliott, John G. Maisey, Xiaobo Yu & Desui Miao (editors)
Verlag Dr. Friedrich Pfeil • München
Morphology, Phylogeny and Paleobiogeography of Fossil Fishes D. K. Elliott, J. G. Maisey, X. Yu & D. Miao (eds.): pp. 247-274, 15 figs., 6 tabs. © 2010 by Verlag Dr. Friedrich Pfeil, München, Germany – ISBN 978-3-89937-122-2
Critical analysis of the impact of fossils on teleostean phylogenies, especially that of basal teleosts Gloria Arratia
Abstract Traditionally, fossils have played little role in most studies of the phylogenetic relationships of teleosts. The usual approach is to study only recent fishes and when fossils are considered their position is assumed in the cladogram. During the last few years this approach has been challenged by the inclusion of both fossil and recent species in phylogenetic studies. Some fossils and their characters play a significant role, shown in phylogenetic studies of basal teleosts, of osteoglossomorphs, and of some paracanthopterygians. An important example is studying whether elopomorphs or osteoglossomorphs are the most basal teleostean lineage. To address this problem, an analysis of the content of both groups and their fossil record is presented first. It is followed by discussion of results obtained in different phylogenetic analyses. When fossil and recent taxa are included in phylogenetic analyses, the elopomorphs stand as the most plesiomorphic group among extant teleosts. When only recent taxa are included in the analysis, the osteoglossomorphs occupy the basal position; however, this scenario is changing because recent molecular phylogenetic studies support the elopomorphs as the most plesiomorphic extant group. Some fossil taxa may present characters or a combination of characters that contradict interpretations based only on extant forms. This may be particularly important for fishes such as elopomorphs with a remarkable fossil record of more than 150 million years. Furthermore, elopiforms had greater species diversity in the Late Jurassic than they do today. The same is true of osteoglossomorphs with a slightly younger fossil record of about 140 million years and a diversity of Cretaceous forms. When studying the evolutionary history of teleosts, to exclude fossils is a mistake. This is especially true of stem members belonging to monophyletic groups with a long history. It biases interpretation of characters, their distribution, and their homology, because the characters present in the recent terminal forms may have undergone significant evolutionary transformation through time to reach their present states.
Introduction Phylogenetic studies of teleostean fishes have taken different approaches during the last 40 years, and have been greatly influenced by Hennig (1966). However, although this was an important publication, it had negative implications concerning the role of fossils in phylogenetic studies of crown taxa. According to Hennig (1966), modern organisms are the starting point for the study of phylogenetic relationships, an idea supported later by Brundin (1968), Nelson (1969, 1978), Løvtrup (1977), and Patterson (1977, 1981a, 1981b, 1982) among others. However, some statements in these publications reveal a lack of understanding of the role of fossils in phylogenetic systematics. For instance, Nelson (1969: 22) noted: “the paleontologist, even with a good fossil record, can contribute very little, if anything, to the solution of this problem [the phylogenetic relationships among extant taxa]. In fact, the paleontologist as a rule cannot do much in determining the relationships of his fossil species unless there are Recent relatives whose relationships are fairly well established. Certainly, as far as teleosts are concerned, and probably other vertebrate groups, as well, paleontology has been of little or no direct help in elucidating the phyletic relationships among any Recent animals. This has been and will continue to be done mainly through 247
detailed comparative study of the diversity of the Recent fauna (see Brundin 1966).” Patterson (1981a: 218) wrote, “. . . instances of fossils overturning theories of relationships based on Recent organisms are very rare, and may be non-existing. It follows that the wide-spread belief that fossils are the only, or best, means of determining evolutionary relationships is a myth.” Such comments and others originated some well-documented publications showing the importance of certain fossil vertebrates in phylogenetic studies of some groups (e. g., Maisey 1986; Gauthier et al. 1988; Arratia 1991, 1997, 1999; Wilson 1992; Murray & Wilson 1999, 2005). There is a general consensus that living organisms are the starting point for the study of phylogenetic relationships. Phylogenetic systematics can formulate hypotheses of relationships of recent species without considering any fossil record and then can translate the hypotheses of relationships into a system with hierarchical structure. However, the aims of phylogenetic systematics are larger than this, because it strives for the phylogenetic relationships of all organisms with each other, whether they exist today as evolutionary species or are known only by fossil remains (Ax 1987). This reasoning raises the question of the phylogenetic systematization of fossil organisms in relation to the recent fauna. For instance, the fossil †Archaeopteryx lithographica – currently known from 10 specimens – from the Upper Jurassic of Solnhofen, Germany, is placed in the taxon Aves on the basis of characters shared with modern birds. †Leptolepis coryphaenoides, a fish from the Lower Jurassic of Europe is placed in the taxon Teleostei on the basis of shared synapomorphies with recent teleosts. †Lycoptera (an Early Cretaceous fish from Asia) and †Anaethalion (a Late Jurassic fish from Europe) are placed in the Osteoglossomorpha and the Elopomorpha, respectively, because of shared synapomorphies with recent members of these taxa. These fossils tell us that if the taxa Aves, Teleostei, Osteoglossomorpha, and Elopomorpha can be justified as monophyletic groups, then the following statement logically applies: “closed descent communities in Nature may comprise recent organisms and also extinct organisms known only as fossils . . . It follows, namely, that all organisms, whether recent or known as fossils, are to be united together in a single phylogenetic system.” This is an inevitable requirement “because the monophyletic taxa of the system we aim to build must always include all descendents of their respective stem system if they are to be valid representatives of the closed descent communities in Nature.” (Ax 1987: 201). Thus, how have studies of phylogenetic relationships of teleosts been approached during the last 40 years? Do they conform to the criteria formulated by Hennig (1966), Nelson (1969), and Patterson (1977, 1981a, 1981b, 1982) or Ax (1987)? According to my interpretation of the literature four main approaches are discernable, as listed below. Approach no. 1: Phylogeny is built on the basis of only recent species. This is the most common tendency observed among ichthyologists because the main aim of their work is to investigate the phylogenetic relationships of living species. It also may be a consequence of the fact that there is a poor fossil record for numerous teleostean groups. Examples of this approach are most teleostean phylogenies in Stiassny et al. (1996), and in most phylogenies published in ichthyological journals during recent years. Approach no. 2: Phylogeny is built only on fossil species because living representatives are unknown or presumably unknown. For instance, †varasichthyids (Arratia 1994: fig. 9) and †ichthyodectiforms (Stewart 1999: figs. 12, 13). Approach no. 3: Phylogeny is built on the basis of recent species and the fossils are added to the phylogeny following an assumption of where the fossils may belong. This conforms to Hennig’s (1966) ideas of the use of fossils. Examples are †Anaethalion and †Leptolepides in Patterson & Rosen (1977: fig. 54) and †Brannerion and †Osmeroides in Forey et al. (1996: fig. 5). Approach no. 4: Phylogeny is built on the basis of fossil and recent species. This approach conforms with Ax’s (1987) ideas as above cited. This is the main tendency developed during recent years among paleontologists studying teleosts and it is beautifully demonstrated in different contributions in the Mesozoic Fishes series of books (Arratia & Viohl 1996, Arratia & Schultze 1999, Arratia & Tintori 2004), the Zoological Journal of Linnean Society, Journal of Vertebrate Paleontology, and others. This approach demands a broader knowledge from the specialist, one that involves education in preparation, observation, and interpretation of both fossil and extant species. The main goal of this contribution is to analyze the effect of these approaches in phylogenetic hypotheses of the intrarelationships of the elopomorphs and osteoglossomorphs and of their positions at the base of the crown-group Teleostei. To reach this goal, I first will introduce the reader to the current understanding of both the elopomorphs and osteoglossomorphs. 248
Institutional abbreviations: IVVP, Institute of Vertebrate Palaeontology and Palaeoanthropology, Academia Sinica, Beijing, China; JME, Jura Museum, Eichstätt, Germany; LACM, Los Angeles County Museum of Natural History, Los Angeles, U.S.A.
Elopomorpha General overview and content The superorder Elopomorpha is represented by approximately 850 species and 156 genera placed in four orders (Elopiformes, Albuliformes, Anguilliformes, and Saccopharyngiformes; Nelson 2006). Currently, the phylogenetic relationships among elopomorphs are disputed by various authors and consequently there are conflicting opinions. According to the phylogenetic studies by Forey et al. (1996) based on morphological and molecular data, elopomorphs include the Elopiformes, Albuliformes, Notacanthiformes, and Anguilliformes (containing the saccopharyngoids), whereas Inoue et al. (2004), based on molecular evidence, included the notacanthiforms within the Albuliformes. Independent of these conflicts affecting the high level classification of elopomorphs, the order Elopiformes, with two extant genera and about 8 species, is smaller than the other three orders. The largest order is the Anguilliformes with 141 extant genera and over 790 species (Nelson 2006). The Elopomorpha, as presently understood, is a heterogeneous group united by the leaf-like, transparent leptocephalous larvae and by the possession of derived sperm morphology (see references in Stiassny et al. 2004). In general, fossil elopomorphs are not well known, especially anguilliforms and saccopharyngiforms. A problematic group is the order †Crossognathiformes, represented by Jurassic to middle Eocene species that were assigned to Elopomorpha by Maisey (1991) and Nelson (1994), to basal clupeocephalans by Taverne (1989), and to basal teleosts by Arratia (2008a). A few forms previously considered to be in the family †Anaethalionidae (e. g., †Anaethalionopsis and †Manchurichthys) by Gaudant (1968) or to be megalopids (e. g., †Pachythrissops) by Forey (1973) are no longer considered within the Elopomorpha (for information about these genera see Taverne 1981, Chang 1999, and Arratia 1997, respectively). The addition of fossils greatly increases the total number of elopomorph genera because some elopomorph subgroups were more diversified in the past than today. The greater generic diversity in the past is not a unique feature of osteoglossomorphs as pointed out by Patterson (1994), but also of elopiforms among elopomorphs. For instance, elopiforms are presently known from two genera in contrast to 15 fossil genera ranging from the Late Jurassic (Kimmeridgian) to late Neogene (see Arratia 2000: table 5). Since the classic revision of elopiforms and albuliforms by Forey (1973), new information has been generated. For instance, some fossils previously assigned to the Elopomorpha incertae sedis or to the elopiforms became better known during the last two decades due to: (1) revision of some Late Jurassic forms such as †Anaethalion angustus (type species of the genus: Figs. 1A, 2A; Arratia 1987, Poyato-Ariza 1999), †A. angustissimus, †A. knorri, †A. subovatus (Arratia 1987, 1997), and †A. mayri (now known as †Eichstaettia mayri; Arratia 1987); (2) revisions of some Early Cretaceous forms such as †‘Anaethalion’ vidali (now known as †Ichthyemidion vidali; Poyato-Ariza 1995) from the Lower Cretaceous of Montsec, Spain, and †Pachythrissops vectensis (now known as †Arratiaelops vectensis; Taverne 1999) from the Lower Cretaceous of Belgium and England; and (3) new findings such as †Anaethalion zapporum (Fig. 1B), †Daitingichthys, †Elopsomolos (Fig. 2C; Arratia 1987, 2000) in the Upper Jurassic of Bavaria, and †Naiathaelon okkidion (Poyato-Ariza & Wenz 1994) in the Upper Jurassic of Canjuers, France. †Anaethalion zapporum (Figs. 1B, 2B) from the Kimmeridgian of Schamhaupten, southern Germany, is the oldest described member of †Anaethalion. It is important to note that recently one specimen of †Anaethalion (pers. obser.) was recovered in Wattendorf, northeast of Bamberg, near the northern tip of the Franconian Alb, which is currently interpreted as the oldest known plattenkalk in southern Germany (late middle Kimmeridgian; Fürsich et al. 2005). In addition to the oldest known elopomorphs mentioned above, several younger forms from the Cretaceous (e. g., †Davichthys, †Elopoides, †Flidersichthys, and †Sedenhorstia; Forey 1973) and from the Paleogene (Elops, †Promegalops, and †Protarpon; Forey 1973, Arratia 2000) can be added to the list. However, none of the Cretaceous and Paleogene forms have been revised recently. The knowledge of albuliforms has also been enriched due to (1) the revisions of †Brannerion (Fig. 3B) by Blum (1991) and Forey & Maisey (2010) and †Paraelops by Maisey & Blum (1991) based on new material from the Santana Formation, Lower Cretaceous of Brazil, and (2) description of the new albuliform 249
A
2 cm
B
C
2 cm
2 cm
Fig. 1. Examples of some of the oldest known elopomorphs. A, †Anaethalion angustus (neotype JME SOS2271) from the Tithonian of Eichstätt, Bavaria, Germany. B, †Anaethalion zapporum (holotype JME SCHA 85) from the Kimmeridgian of Schamhaupten, Bavaria, Germany. C, Eichstaettia mayri (JME SOS2283) from the Tithonian of Blumenberg, Bavaria, Germany. (Photographs executed by C. Radke).
genus †Baugeichthys (Fig. 3A; Filleul 2000a) from the Lower Cretaceous of France. Although the fossil genera of albuliforms (about 13, excluding otolith-based genera) are few in number, they were more diversified in the Cretaceous than today (eight genera; Nelson 1994, 2006). The oldest known Albuloidei is the otolith-based genus “Albuloideorum” from the Lower Cretaceous of Germany and France (Nolf 1985, Patterson 1993). The oldest albulid genus, †Lebonichthys, known from complete specimens is from the Cenomanian of Lebanon (Patterson 1993). The extant genera Istieus (= Pterothrissus) and Albula are known from Upper Cretaceous and Paleogene deposits, respectively, and the halosaurids (e. g., †Echidnocephalus) from the Upper Cretaceous of Westphalia, Germany. Apparently, Albula was already present in the Upper Cretaceous of North America according to Cockerel (1933) who described †Albula antiqua from Florida and to Applegate (1970) who described †A. dunklei from Alabama. 250
A
2 cm
B 2 cm
C 3 cm
Fig. 2. Restorations of some Late Jurassic elopomorphs from Bavaria, southern Germany, to show the extent of preserved morphological details (incomplete lines correspond to missing information). A, †Anaethalion angustus (restoration based on JME SOS2261a, JME SOS2271, JME SOS2284a), Tithonian. B, †Anaethalion zapporum (after Arratia 2000: fig. 11), Kimmeridgian. C, †Elopsomolos frickhingeri (after Arratia 2000: fig. 16).
The North American material assigned to albulids needs to be taxonomically revised to test if the range of Albula extends to the Cretaceous. The Anguilliformes and the Saccopharyngiformes are the most specialized and diversified taxa within the Elopomorpha. Currently, the order Anguilliformes comprises 15 families, whereas the Saccopharyngiformes has only four (Nelson 2006). In contrast to the 141 modern genera, only a few fossil genera are known. The oldest records of anguilliforms (e. g., †Anguillavus, †Urenchelys) are known from the Late Cretaceous (early Cenomanian; Patterson 1993). Robins (1989) questioned the assignment of some of the fossil forms (e. g., †Anguillavus and †Urenchelys) to the order Anguilliformes. According to Patterson (1993) they are “true” eels and he interpreted them as Anguilloidea incertae sedis. Recent studies of the Cretaceous Anguilliformes recognized at least six Lebanese genera: †Urenchelys (with †U. avus), †Anguillavus (with †A. quadripinnis and †A. mazeni), †Luenchelys (with †L. minimus), †Abisaadia (with †A. hakelensis), and †Hayenchelys (with †H. germanus) (Belouze 2002, Belouze et al. 2003a, 2003b). 251
A 2 cm
B 3 cm
Fig. 3. Restorations of some Cretaceous albuliforms, to show the extent of preserved morphological details (incomplete lines correspond to missing information). A, †Baugeichthys caeruleus (slightly modified from Filleul 2001, reversed), Hauterivian of France. B, †Brannerion latum (modified from Blum 1991), Lower Cretaceous of Brazil.
[According to these authors, †Urenchelys anglicus Woodward, 1900 and †Urenchelys abditus Wiley & Stewart (1981) are too poorly preserved or incompletely known to permit an accurate generic determination.] More recently, Taverne (2004) described a new family including a new genus and species from Lebanon, †Libanechelys bultyncki, and another new genus and species from the Upper Cretaceous of Nardò, Italy, †Nardoechelys robinsi (Taverne 2002) that was assigned to the extant family Ophichthidae. The revisions of previously described material and descriptions of new material have greatly increased the knowledge of the morphology of Cretaceous anguilliforms. In addition to much diversity of anguilliforms recovered in the Upper Cretaceous of Lebanon, there is much material from the Paleogene of Bolca, Italy. For example, based on fragmentary material, Blot (1978, 1980, 1984) described 15 fossil genera assigned to five fossil anguilliform families (†Anguilloididae, †Milananguillidae, †Paranguillidae, †Proteomyridae, and †Patavichthidae), and fossil representatives of at least three extant families (e. g., Anguillidae, Congridae, and Ophichthidae). Two genera were left as Anguilliformes incertae sedis. Although Cretaceous fossil anguilliforms have been recently revised and new forms have been described, the knowledge of the 252
material from Bolca has not increased in the last 20 years. The first described fossil ophidiid, †Genypterus valdesensis, from South America was recently recovered in the Miocene of Argentina (Riva Rossi et al. 2000). Overview on geographical distribution and environment Recent elopomorphs are mainly marine forms, living in tropical and subtropical oceans (e. g., Elopidae, Megalopidae, Anguilliformes) or in most tropical seas (e. g., Albulidae) (Nelson 2006). Some families as Notacanthidae and Halosauridae have worldwide distribution. Some elopomorphs occasionally enter freshwater (e. g., Megalopidae, Muraenidae, Moringuidae, Ophichthidae), and the Anguillidae are usually catadromous. Some, like the notacanthoids, live in the deep sea, whereas others are pelagic (e. g., Serrisomeridae) or bathy- and mesopelagic (e. g., Nemichthyidae). Most of the fossil elopomorphs have been recovered in marine localities. For instance, the earliest elopiforms (e. g., †Anaethalion, †Elopsomolos, †Eoprotelops, †Naiathaelon) are found in marine limestones of different localities in the Upper Jurassic of Bavaria, in the Kimmeridgian of Nusplingen (Germany), in the Kimmeridgian of Cerin, and in the Tithononian of Canjuers (France). The oldest albuloid (“Albuloideorum”) was recovered in localities in Germany and France that are interpreted as marine. The oldest anguilliforms (†Anguillavus and †Urenchelys) have been recovered in marine localities in Lebanon. Phylogenetic relationships among elopomorphs In this section, I will concentrate my analysis and evaluation of the role of certain fossils and of their interpretation within elopomorphs based on results published in the last 20 years and my own research. Recent development in elopomorph research includes: (1) Re-assessment of the morphology and re-interpretation of few fossils: e. g.,†Anaethalion from Bavaria, Germany (Fig. 2A; Arratia 1987, 1997), †Brannerion from Brazil (Fig. 3B; Blum 1991, Forey & Maisey 2010), †Anguillavus (Belouze & Gayet 1999, Belouze et al. 2003a), and †Luenchelys from Lebanon (Belouze et al. 2003a). The information on new anguilliforms points to the presence of complete pectoral girdle and pelvic fins in the oldest forms (new specimens of †Anguillavus; Belouze & Gayet 1999). (2) Re-assessment of certain morphological features of some extant forms, e. g., Elops, Albula, and Megalops. Some of these features include the circumorbital series, vertebral column, intermuscular bones, fin rays, and caudal skeleton (1996a; Arratia 1987, 1997, 1999, 2008b; Schultze & Arratia 1988; Arratia et. al. 2001). (3) New discoveries of fossil taxa, including fragmentary or fairly complete or complete specimens. Examples are: species of †Anaethalion, †Daitingichthys and Elopsomolos (Arratia 1987, 1997, 2000), †Naiathaelon (Poyato-Ariza & Wenz 1994), †Baugeichthys (Filleul 2000a), †Libanechelys (Taverne 2004), and †Luenchelys (Belouze at al. 2003a). The three aspects mentioned above have enlightened the knowledge of elopomorphs and of their relationships, as it will be shown below. Studies on phylogenetic relationships among elopomorphs are rare and do not represent the great diversity of the group. For instance, the current available hypotheses, based on extant forms including members of the four orders, is that by Forey et al. (1996; Fig. 4 Node A) and Inoue et al. (2004; Fig. 5, Node A). As shown in Figure 4, only eight elopomorph taxa (1 albulid, 2 elopiforms, and 5 anguilliforms) and five outgoup taxa (3 clupeiforms and 2 osteoglossomorphs) were included in the phylogenetic analysis of elopomorphs of Forey et al. (1996) that was based on molecular and morphological evidence of recent forms. A classification of the group, based on the cladogram shown in Figure 4, was proposed by Forey et al. (1996; Table 1). The molecular study by Inoue et al. (2004; Table 2) shows major differences to that of Forey et al. (1996), in the content of Albuliformes
B A
C
Albula Ophichthus Echiophis Eurypharynx Anguilla Notacanthus Elops Megalops Stolothrissa Limnothrissa Clupea Osteoglossum Petrocephalus
Fig. 4. Hypothesis of phylogenetic relationships of modern elopomorphs after Forey et al. (1996). Labels identifying nodes of elopomorphs are added, as well as their branching is shown in thicker lines, to facilitate comparisons.
253
Table 1. Classification of Elopomorpha (using the sequencing convention). The fossils were added to the classification of recent taxa following approach no. 3 in the Introduction. After Forey et al. 1996. Compare with Figure 4.
Table 2. Classification of Elopomorpha (using the sequencing convention) based on molecular data of living forms after Inoue et al. 2004. Approach no. 1 in the Introduction. Compare with Figure 5.
Cohort Elopomorpha
Cohort Elopomorpha
Order Elopiformes Family Elopidae Elops Family Megalopidae Megalops
Order Elopiformes Family Elopidae Elops Family Megalopidae Megalops
Order Albuliformes Plesion †Brannerion sedis mutabilis Plesion †Osmeroides sedis mutabilis Suborder Albuloidei sedis mutabilis Family Albulidae Albula Family Pterothrissidae
Order Albuliformes Family Albulidae Albula Pterothrissus Family Halosauridae Aldrovandia Family Notacanthidae Notacanthus
Order Notacanthiformes Family Halosauridae Family Notacanthidae Subfamily Notacanthinae Subfamily Polyacanthonotinae Tribe Polyacanthonotini Tribe Lipogenyini Order Anguilliformes Suborder ‘Anguilloidei’ 16 families listed by J. S. Nelson (1994) Suborder Saccopharyngoidei Family Saccopharyngidae sedis mutabilis Family Eurypharyngidae sedis mutabilis Family Monognathidae sedis mutabilis
Order Anguilliformes Family Anguillidae Anguilla Family Muraenidae Gymnothorax Family Synaphobranchidae Synaphobranchus Family Ophichthidae Ophisurus Family Congridae Conger Order Saccopharyngiformes Family Saccopharyngidae Saccopharynx Family Eurypharyngidae Eurypharynx
and the Anguilliformes (*). The notacanthids are included within the Albuliformes in the phylogenetic hypothesis of Inoue et al. (2004; see Fig. 5, Node D), whereas they are members of the Anguilliformes in the phylogenetic hypothesis of Forey et al. (1996; Fig. 4, Node C).
*
254
Changes in the content of a fish order are major changes and should require a well-supported database and justification. It surprises me that Inoue et al.’s (2004) do not provide a well-documented discussion to explain the differences between their phylogenetic hypothesis concerning the elopomorphs, based on molecular evidence, and those based on morphological data. The cladogram (Fig. 5, Node F) also shows another major difference from the current interpretations of Ostariophysi: a new clade including [Gonorynchiformes + Clupeomorpha] as the sister group of Otophysi. Consequently, according to their results, the Ostariophysi as presently interpreted – based on morphological evidence – is not monophyletic according to Inoue et al. (2004). In contrast, Ishiguro et al.’s (2003) molecular analysis indicates another major phylogenetic arrangement: [Clupeomorpha + Alepocephaloidea] as sister group of Ostariophysi. Furthermore, studies of complete nucleotide sequence of the cytochrome b gene of some ictalurids and cyprinids resulted in a nonmonophyletic order Siluriformes (Matsuo et al. 2000). Again, and despite the importance of such different phylogenetic hypotheses, no discussion of these results in contrast to those of morphological studies was offered, but the conclusion that “genetic reclassification of fishes may be necessary to identify the ancestor” (Matsuo et al. 2000: 207).
C
Pagrus major
Polymixia japonica
Chlorophthalmus agassizi
Protacanthopterygii Coregonus lavaretus
Salmo salar
Oncorhynchus mykiss
Otophysi Crossostoma lacustre
Cyprinus carpio
Gonorynchus greyi
Chanos chanos
Anotophysi
Clupeomorpha Sardinops melanostictus
Engraulis japonicus
Saccopharyngiformes D1
F2
D5 D4
D2
A1
Eurypharynx pelecanoides
Saccopharynx lavenbergi
Anguilla japonica
Anguilliformes Conger myriaster
Ophisurus macrorhynchus
Synaphobranchus kaupii
Gymnothorax kidako
Notacanthus chemnitzi
Albuliformes Aldrovandia affinis
Pterothrissus gissu
Albula glossodonta
Megalops atlanticus
Megalops cyprinoides
Elops saurus
Elops hawaiensis
Elopiformes
Osteoglossomorpha Osteoglossum bicirrhosum
Pantodon buchholzi
Hiodon alosoides
D3
F1 G
F
D B
E
A
Fig. 5. Hypothesis of phylogenetic relationships of modern elopomorphs (at node A) and more advanced teleosts (at node E) after Inoue et al. (2004). Nodes identifying taxa are added to facilitate comparisons.
Anguilla is the sister group of saccopharyngiforms and saccopharyngiforms are nested within Anguilliformes so that the later is paraphyletic according to the results by Inoue et al. (2004, see Fig. 7, Node D). Inoue et al.’s (2004) molecular study is based on 15 species of elopomorphs (4 elopiforms, 4 albuliforms including notacanthiforms, 5 anguilliforms and 2 saccopharyngiforms) that correspond to the largest amount of elopomorphs included in a phylogenetic analysis. The phylogenetic relationships of fossil and recent elopomorphs have also been scarcely explored, as explained below. The phylogenetic relationships of six species of †Anaethalion and extant Elops (represented by E. saurus) were explored by Arratia (1997: fig. 38A-D) based on 23 characters. I concluded that †Anaethalion as presently understood is paraphyletic but such a result could be due to incomplete information on certain characters (similar results were obtained by Arratia 2000: fig. 22). In addition, the recognition of two families, the fossil †Anaethalionidae and the extant Elopidae was questioned. In the same paper, I studied also the phylogenetic relationships of 26 fossil and nine recent basal teleostean genera including †Anaethalion and Elops and the results showed that †Anaethalion and Elops are sister groups (Arratia 1997: figs. 99-102). Later, Arratia (1999; see below Fig. 15A) added Megalops into the analysis (because I was interested in exploring the relationships among primitive members of different teleostean subgroups), as well as other taxa. More recently, Arratia (2000; Fig. 6), based on 191 characters and 30 fossil taxa and 14 255
Amia †Aspidorhynchus †Belonostomus †Vinctifer Lepisosteiformes †Mesturus †Allothrissops †Thrissops †Ascalabothrissops †Pachythrissops †Anaethalion angustus †Anaethalion zapporum Elops Megalops Chanos †Gordichthys Opsariichthys †Tischlingerichthys Denticeps Engraulis †Santanaclupea †Diplomystus †Erichalcis †Leptolepides haerteisi †Leptolepides sprattiformis †Orthogonikleithrus leichi †Orthogonikleithrus hoelli †Humbertia Esox Umbra Oncorhynchus Thymallus Hiodon Heterotis †Lycoptera †Ascalabos †Domeykos †Luisichthys †Protoclupea †Varasichthys †Tharsis †Leptolepis coryphaenoides †Pholidophorus bechei †Pachycormiformes
Teleostei
2b
F2
F3 3
2a
G2
F1
L2
K2
G3
K1
G1
2 1
O1
O2 O
N2 L
K
N4
N3 L1
D3
H2
D2
H1 D1
N1 N
J
M
Elopiformes
Osteoglossiformes
I
†Ichthyodectiformes
H G
†Varasichthyidae
F E D
C B A
Fig. 6. Hypothesis of phylogenetic relationships of some basal teleosts including fossil and recent elopiforms after Arratia (2000: fig. 21). Branching of elopiforms and osteoglossomorphs are shown in thicker lines.
extant taxa, studied the phylogenetic position of the newly described †Anaethalion zapporum, the oldest known species of the genus, and the results showed †Anaethalion as the sister group of [Elops + Megalops]. Filleul (2000b; Fig. 7, Nodes A1 and C1), based on the list of characters and the data matrix of Arratia (1997), but using a broader ingroup that included four albuliforms (Albula, †Baugeichthys, †Lebonichthys, and Pterothrissus), concluded that elopomorphs are non monophyletic and albuliforms are closer relatives to euteleosts (Fig. 7, Node C) than to elopiforms. He reached a similar conclusion concerning the monophyly of elopomorphs when studying molecular evidence (nucleotid sequences of ribosomal RNA 18S, 16S, and 12S) of only extant forms (Filleul & Lavoué 2001, see below). Recently, Obermiller & Pfeiler (2003), based on mitochondrial ribosomal DNA sequences, concluded that elopomorphs are not monophyletic. These results contradict those by Forey et al. (1996), Wang et al. (2003), and Inoue et al. (2004), based also on molecular evidence. The phylogenetic studies by Arratia (1997, 1999, 2000) and Filleul (2000b) included fossil and recent forms in the ingroup. Consequently, these studies correspond to approach no. 4 mentioned in the Introduction. The results from these authors agree that †Anaethalion is the sister group of extant Elops or extant [Elops + Megalops], and †Anaethalion is considered an elopiform. In the phylogenetic hypothesis by Patterson & Rosen (1977: fig. 54), the position of †Anaethalion was assumed and it was added at the base of the Elopomorpha (approach no. 3 in the Introduction). Recently, Taverne (2004: fig. 9) published a scheme showing the systematic position of some Lebanese Late Cretaceous fossil eels. Based on 47 characters taken mainly from the literature, he distinguished 256
Combined outgroup †Allothrissops †Thrissops †Pachythrissops †Anaethalion Elops †Daitingichthys Chanos †Gordichthys †Tischlingerichthys †Chongichthys Denticeps Engraulis †Santanaclupea †Diplomystus †Erichalcis †Leptolepides haerteisi †Leptolepides sprattiformis †Orthogonikleithrus hoelli †Orthogonikleithrus leichi †Humbertia Esox Umbra Oncorhynchus Thymallus †Baugeichthys Albula †Lebonichthys Pterothrissus Hiodon †Lycoptera †Pattersonella †Cavenderichthys †Tharsis †Ascalabos †Domeykos †Luisichthys †Protoclupea †Varasichthys †Leptolepis coryphaenoides B1 A1 C1 Albuliformes D Elopiformes
C B A
Fig. 7. Hypothesis of phylogenetic relationships of some fossil and recent teleosts, with special reference to the elopomorphs (after Filleul 2000b: fig. 3). Branching of elopiforms (at node A), albuliforms (at node C), and osteoglossomorphs (at node B) are shown in thicker lines. Nodes identifying taxa are added to facilitate comparisons.
between ancient eels and “anguilliforms of modern anatomical pattern”, and placed †Libanechelys in an intermediate position between the two groups. Knowledge of the phylogenetic relationships among extant eels is incomplete, and this is due mainly to the large taxonomic diversity of the group; additionally, the knowledge of the whole group, including fossil and extant eels is incomplete and new findings of fossils eels are showing important characters such as the presence of structures that are absent in the modern forms. This alone demonstrates the need to enlarge the morphological information on oldest eels for a better understanding of the evolutionary history of anguilliforms. In general, the current knowledge of fossil elopomorphs is scarce. Among them, the Jurassic elopiforms from southern Germany are better known than Cretaceous and Paleogene taxa. Among albulids and anguilliforms, only few taxa are relatively well known. Consequently, as shown above, the phylogenetic studies are based on few fossil taxa. But the situation concerning modern elopomorphs is not more promising. The few phylogenetic studies including modern elopomorphs are also based on few taxa and they usually include one or two species of Elops (as representatives of elopiforms) and Albula vulpes (as representative of albulids). Furthermore, there are contradictory results concerning the monophyly of Elopomorpha and its content.
257
Osteoglossomorpha General overview and content The Superorder Osteoglossomorpha is represented by approximately 220 species and 29 genera placed in two orders, the Osteoglossiformes and Hiodontiformes (Nelson 2006). The osteoglossiforms comprise the suborders Osteoglossoidei and Notopteroidei. The suborder Osteoglossoidei, with five genera, is smaller than the Notopteroidei with 24 genera (Nelson 1994, 2006). The addition of fossils increases these numbers considerably, but research in progress makes it difficult to set a total number at present time. For instance, about 27 osteoglossomorph fossil genera (and more than 40 species) are currently known from Cretaceous to Neogene sites of China (Zhang 2002) and many more are still to be discovered. Additional fossil genera are known by new interpretations of some Cretaceous fishes formerly described as non-osteoglossomorphs but now interpreted as osteoglossomorphs (e. g., †Chanopsis lombardi: Taverne 1984, Poyato-Ariza 1996, Maisey 2000; †Paradercetis kipalaensis and †Kipalaichthys sekirsky: Taverne 1998, Maisey 2000) and by recent discoveries in Africa (e. g., Forey 1997, Cavin & Forey 2001, Murray & Wilson 2005). Three genera of marine osteoglossomorphs have been recently described from the Eocene of Europe (Taverne 1998), and one freshwater genus from the Paleocene of India (Kumar et al. 2005). New discoveries of fragmentary material in Spain and India, presently assigned to the Osteoglossomorpha mainly on the basis of the scales, open the possibility of new additional taxa. A rough account indicates than only the Asiatic fossil genera alone overpass the total number of extant genera. The oldest known records of fossil osteoglossomorphs (e. g., †Lycoptera) have mainly been recovered in China, Japan, Mongolia, and Siberia. A few other genera either referred to †Lycopteridae or Osteoglossomorpha incertae sedis were recovered in many localities in China (see Chang & Miao 2004). Formerly, the fishes were described as from Upper Jurassic deposits in China (e. g., Gaudant 1965, Chang & Chou 1977, Zhang et al. 1994). However, fossiliferous sites bearing †Lycoptera and some stem osteoglossomorphs previously interpreted as Late Jurassic in age are now interpreted as Early Cretaceous in age. For more information and references see the “Lycoptera-Peipiaosteus assemblage” or the “Jehol Fauna” in Chang & Jin (1996) and Chang & Miao (2004). Thus, the oldest known records of osteoglossomorphs are of Early Cretaceous age. They are younger than the oldest known records of elopomorphs of Late Jurassic age (see above). Overview on geographical distribution and environment Extant osteoglossomorphs are distributed in freshwater bodies mainly in tropical and semitropical regions of Africa, South America, Northern Australia, south-eastern Asia (Java and Sumatra), and North America (Berra 1981, 2001; Nelson 1994, 2006). A few of them are able to enter brackish water (Nelson 2006). In the past, osteoglossomorphs were also found in some regions of Europe (during the Paleogene), while they were broadly distributed in different regions of Asia such as China, Japan, Mongolia, and Siberia during the Cretaceous and Paleogene. Although most fossil osteoglossomorphs are thought to live in freshwater environments, a few have been recovered from marine deposits, e. g., in the upper Eocene London Clay (†Brychaetus; Woodward 1901, Taverne 1978), in the Eocene of Monte Bolca (e. g., †Monopteros; Taverne 1998), and in the upper Paleocene Fur Formation in Denmark (osteoglossids; Bonde 1987). Living osteoglossids occur on most continents of the Southern Hemisphere apart from the Indian subcontinent and Antarctica and have an apparently typical Gondwanan distribution that is the result of their extinction in North America, Europe, Central Asia, China, and India during the Eocene (Li 1997, Kumar et al. 2005). In contrast, notopteroids, with a higher number of species than osteoglossids, have a more restricted geographical distribution. Living notopterids occur in Africa, south-eastern Asia and India, while mormyrids and gymnarchids live in Africa (Nelson 1994, 2006); the last two families are poorly known from the fossil record. The living hiodontiforms occur in North America, while stem members of the group were present in China (†Plesiolycoptera: Li & Wilson 1999; †Jiaohichthys: Zhang 2002). Members of the extinct suborder Huashioidei are only known from freshwater localities in the Cretaceous of China (e. g., Li & Wilson 1999; Zhang 1998, 2002). Phylogenetic relationships among osteoglossomorphs Hilton (2003: 2-12) reviewed the history of the classification of the Osteoglossomorpha. To avoid repetitions, I invite the reader to refer to this publication. In the following section, I will analyze and evaluate 258
A
B
1 cm
1 cm
Fig. 8. Examples of some of the oldest known osteoglossomorphs from China. A, †Lycoptera davidi (LACM 4959122316). B, †Lycoptera fuxinensis (IVVP V 12437). (Photo of †L. fuxinensis courtesy of J.-Y. Zhang, IVVP).
the role of certain fossils to understanding the phylogeny within osteoglossomorphs (results that have been published mainly in the last 15 years). In contrast to recent elopomorph research, osteoglossomorphs have captivated the attention of many workers during the last decade or so. This research includes: (1) Re-assessment of the morphology and re-interpretation of numerous fossils. For instance: †Lycoptera from China (Figs. 8A,B, 9A; Zhang 2002), †Xixiaichthys (Figs. 9B; Zhang 2002, 2004); †Kuntulunia (Zhang 1998), †Tongxinichthys (Zhang & Jin 1999), †Eohiodon from China and North America (Shen 1989 and Li, Wilson & Grande 1997, respectively), †Phaerodus from North America (Li, Grande & Wilson 1997), †Monopteros, †Thrissopterus and †Foreyichthys from the Eocene of Monte Bolca (Taverne 1998), †Palaeonotopterus from the Lower Cretaceous of Morocco (Cavin & Forey 2001, Taverne & Maisey 1999, Taverne 2000), and †Singida from the Paleogen of Tanzania (Murray & Wilson 2005). (2) Re-assessment of the morphology of some extant forms, e. g., Hiodon (Hilton 2002). (3) New discoveries of fossil taxa, on the basis of otoliths (e. g., “genus Osteoglossidarum” from the Upper Cretaceous of U.S.A.; Nolf & Stringer 1996) and fragmentary and fairly complete specimens. Examples of the later are: †Cretophareodus (Li 1996), †Joffrichthys (Li & Wilson 1996a), †Chauliopareion (Murray & Wilson 2005), and †Taverneichthys (Kumar et al. 2005). The three aspects mentioned above have contributed enormously to the development of phylogenetic studies of osteoglossomorphs, as shown below.
259
A 1 cm
B
5 cm
Fig. 9. Restorations of some Cretaceous osteoglossomorphs from China, to show the extent of preserved morphological details. A, †Lycoptera davidi (after Arratia 1996b: fig. 2B). B, †Xixiaichthys tongxinensis (after Zhang 2002, but reversed).
(4) New phylogenetic hypotheses concerning relationships within osteoglossomorphs and proposals of new classifications in some cases. For instance, of the Osteoglossomorpha (Li 1994; Li & Wilson 1996b; Taverne 1998; Zhang 2002, 2006; Hilton 2003; Lavoué & Sullivan 2004), of the osteoglossomorphs with particular reference to the Hiodontiformes (Li & Wilson 1999), of the Hiodontidae (Li, Wilson & Grande 1997), of some heterotidinids (Li & Wilson 1996a), and of osteoglossids (e. g., Murray & Wilson 2005). Most of these studies dealt with computerized phylogenetic programs able to handle large matrices, with the exception of Taverne (1998). Taverne included numerous fossil and modern taxa in his study and his results largely agree with those by Li & Wilson (1996b) concerning the relationships of extant families. The position of most fossils, however, differs from that postulated in other studies. Taverne listed 344 characters, all of them interpreted as uniquely derived and supporting different phylogenetic levels; however, the homoplastic condition of many of them was not recognized or evaluated (Cavin & Forey 2001, pers. obs.). A recent analysis of five molecular markers of 12 living osteoglossomorphs (Lavoué & Sullivan 2004: fig. 3) provides a different hypothesis to those of Kumazawa & Nishida (2000) and Al-Mahrouki et al. (2001) also based on molecular evidence. Although the results of (Lavoué & Sullivan 2004: fig. 3) mostly agree with phylogenetic hypotheses based on morphological data (e. g., Hiodon occupies a basal position among living osteoglossomorphs), there is a major difference in the position of Pantodon that appears as the sister group of all living osteoglossomorphs excluding Hiodon. Consequently with their results Lavoué & Sullivan (2004: 183) propose that Pantodon buchholzi should be placed in its own family, the Pantodontidae, and that this new result should stimulate reappraisal of phylogenetically informative morphological characters within Recent and fossil osteoglossomorphs.
260
Chanos
Clupeoidea
Mormyroidea
Papyrocranus
Notopterus
Xenomystus
†Ostariostoma
Osteoglossum
Scleropages
†Singida
Pantodon
†Brychaetus muelleri
†Phareodus queenslandicus
†Phareodus encaustus
†Phareodus testis
Arapaima
Heterotis
†Sinoglossus
†Laeliichthys
†Tanichthys
Hiodon alosoides
Hiodon tergisus
†Hiodon consteniorum
†Eohiodon woodruffi
†Eohiodon rosei
†Yanbiania
†Lycoptera
Elopoidei
†Leptolepis
†Lycopteridae
D Hiodontiformes
Osteoglossoidei
Notopteroidei
E Osteoglossiformes
Clupeomorphs plus Euteleosts
C B Osteoglossomorpha A
Fig. 10. Hypothesis of phylogenetic relationships of some basal teleosts, especially of fossil and recent osteoglossomorphs (after Li & Wilson 1996). Nodes identifying taxa are added and branching of elopomorphs and osteoglossomorphs are shown in thicker lines to facilitate comparisons.
The phylogenetic hypotheses discussed below are based on fossil and living osteoglossomorphs and correspond to the approach listed as number 4 in the Introduction. However, the relation between the studied fossil and living taxa supporting these hypotheses is variable. More fossil taxa than living ones were considered in the studies by Li & Wilson (1996b, 1999) and Zhang (2002, 2006). In contrast, the number of fossil taxa is almost the half of the living ones in Hilton (2003). Another interesting point is that the selection of taxa makes comparisons between studies difficult because only Arapaima, †Eohiodon, Heterotis, Hiodon, †Lycoptera, Osteoglossum, Pantodon, †Phareodus, and Scleropages are included in all analyses. In other words, only three fossil and 6 living genera are common to the cladograms in Figs. 10-13. Certainly, access to the material is a fundamental difference in these works. For instance, some important Asiatic fossils are included only in studies by Chinese authors because that material has not yet been accessible to non-Asiatic researchers. A comparison between the four cladograms in Figs. 10-13 reveals some major agreements, but also major disagreements. The following are the common results: (1) The basal position of †Lycoptera (see Figs. 10-13 and Tables 3-6). Although †Lycoptera is interpreted as a basal osteoglossomorph, it is important to recall that most of the knowledge of the taxon is based on numerous incompletely known species. However, and unlike other fossil taxa, otoliths of †Lycoptera middendorfi were found in situ in the skull of several specimens from the Lower Cretaceous of Transbaikalia, Russia (Nolf 1985). 261
Table 3. Classification of Osteoglossomorpha (using the sequencing convention) by Li & Wilson 1996. Compare with Figure 10.
Table 4. Classification of Osteoglossomorpha (using the sequencing convention) based on Li & Wilson 1999: fig. 3. Compare with Figure 11.
Superorder Osteoglossomorpha
Superorder Osteoglossomorpha
Family †Lycopteridae (possibly also including †Tongxinichthys) †Lycoptera Order Hiodontiformes Family Hiodontidae †Yanbiania †Plesiolycoptera †Eohiodon Hiodon Order Osteoglossiformes †Tanichthys Suborder Osteoglossoidei Family Osteoglossidae Subfamily Osteoglossinae †Phareodus (including †Phareoides and †Brychaetus) Pantodon Unnamed genus group †Singida Osteoglossum Scleropages Subfamily Heterotidinae †Laeliichthys †Sinoglossus Heterotis Arapaima Suborder Notopteroidei Family †Ostariostomidae (possibly including †Thaumaturus) †Ostariostoma Family Notopteridae Xenomystus Notopterus Papyrocranus Family Mormyridae Family Gymnarchidae
†Tongxinichthys †Jiuquanichthys †Kuyangichthys †Lycoptera Order Hiodontiformes s. str. †Plesiolycoptera Family Hiodontidae †Yanbiania †Eohiodon Hiodon Order Osteoglossiformes Suborder Osteoglossoidei †Paralycoptera †Tanolepis Family Huashiidae Kuntulunia Huashia Family Osteoglossidae Subfamily Osteoglossinae †Phareodus (including †Phareoides and †Brychaetus) Pantodon Unnamed genus group †Singida Osteoglossum Scleropages Subfamily Heterotidinae †Laeliichthys †Sinoglossus Heterotis Arapaima
(2) The basal position of other Chinese forms that together with †Lycoptera are interpreted as stem osteoglossomorphs (e. g., †Tongxinichthys: Figs. 10-13; Tables 3-5; †Kuyangichthys and †Jiuquanichthys: Figs. 11, 13, Tables 4-5). The available studies indicate that a better knowledge of †Lycoptera is required to clarify the phylogenetic position of other taxa currently interpreted as lycopterids, osteoglossomorphs incertae sedis or stem osteoglossomorphs. (3) The recognition of the order Hiodontiformes including at least †Eohiodon and Hiodon. (4) The recognition of the order Osteoglossiformes and suborder Osteoglossoidei as high level taxa, but with an unclear content (compare Fig. 14 and Figs. 10, 11, 13). Hilton (2003; Fig. 13, Table 6) proposed a new usage of Osteoglossoidei, one comprising the families Osteoglossidae and Notopteridae, the later included in the suborder Notopteroidei of Nelson (1994) and Li & Wilson (1996b). Recently, Murray & Wilson (2005) published important new evidence concerning the Eocene genera Singida and †Chauliopareion from Mahenge site in Tanzania, Africa. The authors tested the phylogenetic position 262
†Huashiidae
†Lycopteridae
†Leptolepis coryphaenoides †Anaethalion knorri Elopoidei †Tongxinichthys microdus †Jinquanichthys liui †Kuyangichthys microdus †Lycoptera davidi †Plesiolycoptera daqigensis †Yanbiania wangqingica †Eohiodon rosei †Eohiodon woodruffi †Hiodon consteniorum Extant Hiodon spp. †Paralycoptera wui †Tanolepis ningjigouensis †Kuntulunia longipterus †Huashia gracilis †Laeliichthys †Joffrichthys symmetropterus †Sinoglossus lushanensis Heterotis Arapaima †Cretophareodus alberticus †Phareodus testis †Phareodus encaustus †Phareodus queenslandicus †Phareodus (Brychaetus) muelleri Pantodon †Singida Scleropages Osteoglossum †Thaumaturus †Ostariostoma Mormyroidea Notopteridae Clupeoidea Chanos Hypomesus (Osmeridae) Coregonus (Salmonidae) C1
E1
Clupeomorphs plus Euteleosts Osteoglossoidei
Notopteroidei
Hiodontiformes s.str. (Li & Wilson 1996)
E2 Osteoglossiformes E
D C Osteoglossomorpha B A
Fig. 11. Hypothesis of phylogenetic relationships of some fossil and recent osteoglossomorphs after Li & Wilson (1999). Nodes identifying taxa are added and branching of elopomorphs and osteoglossomorphs are shown in thicker lines to facilitate comparisons.
of the African taxa using the data sets of Li, Wilson & Grande (1997: fig. 7) and Hilton (2003: fig. 5) and concluded that the revised informations for †Singida jacksonoides causes the subfamily Osteoglossinae to be non-monophyletic. Here we have an example of how some fossils and their characters may impact previous phylogenetic hypotheses. Osteoglossomorphs or elopomorphs as the most primitive teleocephalans? One of the most important contributions to our understanding of teleostean relationships is that of Greenwood et al. (1966) where a tentative scheme of relationships among three main teleostean lineages are presented: Division I (Elopomorpha and Clupeomorpha), Division II (Osteoglossomorpha), and Division III (Ostariophysi, Protacanthopterygii, Atherinomorpha, and Acanthopterygii). I interpret Division I as the most primitive in this scheme. From this classic contribution, there has been much rearrangement of the main teleostean lineages. A historical review can be found in Hilton (2003). Later, Patterson & Rosen (1977) presented a hypothesis of phylogenetic relationships where the osteoglossomorphs are the sister group of elopomorphs plus more advanced teleosts. This hypothesis was mainly built on evidence from extant taxa, to which the fossils were added (approach no. 3 in the Introduction). A few years later, Nelson (1989) presented a summary tree that showed a fully resolved 263
Osteoglossiformes Osteoglossoidei (new usage) Osteoglossidae
Notopteridae
Mormyridae
Ostariostoma
Campylomormyrus
Gnathonemus
Petrocephalus
†Palaeonotopterus
Xenomystus
Papyrocranus
Chitala
Mormyrinae
Osteoglossum
†Singida
Scleropages
Osteoglossinae
Phareodus
Arapaima
Heterotis
†Joffrichthys
Hiodon
†Eohiodon
†Lycoptera
Heterotinae
Pantodon
Hiodontidae
A1 G F E D C B A
Fig. 12. Hypothesis of phylogenetic relationships of osteoglossomorphs after Zhang (2006).
scheme of the major teleostean lineages that was thought to be solved at the base, but contained problems at the top. Nevertheless, the inclusion of new high quality fossil evidence, large and refined data sets of characters and larger data set of taxa, of more refined morphological studies, new molecular data, and assistance of phylogenetic programs are showing that many problems remain at different teleostean nodes, as well as different nodes within Actinopterygii (Arratia 2004, Stiassny et al. 2004, Nelson 2006). One example is the phylogenetic position of elopomorphs and osteoglossomorphs, for which different hypotheses are available (Fig. 15A-E): Osteoglossomorpha as sister group of Elopomorpha + more advanced teleosts (Fig. 14A). This sistergroup relationship is mainly or only based on extant forms (approach no. 1 listed in the Introduction: e. g., Forey et al. 1996, Inoue et al. 2001, and Wang et al. 2003, based on molecular evidence of recent species) or the fossils are mapped on the cladogram (approach no. 3: Patterson & Rosen 1977). For example, all fossils above the level of †Tharsis dubius in Patterson & Rosen (1977: fig. 54) were added to the phylogenetic hypothesis, e. g., †Anaethalion and †‘Anaethalion’ vidali at the base of Elopomorpha and †Lycopteridae within Osteoglossomorpha. No character, in the list provided by the authors, supports these fossil assignments (see details in Arratia 1998). Elopomorpha as sister group of Osteoglossomorpha + more advanced teleosts (Fig. 14B). This sister-group relationship has been consistently demonstrated when fossils and recent fishes are included in the phylogenetic analysis. So far I know, the only exception is Li & Wilson (1996; Fig. 14D), in contrast to other studies by the same authors (e. g., Li 1994, 1996; Li & Wilson 1999; compare Fig. 10 and Fig. 11). Elopomorphs are non-monophyletic after Filleul (2000b; Fig. 7), with Elopiformes occupying a more basal position than osteoglossomorphs, but Albuliformes more advanced than osteoglossomorphs. All these studies correspond to approach no. 4 listed in the Introduction. Recent studies based on molecular evidence (approach no. 1 listed in the Introduction) also have reached similar results concerning the elopomorphs as the most plesiomorphic group among extant teleosts (e. g., Alfaro et al. 2009, Santini et al. 2009). It is important to remark that all these studies are based on different sets of characters but several taxa and some characters are common to all of them. Elopomorpha + Osteoglossomorpha as sister group of more advanced teleosts (Fig. 14C). This sistergroup relationship was obtained in a study of molecular evidence of only recent species (approach no. 1 listed in the Introduction) (Lê et al. 1993, Broughton 2010). 264
Table 5. Classification of Osteoglossomorpha (using the sequencing convention) by Zhang 2006. Compare with Figure 12.
Table 6. Classification of Osteoglossomorpha (using the sequencing convention) after Hilton 2003. Compare with Figure 13.
Superorder Osteoglossomorpha
Superorder Osteoglossomorpha
†Jinanichthys †Lycoptera †Kuyangichthys †Jiuquanichthys †Tongxinichthys †Xixiaoichthys †Kuntulunia †Huashia Order Hiodontiformes Family Hiodontidae †Eohiodon Hiodon †Jiaohichthys Order Osteoglossiformes †Thaumaturus Suborder Osteoglossoidei Family Osteoglossidae Unnamed Group †Paralycoptera Subfamily Osteoglossinae Osteoglossum Scleropages Pantodon Subfamily Heterotidinae Heterotis Arapaima †Sinoglossus Subfamily Phareodontinae †Phareodus (†Brychaetus) Singida Suborder Notopteroidei Family Mormyridae Family Ostariostomidae Ostariostoma Family Notopteridae Notopterus Papyrocranus Xenomystus
Osteoglossomorpha incertae sedis Family †Lycopteridae †Lycoptera Order Hiodontiformes Family Hiodontidae †Eohiodon Hiodon Order Osteoglossiformes plesion †Ostariostoma Osteoglossifomes incertae sedis †Palaeonotopterus Family Mormyridae Petrocephalus Gnathonemus Campylomormyrus Suborder Osteoglossoidei (new usage, Notopteridae + Osteoglossidae) Family Notopteridae Chitala Papyrocranus Xenomystus Family Osteoglossidae plesion †Joffrichthys Subfamily Heterotidinae Heterotis Arapaima Subfamily Osteoglossinae Osteoglossinae incertae sedis †Phareodus Osteoglossinae incertae sedis Pantodon Osteoglossinae incertae sedis †Singida Osteoglossum Scleropages
Non monophyletic Elopomorpha and Osteoglossomorpha (Fig. 14E). Elopomorpha and Osteoglossomorpha as non monophyletic taxa and with unresolved phylogenetic positions were the conclusions reached by Filleul & Lavoué (2001), on the basis of molecular evidence. Elopomorphs as a non monophyletic group and in an unresolved position with osteoglossomorphs and clupeopeocephalans were the results of Li et al. (2008) based also on molecular evidence. Thus, while the morphological evidence of recent taxa has given one topology (Fig. 14A), the morphological evidence of fossil and recent taxa has given another (Fig. 14B), but the molecular data of modern taxa have given different results (compare Fig. 14A, C, and E). Molecular studies and studies based only on recent teleosts are based only on terminal taxa of groups that may have a small representation today in comparison to the past (e. g., elopiforms, osteoglossomorphs). Consequently, inevitable questions arise: 265
†Jiuquanichthys
†Lycoptera
†Kuyangichthys
†Jinanichthys
†Tongxinichthys
†Xixiaichthys
†Kuntulunia
†Huashia
Hiodon
†Jiaohichthys
†Eohiodon
†Thaumaturus
†Ostariostoma
Xenomystus
Papyrocranus
Notopterus
Mormyridae
†Paralycoptera
Pantodon
Scleropages leichardti
Scleropages formosus
Osteoglossum
†Singida
†Phareodus testis
†Phaerodus queenslandicus
†Phareodus encaustus
†Brychaetus
†Sinoglossus
Heterotis
Arapaima
†Leptolepis
F E
I
H G D C B A
Fig. 13. Hypothesis of phylogenetic relationships of some fossils and recent osteoglossomorphs after Hilton (2003).
Do the fossils play a role in phylogenetic hypotheses of relationships of teleocephalans (= containing modern teleostean major lineages)? Should the characters present in stem-lineage members be excluded from phylogenetic analyses involving their recent representatives, and consequently possible primitive conditions? Arratia’s (1997, 1999; Fig. 15A) studies of fossil and recent basal teleosts and of their possible sister group demonstrated that elopiforms are the basal group among teleocephalans. This result does not change when the composition of the outgroup is changed. Arratia’s (1999: figs. 19-21) studies revealed that changes in the composition of the outgroup did not affect the position of the elopiforms versus osteoglossomorphs. However, when all the fossils are deleted (in the ingroup and outgroup) from the analysis there is a change in the topology and osteoglossomorphs are now in the most basal position (see Fig. 15B) and they do not form a monophyletic group. Certainly, there is a significant change in the interpretation of characters and of their distribution when the data sets include fossil and recent forms or only recent or fossil forms. Both elopomorphs and osteoglossomorphs have numerous recent members, but also have numerous well-preserved stem-lineage representatives with different quantitative and qualitative expressions (states) of the synapomorphies of each taxon (as defined based on recent members of each taxon). Thus, understanding the different states for each character discovered in the stem-lineage representatives may change the coding of some of them as well as interpretations on character distribution. 266
Li & Wilson (1996b)
E
Lê et al. (1993) Broughton (2010)
More advanced teleosts
Osteoglossiformes
C
Anguilliformes
Elopiformes
More advanced teleosts
Osteoglossomorpha
Elopomorpha
More advanced teleosts Notacanthus
Albula
Arratia (1991, 1996b, 1999, 2000) Li (1994, 1996); Shen (1996) Zhang (1998); Li & Wilson (1999) Alfaro et al. (2009) Santini et al. (2009)
Hiodon
More advanced teleosts
Osteoglossomorpha
B
Elopomorpha
A
D
Osteoglossomorpha
Elopomorpha
More advanced teleosts
Elopomorpha
Osteoglossomorpha
Patterson & Rosen (1977) Forey et al. (1996) Inoue et al. (2001, 2004)
Filleul & Lavoué (2001)
Fig. 14. Hypotheses of phylogenetic relationships of teleosts showing changes in the position of elopomorphs and osteoglossomorphs when using different sets of taxa (fossil and/or recent forms) and characters (e. g., morphological or molecular data).
When the phylogenetic analysis includes extant taxa in the ingroup and Amia or Amia plus Lepisosteus as outgroup, the osteoglossomorphs are the sister group of elopomorphs plus more advanced teleosts (Figs. 14A, 15B). However, when fossils are included in the ingroup and outgroup, the elopomorphs are in the basal position (Fig. 14B). Why is this? Not only the recent members of the ingroup have wellpreserved fossil stem-lineage representatives, but also the outgroups, so that their addition to the analysis changes the polarization of characters. We would expect to have the same results in both phylogenetic analyses, and because of this expectation it is believed that the evolutionary history of a group can be extrapolated from its recent taxa alone. Probably, this is an acceptable approach when the history of a group (e. g., cypriniforms) involves a few million years, but apparently our expectations and interpretations of a “linear” evolution or of rates of evolutionary changes may prove seriously wrong when we are dealing with groups like the elopomorphs, with more than 150 million years of evolutionary history, or the osteoglossomorphs with about 140 million years of history.
267
†Allothrissops †Thrissops †Pachythrissops †Anaethalion Elops Megalops Chanos †Gordichthys Opsariichthys †Tischlingerichthys Denticeps Engraulis †Santanaclupea †Diplomystus †Erichalcis †Leptolepides haerteisi †Leptolepides sprattiformis †Orthogonikleithrus leichi †Orthogonikleithrus hoelli †Humbertia Esox Umbra Oncorhynchus Thymallus Hiodon Heterotis †Lycoptera †Ascalabos †Domeykos †Luisichthys †Protoclupea †Varasichthys †Tharsis †Leptolepis coryphaenoides †Pholidophorus bechei †Dapedium †Hypsocormus †Pachycormus †Aspidorhynchus †Belonostomus †Vinctifer Lepisosteus †Obaichthys †Mesturus †Prohalecites Amia calva †Amia pattersoni †Watsonulus
Teleostei
G2
F2
L2
K2
N3
K
N4
O1 O2 O
N2
L1
K1
G1
F1
L
4b
D3
H2 H1
D2
1a
3d
3b 3a
D1
N1
J
3e
3c
4a
N M I H G
F E
A
D C B A 4
E2
E1
F2
F1
2 1
C1
F
E D
C B
B
Lepisosteus
Amia calva
Heterotis
Hiodon
Megalops
Elops
3
Thymallus
Oncorhynchus
Umbra
Esox
Engraulis
Denticeps
Chanos
Opsariichthys
Teleostei
A
Fig. 15. Hypothesis of phylogenetic relationships of some teleosts (after Arratia 1999). Extant osteoglossomorphs are represented by thicker lines to facilitate comparisons. A, including fossil and recent taxa. B, including only the recent taxa.
Final comments and conclusions “Very few studies have succeeded in including fossils to overturn a theory of relationships based on modern taxa. Some that I know in the fish world are Arratia (1997), Murray & Wilson (1999) and Wilson (1992).” (Forey 2004: 169). [The quotation of Arratia (1997; Fig. 14B) corresponds to the phylogenetic study of fossil and recent basal teleosts that overturn the hypothesis of phylogenetic relationships based on extant forms where the osteoglossomorphs stand at the base of the modern teleostean lineages (see Fig. 14A).] Forey’s statement is correct; however, he did not address the particular point that only very 268
few fish studies include both fossil and recent members in phylogenetic studies of Teleostei or within teleostean subgroups. Thus, the “lack of success” of fossils is not due to their condition of preservation – and consequently as to how much morphological information they may provide – but to the fact that they are usually not included in phylogenetic studies. For instance, the important contribution of Johnson & Patterson (1996) concerning the phylogenetic relationships of basal euteleosts is based solely on modern representatives despite the fact that very old fossil euteleosts have been known in the literature for a long period of time. The siluriforms are another example; the available hypotheses of relationships within siluriforms (e. g., Grande 1987, Arratia 1992) consider the modern Diplomystidae at the basal node and include the fossil †Hypsidoris from the Paleogene of North America. Would it be a surprise if these hypotheses are overturned when some of the Late Cretaceous-Paleocene catfishes from Bolivia, with very different morphological patterns, are included in phylogenetic studies of Siluriformes? Forey (e. g., 2004) identified five aspects to illustrate the strengths of fossils: (1) Ability to break up long branches (dependent on being able to assign a fossil to a Recent taxon on the basis of synapomorphies). Both elopomorphs and osteoglossomorphs are groups with long evolutionary history and their oldest representatives share with modern members the elopomorph and osteoglossomorph synapomorphies, respectively. (2) Stabilization of theories of character evolution. (3) Elucidation of the sequential steps involved in the evolution of a complex character. Aspects 2 and 3 are problematic because many morphological characters of recent teleosts are incompletely known and their homology is assumed. If the fossils are generally excluded in phylogenetic studies of teleosts, hardly the evolutionary steps of certain characters can be understood. (4) Arbitration between equally parsimonious theories based on Recent taxa [morphological data] or molecular data. Phylogenetic studies of teleosts including both fossil and recent elopomorphs and osteoglossomorphs together with other fossil basal teleosts and more advanced teleosts produce a different theory than studies based on only recent species. (5) Acting as outgroups for modern taxa. There are numerous examples of this role in the literature. However, this role can be misleading when fossils that are phylogenetically very distant of the group under study are used as outgroups. As shown in sections above, the position of elopomorphs change when only recent taxa or fossil plus recent taxa are included in phylogenetic studies of teleosts. This result is not due to the age of the fossils (e. g., oldest records of elopomorphs are older than the oldest records of osteoglossomorphs); age and primitiveness are not synonymous, and not every fossil is phylogenetically important. If elopomorphs are more primitive than other teleostean lineages this is due to the characters and/or combination of certain characters present in the members of the group. It follows that the inclusion or exclusion of fossils in the phylogenetic analysis of basal teleosts or not, biases the results (discussed by Arratia 2004), and when we select one or another set of taxa, inevitably, we have already biased the results (as above illustrated). It is clear to me that we have accumulated many data – e. g., morphological and molecular – but we still are unable to interpret them properly. It is a major task to interpret the sequence of evolutionary changes in the characters distribution of fossil and recent representatives of supposedly monophyletic taxa within the time dimension. Finally, as I have pointed out earlier (Arratia 1998, 2001, 2004), when studying the evolutionary history of a group, fossils should be included, otherwise we are only dealing with a taxon at one-time level (0 time). Obviously, the molecular characteristics and the genome as well as the morphological characters of this recent group are very important, but they alone will tell us little about the evolutionary history of the group. Acknowledgements I am grateful to Jiangyong Zhang (Beijing) for permission to use information contained in his Ph.D. dissertation and to use some photographs of certain Chinese specimens; to Arnaud Filleul (Paris) for permission to use one of his published figures; to Matthias Mäuser (Bamberg) for letting me identify the fish material recently collected by him in Wattendorf; to Mr. J.-P. Mendau (Berlin) for his help with the illustrations based on my original drawings; to Mrs. C. Radke (Berlin) for the photographs illustrated in Figure 1; to Pingfu Chen (Kansas) for his assistance with the computerized preparation of the illustrations; and to Günter Viohl and Martina Kölbl-Ebert (Eichstätt) for permission to study material under her care. Sincerest thanks to Eric Hilton, Joseph Nelson, Hans-Peter Schultze, and John Chorn for their helpful comments that greatly improved the manuscript. This is a contribution of grants NSF EF 0431262 and DFG AR 275/10-1-1 and DFG KA 1525/3-1.
269
References Alfaro, M. E., Santini, F., Brock, C., Alamillo, H., Dornburg, A., Rabosky, D. L., Carnevale, G. & Harmon, L. (2009): Nine exceptional radiations plus high turnover explain species diversity in jawed vertebrates. – Proc. Nat. Acad. Sci. 106 (32): 13410-13414. Al-Mahrouki, A. A., Irwin, D. M., Graham, L. C. & Youson, J. H. (2001): Molecular cloning of preproinsulin cDNAs from several osteoglossomorphs and a cyprinid. – Mol. Cell. Endocrinol. 174: 51-58. Applegate, S. P. (1970): The vertebrate fauna of the Selma Formation of Alabama. – Fieldiana: Geol. Mem. 3: 383-433. Arratia, G. (1987): Anaethalion and similar teleosts (Actinopterygii, Pisces) from the Late Jurassic (Tithonian) of southern Germany and their relationships. – Palaeontogr. A 200: 1-44. – (1991): The caudal skeleton of Jurassic teleosts; a phylogenetic analysis. – In: Chang M.-M., Liu Y.-H. & Zhang G.-R. (eds.): Early Vertebrates and Related Problems in Evolutionary Biology; p. 249-340; Beijing (Science Press). – (1992): Development and variation of the suspensorium of primitive catfishes (Teleostei: Ostariophysi) and their phylogenetic relationships. – Bonner zool. Monogr. 32: 1-140. – (1994): Phylogenetic and paleobiogeographic relationships of the varasichthyid group (Teleostei) from the Late Jurassic of Central and South America. – Rev. Geol. Chile 21: 119-165. – (1996a): The Jurassic and the early history of teleosts. – In: Arratia, G. & Viohl, G. (eds.): Mesozoic Fishes – Systematics and Paleoecology; p. 243-260; München (Verlag Dr. Friedrich Pfeil). – (1996b): Reassessment of the phylogenetic relationships of certain Jurassic teleosts and their implications on teleostean phylogeny. – In: Arratia, G. & Viohl, G. (eds.): Mesozoic Fishes – Systematics and Paleoecology; p. 219-242; München (Verlag Dr. Friedrich Pfeil). – (1997): Basal teleosts and teleostean phylogeny. – Palaeo Ichthyologica 7: 1-168. – (1998): Basal teleosts and teleostean phylogeny: response to C. Patterson. – Copeia 1998 (4): 1-168. – (1999): The monophyly of Teleostei and stem-group teleosts. Consensus and disagreements. – In: Arratia, G. & Schultze, H.-P. (eds.): Mesozoic Fishes 2 – Systematics and Fossil Record; p. 265-334; München (Verlag Dr. Friedrich Pfeil). – (2000): Remarkable teleostean fishes from the Late Jurassic of southern Germany and their phylogenetic relationships. – Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 3: 137-179. – (2001): The sister-group of Teleostei: consensus and disagreements. – J. Vert. Paleontol. 21 (4): 767-773. – (2004): Mesozoic Halecostomes and the early radiation of teleosts. – In: Arratia, G. & Tintori, A. (eds.): Mesozoic Fishes 3 – Systematics, Paleoenvironments and Biodiversity; p. 279-316; München (Verlag Dr. Friedrich Pfeil). – (2008a): The varasichthyids and other crossognathiform fishes, and the break-up of Pangaea. – In: Cavin, L., Longbottom, A. & Richter, M. (eds.): Fishes and the break-up of Pangaea; Special Publication 295: 71-92. Geol. Soc. London. – (2008b): Actinopterygian postcranial skeleton with special reference to the diversity of fin ray elements, and the problem of identifying homologies. – In: Arratia, G., Schultze, H.-P. & Wilson, M. V. H. (eds.): Mesozoic Fishes 4 – Homology and Phylogeny; p. 49-101, München (Verlag Dr. Friedrich Pfeil). Arratia, G. & Schultze, H.-P. (eds.) (1999): Mesozoic Fishes – Systematics and Fossil Record. – 604 pp.; München (Verlag Dr. Friedrich Pfeil). Arratia, G., Schultze, H.-P. & Casciotta, J. (2001): Vertebral column and associated elements in Dipnoans and comparison with other fishes: Development and homology. – J. Morphol. 250: 101-172. Arratia, G. & Tintori, A. (eds.) (2004): Mesozoic Fishes – Systematics, Paleoenvironments and Biodiversity. – 649 pp; München (Verlag Dr. Friedrich Pfeil). Arratia, G. & Viohl, G. (eds.) (1996): Mesozoic Fishes – Systematics and Paleoecology. – 575 pp.; München (Verlag Dr. Friedrich Pfeil). Ax, P. (1987): The Phylogenetic System. The Systematization of Organisms on the Basis of their Phylogenesis. – 340 pp.; Chichester, New York, Brisbane, Toronto, Singapore (J. Wiley and Sons). Belouze, A. (2002): Compréhension morphologique et phylogénétique des taxons actuels et fossils rapports aux Anguilliformes (“poissons”, téléostéens). – Docum. Lab. Géol. Lyon 158: 401 pp. Belouze, A. & Gayet, M. (1999): Première observation de la presence simultanée d’ une ceinture pectorale complete et de nageoires pelviennes chez des Anguilliformes (Teleostei) du Crétacé du Liban. – C. R. Acad. Sci. Paris, Earth & Plan. Sci. 329: 683-688. Belouze, A., Gayet, M. & Atallah, C. (2003a): Les premiers Anguilliformes: I. Révision des genres cénomaniens Anguillavus Hay, 1903 et Luenchelys nov. gen. – Geobios 36: 241-273. – (2003b): Les premiers Anguilliformes. II. Paraphylie du genre Urenchelys Woodward, 1900 et relations phylogénétiques. – Geobios 36: 351-378.
270
Berra, T. M. (1981): An atlas of distribution of fresh water fish families of the world. – 197 pp.; Lincoln and London (University of Nebraska Press). – (2001): Freshwater Fish Distribution. – 604 pp.; San Diego (Academic Press). Blot, J. (1978): Les Apodes fossils du Monte Bolca. – Studi e Ricerche sui giacimenti terziari di Bolca, 3 (1): 1-260, Verona. – (1980): La faune ichthyologique des gisements du Monte Bolca (Province de Vérona, Italie). – Bull. Mus. Natl. Hist. Natur., Paris (4), 2, C: 339-396. – (1984): Les Apodes fossils du Monte Bolca. – II. – Studi e Ricerche sui giacimenti terziari di Bolca, 4: 61-238, Verona. Blum, S. (1991): Brannerion Jordan, 1919. – In: Maisey, J. G. (ed.): Santana Fossils. An Illustrated Atlas; p. 218237; Neptune City (T.F.H. Publications, Inc.). Bonde, N. (1987): Moler, its origin and its fossils especially fishes. – 53 pp.; Skamol (Nykobing Mors). Broughton, R. E. (2010): Phylogeny of teleosts based on mitochondrial genome sequences. – In: Nelson, J. S., Schultze, H.-P. & Wilson, M. V. H. (eds.): Origin and Phylogenetic Interrelationships of Teleosts; p. 61-76, München (Verlag Dr. Friedrich Pfeil). Brundin, L. (1966): Transantarctic relationships and their significance, as evidence by chironomid midges with a monograph of the subfamilies Podonominae and Aphroteniinae and the Austral Heptaginae. – K. Svenka VetenskAkad. Handl., ser. 4, 11 (1): 1-471. – (1968): Application of the phylogenetic principles in systematics and evolutionary theory. – In: Ørvig, T. (ed.): Current Problems of Lower Vertebrate Phylogeny; Nobel Symposium 4: 473-495. Cavin, L. & Forey, P. L. (2001): Osteology and systematic affinities of Palaeonopterus greenwoodi Forey 1997 (Teleostei: Osteoglossomorpha). – Zool. J. Linn. Soc. 133: 25-52. Chang, M.-M.(1999): ‘Mid’-Cretaceous fish fauna from northeast China. – In: Arratia, G. & Schultze, H.-P. (eds.): Mesozoic Fishes 2 – Systematics and Fossil Record; p. 469-480; München (Verlag Dr. Friedrich Pfeil). Chang, M.-M. & Chou, C.-C (1977): On Late Mesozoic fossil fishes from Zhejiang Province, China. – Mem. Inst. Vert,. Palaeontol., Palaeoanthropol., Acad. Sinica 12: 1-59. (In Chinese with English summary.) Chang, M.-M. & Jin, F. (1996): Mesozoic Fishes of China. – In: Arratia, G. & Viohl, G. (eds.): Mesozoic Fishes – Systematics and Paleoecology; p. 461-478; München (Verlag Dr. Friedrich Pfeil). Chang, M.-M. & Miao, D.-S. (2004): An overview of Mesozoic Fishes in Asia. – In: Arratia, G. & Tintori, A. (eds.): Mesozoic Fishes 3 – Systematics, Paleoenvironments and Biodiversity; p. 535-563; München (Verlag Dr. Friedrich Pfeil). Cockerel, T. D. A. (1933): The antiquity of Albula. – Copeia: 226. Filleul, A. (2000a): Baugeichthys caeruleus, gen. et sp. nov., a new albuliform fish from the Hauterivian of the Massif des Bauges (France). – J. Vert. Paleontol. 20: 637-644. – (2000b): Analyse critique des synapomorphies des Elopomorphes et analyse phylogénétique du groupe. – Cybium 4 (3): 75-83. Filleul, A. & Lavoué, S. (2001): Basal teleosts and the question of elopomorph monophyly. – C. R. Acad. Sci., Paris, Sci de la vie 324: 393-399. Forey, P. L. (1973): A revision of the elopiform fishes, fossil and recent. – Bull. Brit. Mus. (Natur. Hist.), Geol., Suppl. 10: 4-222. – (1997): A Cretaceous notopterid (Pisces: Osteoglossomorpha) from Morocco. – South African J. Sci. 93: 564-569. – (2004): Systematics and Paleontology. – In: Williams, D. M. & Forey, P. L. (eds.): Milestones in Systematics; p. 149-180. Boca Raton, London, New York, Washington, D.C. (CCR Press). Forey, P., Littlewood, D. T. J., Ritchie, P. & Meyer, A. (1996): Interrelationships of elopomorphs fishes. – In: Stiassny, M. L. J., Parenti, L. R. & Johnson G. D. (eds.): Interrelationships of Fishes; p. 175-191; San Diego (Academic Press). Forey, P. & Maisey, J. G. (2010): Structure and relationships of †Brannerion (Albuloidei), an Early Cretaceous teleost from Brazil. – In: Nelson, J. S., Schultze, H.-P. & Wilson, M. V. H. (eds.): Origin and Phylogenetic Interrelationships of Teleosts; p. 183-218, München (Verlag Dr. Friedrich Pfeil). Fürsich, F. T., Mäuser, M., Schneider, S. & Werner, W. (2005): Sedimentology and palaecology of laminated limestones from the Kimmeridgian of the northern Franconian Alb (southern Germany). – Zitteliana B26: 13. (Abstract of 4th Intl. Symp. Lithographic Limestone and Plattenkalk, Eichstätt and Solnhofen, 2005.) Gaudant, J. (1965): Lycoptera wangi nov. sp. (Poisson Téléostéen) dans le Jurassique des environs de Hengshan (Shensi, Chine). – C. R. Soc. Géol., France 10: 337-339. – (1968): Contribution à une révision des Anaethalion de Cerin (Ain). – Bull. Bur. Rech. Geol. miner. Paris (2), 4, 1: 95-115. Gauthier, J., Kluge, A. G. & Rowe, T. (1988): Amniote phylogeny and the importance of fossils. – Cladistics 4, 2: 105-209. Grande, L. (1987): Redescription of Hypsidoris farsonensis (Teleostei: Siluriformes), with a reassessment of its phylogenetic relationships. – J. Vert. Paleontol. 7 (1): 24-54.
271
Greenwood, P. H., Rosen, D. E., Weitzman, S. H. & Myers, G. S. (1966): Phyletic studies of teleostean fishes with a provisional classification of living forms. – Bull. Amer. Mus. Natur. Hist. 131: 339-456. Hennig, W. (1966): Phylogenetic Systematics. – Urbana, Chicago, London (University of Illinois Press). Hilton, E. (2002): Osteology of the extant North American fishes of the genus Hiodon Lesueur, 1818 (Teleostei, Osteoglossomorpha): Hiodontiformes). – Fieldiana: Zool., n. ser. 100: 1-142. – (2003): Comparative osteology and phylogenetic systematics of fossil and living bony-tongue fishes (Actinopterygii, Teleostei, Osteoglossomorpha). – Zool. J. Linn. Soc. 137: 1-100. Inoue, J. G. & Miya, M. (2001): Phylogeny of the basal teleosts, with special reference to the Elopomorpha. – Jpn. J. Ichthyol. 48 (2): 75-91. Inoue, J. G., Miya, M., Tsukamoto, K. & Nishida, M. (2004): Mitogenomic evidence for the monophyly of elopomorph fishes (Teleostei) and the evolutionary origin of the leptocephalous larva. – Mol. Phylogenetics Evol. 32 (1): 274-286. Ishiguro, N. B., Miya, M. & Nishida, M. (2003): Basal euteleostean relationships: a mitogenomic perspective on the phylogenetic reality of the ‘Protacanthopterygi’. – Mol. Phylogenetics Evol. 27: 476-488. Johnson, G. D. & Patterson, C. (1996): Relationships of lower euteleostean fishes. – In: Stiassny, M. L. J., Parenti, L. R. & Johnson, G. D. (eds.): Interrelationships of Fishes; p. 251-332. San Diego (Academic Press). Kumar, K., Rana, R. S. & Paliwal, B. S. (2005): Osteoglossid and lepisosteid fish remains from the Paleocene Palana Formation, Rajasthan, India. – Palaeontology 48 (6): 1187-1209. Kumazawa, Y. & Nishida, M. (2000): Molecular phylogeny of osteoglossoids: a new model for Gondwanian origin and plate tectonic transportation of the Asian arowana. – Mol. Biol. Evol. 17: 1869-1878. Lavoué, S. & Sullivan, J. P. (2004): Simultaneous analysis of five molecular markers provides a well-supported phylogenetic hypothesis for the living bony-tongue fishes (Osteoglossomorpha: Teleostei). – Mol. Phylog. Evol. 33: 171-185. Lê, H. L. V., Lecointre, G. & Peraso, R. (1993): A 28S rRNA-based phylogeny of the Gnathostomes: First steps in the analysis of conflict and congruence with morphologically based cladograms. – Mol. Phylogen. Evol. 2: 31-51. Li, G.-Q. (1994): New osteoglossomorphs (Teleostei) from the Upper Cretaceous and Lower Tertiary of North America and their phylogenetic significance. – Unpublished Ph.D. dissertation. 290 pp. University of Alberta, Edmonton, Canada. – (1996): A new species of Late Cretaceous osteoglossid (Teleostei) from the Oldman Formation of Alberta, Canada, and its phylogenetic relationships. – In: Arratia, G. & Viohl, G. (eds.): Mesozoic Fishes – Systematics and Paleoecology; p. 285-298; München (Verlag Dr. Friedrich Pfeil). – (1997): Notes on the historical biogeography of the Osteoglossomorpha. – Proc. 30th Intl. Geol. Congr. 12: 54-66. Li, G.-Q., Grande, L. & Wilson, M. V. H. (1997): The species of †Phaerodus (Teleostei: Osteoglossidae) from the Eocene of North America and their phylogenetic relationships. – J. Vert. Paleontol. 17: 487-505. Li, G.-Q. & Wilson, M. V. H. (1996a): The discovery of Heterotidinae (Teleostei: Osteoglossidae) from the Paleocene Paskapoo Formation of Alberta, Canada. – J. Vert. Paleontol. 16: 198-209. – (1996b): Phylogeny of the Osteoglossomorpha. – In: Stiassny, M. L. J., Parenti, L. R. & Johnson, G. D. (eds.): Interrelationships of Fishes; p. 163-174; San Diego (Academic Press). – (1999): Early divergence of Hiodontiformes sensu stricto in East Asia and phylogeny of some Late Mesozoic teleosts from China. – In: Arratia, G. & Schultze, H.-P. (eds.): Mesozoic Fishes 2 – Systematics and Fossil Record; p. 369-384; München (Verlag Dr. Friedrich Pfeil). Li, G.-Q., Wilson, M. V. H. & Grande, L. (1997): Review of Eohiodon (Teleostei: Osteoglossomorpha) from the western North America, with a phylogenetic reassessment of Hiodontidae. – J. Paleontol. 71: 1109-1124. Li, C., Lu, G. & Orti, G. (2008): Optimal data partitioning and a test case for ray-finned fishes (Actinopterygii) based on ten nuclear loci. – Syst. Biol. 57: 519-539. Løvtrup, S. (1977): The Phylogeny of Vertebrata. – London (J. Wiley). Maisey, J. G. (1986): Heads and tails: a chordate phylogeny. – Cladistics 2: 20-256. – (ed.) (1991): Santana Fossils. An Illustrated Atlas. – 496 pp.; Neptune City (T.F.H. Publications, Inc.). – (2000): Continental break up and the distribution of fishes of Western Gondwana during the Early Cretaceous. – Cretaceous Research 21: 281-314. Maisey, J. G. & Blum, S. (1991): Paraelops Silva Santos, 1971. – In: Maisey, J. G. (ed.): Santana Fossils. An Illustrated Atlas; p. 238-247; Neptune City (T.F.H. Publications, Inc.). Murray, A. M. & Wilson, M. V. H. (1999): Contributions of fossils to the phylogenetic relationships of the percopsiform fishes (Teleostei: Paracanthopterygii): order restored. – In: Arratia, G. & Schultze, H.-P. (eds.): Mesozoic Fishes 2 – Systematics and Fossil record; p. 369-384; München (Verlag Dr. Friedrich Pfeil). – (2005): Description of a new Eocene osteoglossid fish and additional information on Singida jacksonoides Greenwood and Patterson 1967 (Osteoglossomorpha), with an assessment of their phylogenetic relationships. – Zool. J. Linn. Soc. 144: 213-228.
272
Nelson, G. J. (1969): Origin and diversification of teleostean fishes. – Ann. New York Acad. Sci. 167: 18-30. – (1978): Ontogeny, phylogeny, paleontology and the biogenetic law. – Syst. Zool. 27: 324-345. – (1989): Phylogeny of major fish groups. – In: Fernholm, B., Bremer, K., Brundin, L., Jornvall, H., Rutberg, L. & Wanntorp, H.-E. (eds.): The Hierarchy of Life; p. 315-336; Amsterdam (Elsevier Science). Nelson, J. (1994): Fishes of the World. – Third ed., xvii + 600 pp.; New York (J. Wiley & Sons, Inc.). – (2006): Fishes of the World. – Fourth ed., xix + 601 pp.; New York (J. Wiley & Sons, Inc.). Nolf, D. (1985): Otolithi Piscium. – In: Schultze, H.-P. (ed.): Handbook of Paleoichthyology, Volume 10, 145 pp.; Stuttgart, New York (Gustav-Fischer-Verlag). Nolf, D. & Stringer, G. L. (1996): Cretaceous fish otoliths – a synthesis of the North American record. – In: Arratia, G. & Viohl, G. (eds.): Mesozoic Fishes – Systematics and Paleoecology; p. 433-459; München (Verlag Dr. Friedrich Pfeil). Obermiller, L. E. & Pfeiler, E. (2003): Phylogenetic relationships of elopomorph fishes inferred from mitochondrial ribosomal DNA sequences. – Mol. Phylogen. Evol. 26: 202-214. Patterson, C. (1977): The contribution of paleontology to teleostean phylogeny. – In: Hetch, M. K., Goody, & Hetch, B. M. (eds.): Major Patterns of Vertebrate Evolution; p. 579-643; New York (Plenum Press). – (1981a): Significance of fossils in determining evolutionary relationships. – Ann. Rev. Ecol. Syst. 12: 195-223. – (1981b) Agassiz, Darwin, Huxley, and the fossil record of teleost fishes. – Bull. Brit. Mus. Natur. Hist. (Geol.) 12: 195-223. – (1982): Morphological characters and homology. – In: Jorsey, K. A. & Friday, A. E. (eds.): Problems of Phylogenetic Reconstructions; p. 21-74; New York (Academic Press). – (1993): Osteichthyes: Teleostei. – In: Benton, M. (ed.): The Fossil Record; p. 621-663; London, Glasgow, New York, Tokyo, Melbourne, Madras (Chapman & Hall). – (1994): Bony fishes. – In: Spencer, R. (ed.): Major Features of Vertebrate Evolution; p. 57-84; Knoxville (University of Tennessee Press). Patterson, C. & Rosen, D. E. (1977): Review of the ichthyodectiform and other Mesozoic teleost fishes and the theory and practice of classifying fossils. – Bull. Amer. Mus. Natur. Hist. 158: 83-172. Poyato-Ariza, F. J. (1995): Ichthyemidion a new genus for the elopiform fish ‘Anaethalion vidali’, from the Early Cretaceous of Spain: phylogenetic comments. – C. R. Acad. Sci., Paris 320, sér. IIa: 133-139. – (1996): A revision of the ostariophysin fish family Chanidae, with special reference to the Mesozoic forms. – Palaeo Ichthyologica 6: 5-52. – (1999): The elopiform fish Anaethalion angustus restored, with comments on individual variation. – In: Arratia, G. & Schultze, H.-P. (eds.): Mesozoic Fishes 2 – Systematics and Fossil Record; p. 361-368; München (Verlag Dr. Friedrich Pfeil). Poyato-Ariza, F. J. & Wenz, S. (1994): Naiathaelon okkidion n. g. n. sp. (Teleostei, Elopomorpha) from the Lower Tithonian of Canjuers (Var, France). – Geobios, Mém. Table Redonde Intl. Calcaires lithogr., Lyon 1991: 157-166. Riva Rossi, C. M., Gosztonyi, A. E. & Cozzuol, M. (2000): A Miocene cusk-eel (Ophidiiformes: Ophidiidae) from Peninsula Valdes, Argentina. – J. Vert. Paleontol. 20 (4): 645-650. Robins, C. R. (1989): The phylogenetic relationships of the anguilliform fishes. – In: Böhlke, E. B. (ed.): Orders Anguilliformes and Saccopharyngiformes (Fishes of the Western North Atlantic). Vol. 1; p. 9-23; New Haven, Conn. (Sears Foundation for Marine Research). Santini, F., L. J. Harmon, G. Carnevale, G. & Alfaro, M. E. (2009): Did genome duplication drive the origin of teleosts? A comparative study of diversification in ray-finned fishes. – BMC Evol. Biol. 9: 194-209. Shen, M. (1989): Eohiodon from China and the distribution of osteoglossomorphs. – Vert. PalAsiat. 27: 237-247. (In Chinese with English summary.) – (1996): Fossil ‘osteoglossomorphs’ from East Asia and their implications for teleostean phylogeny. – In: Arratia, G. & Viohl, G. (eds.): Mesozoic Fishes – Systematics and Paleoecology; p. 261-272; München (Verlag Dr. Friedrich Pfeil). Schultze, H.-P. & Arratia, G. (1988): Reevaluation of the caudal skeleton of some actinopterygians fishes. II. Hiodon, Elops and Albula. – J. Morphol. 195: 257-303 Stewart, J. D. (1999): A new genus of Saurodontidae (Teleostei: Ichthyodectiformes) from Upper Cretaceous rocks of the Western Interior of North America. – In: Arratia, G. & Schultze, H.-P. (eds.): Mesozoic Fishes 2 – Systematics and Fossil Record; p. 335-360; München (Verlag Dr. Friedrich Pfeil). Stiassny, M. L. J., Parenti, L. R. & Johnson, G. D. (eds.) (1996): Interrelationships of Fishes. – 496 pp.; San Diego, London, Boston, New York, Sydney, Tokyo, Toronto (Academic Press). Stiassny, M., Wiley, E. O., Johnson, G. D. & Carvalho, M. R. de (2004): Gnathostome fishes. – In: Cracraft, J. & Donoghue, M. J. (eds.): Assembling the Tree of Life; p. 410-429; Oxford, Ohio (Oxford University Press). Taverne, L. (1978): Ostéologie, phylogénèse et systématique des Téléostéens fossils et actuels Ostéoglossomorphes. Deuxième partie. Ostéologie des genres Phareodus, Phareoides, Brychaetus, Musperia, Pantodon, Singida, Notopterus, Xenomystus et Papyrocranus. – Acad. Roy. Belgique, Mém. Cl. Sci., Coll. in 8°, 2e sér. XLII, 6: 213 pp.
273
–
(1981): Ostéologie et position systématique d’ Aethalionopsis robustus (Pisces, Teleostei) du Crétacé inférieur de Bernissart (Belgique) et considerations sur les affinities des Gonorynchiformes. – Bull. Acad. Roy. Belgique, Class Sci., Sér. 5, 67 (12): 958-982. – (1984): A propos de Chanopsis lombardi du Crétacé inférieur du Zaïre (Teleostei, Osteoglossiformes). – Rev. Zool. Afriçain 98 (3): 578-590. – (1989): Crossognathus Pictet, 1858 du Cretacé inférieur de l’ Europe et systématique, paleozoogeographie et biologie des Crossognathiformes nov. ord. (Téléostéens) du Cretacé et du Tertiare. – Palaeontogr. A 207: 79-105. – (1998): Les ostéoglossomorphes marins de l’Éocène du Monte Bolca (Italie): Monopteros Volta 1796, Thrissopterus Heckel, 1856 et Foreyichthys Taverne, 1979. Considérations sur la phylogénie des Téléostéens Ostéoglossomorphes. – Studi e Ricerche sui Giacimenti Terziari di Bolca. VII. Museo Civico di Storia Naturale: 67-158. – (1999): Ostéologie et affinities systématiques de Arratiaelops vectensis gen. nov., téléostéen élopiforme du Wealdien (Crétacé inférieur) d’Angleterre et de Belgique. – Bull. Inst. Roy. Sci. Natur. Belgique, Sci. Terre 69: 77-96. – (2000): Nouvelle données ostéologiques et phylogénétiques sur Palaeonotopterus greenwoodi, notoptéridé (Teleostei, Osteoglossomorpha) du Cénomanian inférieur continental (Crétacé) du Maroc. – Stuttgarter Beitr. Naturk., Ser. B (Geol. Paläontol.) 293: 1-24. – (2002): Les Poissons crétacés de Nardo. 13º. Nardoechelys robinsi gen. et sp. nov., la plus ancienne anguille de type moderne connue par des elements squelettiques (Teleostei, Anguilliformes, Ophichthidae). – Boll. Mus. Civ. St. Natur. Verona, Geol. Paleontol. Preist. 26: 25-31. – (2004): Libanechelys bultyncki gen. et sp. nov., une nouvelle anguille primitive (Teleostei, Anguilliformes) du Cénomanian marin du Liban. – Bull. Inst. Roy. Sci. Natur. Belgique. Sci. Terre 74: 73-87. Taverne, L. & Maisey, J. (1999): A notopterid skull (Teleostei, Osteoglossomorpha) from the continental Early Cretaceous of Southern Morocco. – Amer. Mus. Novitates 3260: 1-12. Wang, C.-H., Kuo, C.-H., Mok, H.-K. & Lee, S.-C. (2003): Molecular phylogeny of elopomorph fishes inferred from mitochondrial 125 ribosomal RNA sequences. – Zool. Scr. 32 (3): 231-241. Wiley, E. O. & Srewart, J. D. (1981): Urenchelys abditus, new species, the first undoubted eel (Teleostei: Anguilliformes) from the Cretaceous of North America. – J. Vert. Paleontol. 1 (1): 43-47 Wilson, M. V. H. (1992): Importance of phylogeny of single and multiple stem-group fossil species with examples of freshwater fishes. – Syst. Biol. 41: 462-470. Woodward, A. S. (1900): Evidence of an extinct eel (Urenchelys anglicus n. sp.) from the English Chalk. – Ann. Mag. Natur. Hist., ser. 7, 5: 321-323. – (1901): Catalogue of the Fossil Fishes in the British Museum (Natural History). – Vol. 4. xxxix + 636 pp. British Museum (Natural History), London. Zhang, J.-Y. (1998): Morphology and phylogenetic relationships of †Kuntulunia (Teleostei: Osteoglossomorpha). – J. Vert. Paleontol. 18 (2): 280-300. – (2002): New fossil osteoglossomorphs from China and the phylogeny of Osteoglossomorpha. – Unpublished Ph.D. dissertation. 172 pp. Graduate School of the Chinese Academy of Sciences, Beijing. – (2004): New fossil osteoglossomorph from Ningxia, China. – J. Vert. Paleontol. 24 (3): 515-524. – (2006): Phylogeny of Osteoglossomorpha, – Vert. PalAsiat. 44 (1): 43-59. Zhang, J.-Y. & Jin, F. (1999): A revision of †Tongxinichthys Ma 1980 (Teleostei: Osteoglossomorpha) from the Lower Cretaceous of northern China. – In: Arratia, G. & Schultze, H.-P. (eds.): Mesozoic Fishes – Systematics and Fossil Record; p. 385-396; München (Verlag Dr. Friedrich Pfeil). Zhang, J.-Y., Jin, F. & Zhou, Z. (1994): A review of Mesozoic osteoglossomorph fish Lycoptera longicephalus. – Vert. PalAsiat. 32 (1): 41-59. (In Chinese with English summary.)
Author’s address: Gloria Arratia, Biodiversity Center, University of Kansas, Dyche Hall, Lawrence, Kansas 66045-7561, USA. E-mail:
[email protected] 274
The study of fossil fishes has advanced significantly over the past few years, giving scientists a rare opportunity to understand the origin and early evolution of major vertebrate groups, ranging from the jawless agnathans to piscine gnathostomes (placoderms, acanthodians, chondrichthyans and osteichthyans). This book presents recent findings on the morphology, phylogeny and paleobiogeography of fossil fishes, as a tribute to Professor Meemann Chang for her contributions to paleoichthyology and to the study of early vertebrate evolution. With a foreword by Dr. Henry Gee (Senior Science Editor of Nature), an introduction, 22 research papers by leading vertebrate paleontologists from 14 countries, and 220 photos and illustrations, this book covers important fossil forms ranging from the Paleozoic to the Cenozoic and reflects research advances based on traditional paleontological methods as well as new techniques such as CT scanning. For fossil agnathans, a new heterostracan is described from the western U.S., the interrelationships and evolutionary history of anaspids are discussed, and evidence is presented showing that anaspids or anaspid-like agnathans may have had a spiral intestine similar to that of gnathostomes. One paper on acanthodians shows that the enigmatic Machaeracanthus may have had ‘paired pairs’ of pectoral fin spines and a perichondrally ossified scapulocoracoid. New placoderms from northern Siberia and western Australia are described, and the pectoral fin development in gnathostomes is reviewed based on a revision of previous hypotheses and new fossil arthrodire material. Chondrichthyans are represented by the description of a giant electric ray from the Eocene of Italy, and by new articulated material from the Early Devonian of the Northwest Territories showing that the scale- and spine-based distinctions between acanthodians and chondrichthyans do not account for the diversity that is now apparent. Nine papers on osteichthyans cover wide ranging topics from the cosmine histology of a stem sarcopterygian, to the characters of the stem tetrapod neurocranium, and to new Tertiary osteoglossid fishes. New morphological and phylogenetic information on the snout of Devonian dipnoans and the neurocranium of Powichthys is presented based on CT scanning. An uncrushed specimen of Eusthenopteron enables a revision of the ethmosphenoid morphology bearing on the choana, while a lungfish study indicates that the postcranial anatomy may be an underexploited source of characters for phylogenetic studies. The role of fossils in phylogenetic studies is also examined based on teleost phylogenies. While the link between morphology, phylogeny and paleobiogeography permeates many papers, two papers have a predominant focus on paleoecology and paleobiogeography – one reviewing the ecological connections and paleobiogeographic implications of the Jehol Biota, and the other reviewing the South American Devonian vertebrate record, demonstrating the presence of two faunal assemblages of which the earlier one equates with the “Malvinokaffric Realm” based on invertebrate communities. Professor Meemann Chang is a Research Professor at the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP), Beijing, China. She is an honorary member of the Society of Vertebrate Paleontology, a past president of the International Paleontological Association, and a Member (Academician) of the Chinese Academy of Sciences.
ISBN 978-3-89937-122-2
www.pfeil-verlag.de
