The Early Diversification of Ray-Finned Fishes (Actinopterygii) a N an Ecomorphological Approach
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The early diversification ofray-finned fishes (Actinopterygii) anecomorphological approach Weronica Klasson Degree project inbiology, 2008 Examensarbete ibiologi, 20 p,2008 Biology Education Center and Department ofPhysiology and Developmental Biology Supervisor: Henning Blom The early diversification of ray-finned fishes (Actinopterygii); an ecomorphological approach Weronica Klasson Uppsala University, Subdepartment of Evolutionary Organismal Biology, Department of Physiology and Developmental Biology. Examensarbete 30hp Contents Sammanfattning 2 Abstract 3 1. Introduction 4 2. Taxonomical and morphological framework 6 2.2 Shape, habitat and diet 6 2.3 Adaptation to Salt & freshwater 8 3. Environmental framework 10 3.1 Environment vs. morphology 10 3.2 Paleogeographic and paleoenvironments 11 3.3 Localities and there environments 12 3.3.1 Devonian 14 3.3.2 Carboniferous 16 4. Morphometrics analysis 19 4.1 Relative warp analysis 20 4.2 Disparity measures 21 5. Discussion 22 5.1 Ecomorphology 25 Acknowledgments 28 References 29 Appendix 1 33 List of Devonian taxa 33 List of Carboniferous taxa 34 Appendix 2 41 Relative Warp 1.1 41 Relative Warp 1.2 42 Appendix 3 43 Relative warp scores matrix 43 Relative warp scores 58 Landmarks 58 Appendix 4 64 Reconstructions of the Devonian and Carboniferous fishes 64 1 De strålfeniga fiskarnas (Actinopterygii) tidiga uppblomstring; en ekomorfologisk studie Sammanfattning Strålfeniga fiskar är idag den största och mest framgångsrika gruppen av fiskar och omfattar omkring 27000 arter, och finns i nästan alla olika typer av miljöer (habitat). Dagens mångfald är ett resultat av en rad uppblomstringar av vilka dom tidigaste verkar sammanfalla med såväl uppkomst och tidigaste utveckling under devon (416-359.2 miljoner år sedan) och karbon (359.2-299 miljoner år sedan). En morfometrisk analys (jämförande av kroppsformer) är gjord för att undersöka eventuella evolutionära mönster som kan urskiljas bland de devonska och karbonska strålfeniga fiskarnas kroppsformer (morfologier). Denna studie visar att den morfologiska mångfalden, som undersöks och kvantifierats med hjälp av olika numeriska metoder, kan jämföras med den taxonomiska mångfalden. Denna approach uppvisar en trend i vilken en jämn ökning av den morfologiska och taxonomiska mångfalden under den senare hälften av Devon, med en efterföljande mer explosionsartad ökning under tidig Karbon. Slutet av karbon uppvisar däremot en dramatisk minskning av såväl den morfologiska som taxonomiska mångfalden, en händelse som är svår att förstå och som kräver ytterligare studier. Denna studie har också visat att skillnader i kroppsform, även under dessa tidiga episoder av evolution, snarare beror på hur fiskarna levde och jagade (d.v.s. s.k. mikrohabitat) snarare än fysiska miljöfaktorer (t.ex. salinitet). Sex ”ekologiska kroppsformsgrupper” (ekomorfologiska grupper) har föreslagits och genom att jämföra dessa med motsvarande ”ekomorfologiska grupper” bland nulevande strålfeniga fiskar är det möjligt att dra mer omfattande slutsatser om hur de levde och jagade under devon och karbon. Noterbart med denna analys är att de devonska formerna hamnar i den ”ekomorfologiska grupper” som anses överraska sina byten, s.k. ”ligga och vänta predatorer” (“lay-in-wait-predators”), eller de som aktivt följer sina byten, s.k. ”följande predatorer” (“rover-predators”). Detta tyder på att de devonska formerna troligtvis var fiskätare snarare än plankton- och bryozo-ätare, vilka verkar representera en senare evolutionär utveckling. 2 The early diversification of ray-finned fishes (Actinopterygii); an ecomorphological approach Abstract The actinopterygians are the now largest and most successful group of living fishes with about 27 000 species in almost every aquatic environment. This diversity is a result of numerous radiations through time, including the origins and early diversification in the Devonian and Carboniferous. Morphometric analyses have been performed in order to investigate patterns of morphological diversity during these early episodes of actinopterygian evolution. This study shows that the disparity, which has been quantified by various methods, can be correlated to the overall taxonomic diversity. This pattern of morphological and taxonomical diversification starts with a steady increase in the Devonian, followed by what appears to be a major radiation event in the early Carboniferous. However, in the late Carboniferous both the diversity and disparity drastically decreases and the reason for this is unknown. This study also shows that there are no relation between body shape and the environments salinity. Instead it seems to be the microhabitats and the way to hunt that have the most effect on body shape. Six ecomorphological groups have been detected in the studied data set, which can give clues about feeding strategies when compared to ecomorphological groups established for recent fishes. Interestingly, the ecomorphological groupings suggest that the Devonian fishes seem to be “lay-in- wait-predators” and “rover-predators”. This means that the early ray-finned fishes probably were piscivores, rather then browsers and plankton-eaters, which seems to be a later evolutionary invention. 3 1. Introduction Eifelian and Frasnian of Scotland and Canada (Friedman and Blom 2006). The Actinopterygii, or ray-finned Cheriolepis was followed by an fishes, belong to the osteichthyans increased record of taxa in the late (bony fishes) together with the sister Devonian, probably due to the group lobe-finned fishes extension of floodplains and deltaic (Sarcopterygii). environments. The diversification then The actinopterygians are defined by speeded up in the Early Carboniferous several characters, including the (Viséan) when more ecological niches median fin rays that are inserted became available after the placoderms directly into the body, with no became extinct. Many niches also intervening basal lobe. The primitive become available when the tetrapods diamond-shaped scales, covered with went up on land and the sarcopterygian ganoine, the pelvic girdle that are decrease in the marine environments replaced with part of the (Janvier 1996). Today there are about metapterygium and soft tissue, and the 27 000 species of ray-finned fishes presence of acrodine (a transparent cap (Hurley et al. 2007). of mineralized tissue on the tip of the The phylogeny of the basic teeth) (Janvier 1996). (oldest) Actinopterygii is based on Ray-finned fishes are the different morphological characters largest and most successful group of (Figure 1), while the recent and living fishes. They are found in every younger is based on molecular data. aquatic habitat, from the high-pressure This makes it difficult to combine the depth and salinity of the ocean to living forms with the fossil record in freshwater streams and ponds, from the phylogenetic analyses. The fossil artic coldness (-1.8C) to the tropical record may involve some problems in warmth (+40C). They are also tolerant that some taxa can be “misplaced”. For to pH level from 4 to 10 and low example, two specimens of the same oxygen levels. Some can breathe air species could be put in two different and some can even cope without taxa (or the contrary two different taxa oxygen for periods of time (Moyle et put in to one) on the basis of; sexual al. 2004). There are even species that dimorphism, different size in different can crawl on land. habitat (young/adult), local The earliest record of possible morphology changes (ecological ray-finned fishes are isolated scales species can be more common then first and fragments from the Silurian, but realized) or the taphonmy (the the oldest uncontested preservation of the fossil that could actinopterygianas to be described alter the body shape) (Moyle et al. based on articulated material is 2004). Cheriolepis (Agassiz 1835) from the 4 Figure 1. Phylogenetic tree, showing the basal actinopterygians from the Devonian (bold) and Carboniferous (C. wilidi & P. decorus are Permian), based on 185 morphological characters. Also note the different environments in which the fishes lived: M = Marine F = freshwater and B = Brackish (modified from Cloutier et al. 2004). In this study the taxonomical and (2006) who explore the early morphological diversity, of several morphological diversification in the Devonian and Carboniferous coelacanth clade. actinopterygians were investigated in These types of morphometrics an environmental and temporal context methods are not mainly used on fossils to answer questions relating to the but are frequently used on recent fishes early radiation of the group. The to test various hypotheses on morphological diversity between ecomorphological questions (Costa et species have been studied to show al. 2007). A link between morphology differences in morphology depending and ecology has been shown in studies on the environment, more specifically on recent fish and Costa and if the species lived in marine water or Cataudella (2007) showed a fresh water environments have relationship between trophic ecology different body shapes. Such questions and morphology. Ruben and Adams can be referred to the concept of (2001) showed that ecology control the ecomorphology, which can be defined morphology more than the genes, as, “a study of the relationship between which means that the plasticity of the the ecological role of an individual and fish was more than first expected. The its morphological adaptations” morphology of a species is then shaped (Rickleffs 1990). One way to study