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      Skeletal Transformations and the Origin of Baleen Whales (Mammalia, Cetacea, Mysticeti): A Study on Evolutionary Patterns

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          Abstract

          A review of the morphological patterns exhibited by all the main radiations of mysticete (baleen whale) cetaceans provided a broad assessment of the fundamental morphological transformations that occurred in the transition to the Mysticeti clade. Skull and postcranial characters were illustrated, described and compared, and their distribution was mapped on a combined phylogeny in the search for morphological support for the principal mysticete clades (i.e., Mysticeti, Chaeomysticeti and Balaenomorpha). In particular, characters of the skull (rostrum, vertex, temporal fossa, tympanic bulla and dentary) and the postcranial appendicular skeleton (scapula, humerus, radius and ulna) were all involved at different degrees in the process of morphological transformations leading to the modern-day mysticetes. Apart from a few typical characteristics of the rostrum that were already present in the earliest-diverging mysticetes (presence of lateral process of the maxilla, presence of multiple dorsal infraorbital foramina, thin lateral border of maxilla and presence of mesorostral groove), most of the other anatomical districts were unaffected by the transition so the earliest mysticetes show a number of archaeocete characters in the tympanic bulla, dentary and skull roof. The analysis of the whole dataset supported the hypothesis that the origin and evolution of mysticetes constituted a step-wise process and that the bauplan of the modern-day mysticetes was fully assembled at the level of the common ancestor of all Balaenomorpha.

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          The therian skull : a lexicon with emphasis on the odontocetes

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            Morphological and molecular evidence for a stepwise evolutionary transition from teeth to baleen in mysticete whales.

            The origin of baleen in mysticete whales represents a major transition in the phylogenetic history of Cetacea. This key specialization, a keratinous sieve that enables filter-feeding, permitted exploitation of a new ecological niche and heralded the evolution of modern baleen-bearing whales, the largest animals on Earth. To date, all formally described mysticete fossils conform to two types: toothed species from Oligocene-age rocks ( approximately 24 to 34 million years old) and toothless species that presumably utilized baleen to feed (Recent to approximately 30 million years old). Here, we show that several Oligocene toothed mysticetes have nutrient foramina and associated sulci on the lateral portions of their palates, homologous structures in extant mysticetes house vessels that nourish baleen. The simultaneous occurrence of teeth and nutrient foramina implies that both teeth and baleen were present in these early mysticetes. Phylogenetic analyses of a supermatrix that includes extinct taxa and new data for 11 nuclear genes consistently resolve relationships at the base of Mysticeti. The combined data set of 27,340 characters supports a stepwise transition from a toothed ancestor, to a mosaic intermediate with both teeth and baleen, to modern baleen whales that lack an adult dentition but retain developmental and genetic evidence of their ancestral toothed heritage. Comparative sequence data for ENAM (enamelin) and AMBN (ameloblastin) indicate that enamel-specific loci are present in Mysticeti but have degraded to pseudogenes in this group. The dramatic transformation in mysticete feeding anatomy documents an apparently rare, stepwise mode of evolution in which a composite phenotype bridged the gap between primitive and derived morphologies; a combination of fossil and molecular evidence provides a multifaceted record of this macroevolutionary pattern.
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              A supermatrix analysis of genomic, morphological, and paleontological data from crown Cetacea

              Background Cetacea (dolphins, porpoises, and whales) is a clade of aquatic species that includes the most massive, deepest diving, and largest brained mammals. Understanding the temporal pattern of diversification in the group as well as the evolution of cetacean anatomy and behavior requires a robust and well-resolved phylogenetic hypothesis. Although a large body of molecular data has accumulated over the past 20 years, DNA sequences of cetaceans have not been directly integrated with the rich, cetacean fossil record to reconcile discrepancies among molecular and morphological characters. Results We combined new nuclear DNA sequences, including segments of six genes (~2800 basepairs) from the functionally extinct Yangtze River dolphin, with an expanded morphological matrix and published genomic data. Diverse analyses of these data resolved the relationships of 74 taxa that represent all extant families and 11 extinct families of Cetacea. The resulting supermatrix (61,155 characters) and its sub-partitions were analyzed using parsimony methods. Bayesian and maximum likelihood (ML) searches were conducted on the molecular partition, and a molecular scaffold obtained from these searches was used to constrain a parsimony search of the morphological partition. Based on analysis of the supermatrix and model-based analyses of the molecular partition, we found overwhelming support for 15 extant clades. When extinct taxa are included, we recovered trees that are significantly correlated with the fossil record. These trees were used to reconstruct the timing of cetacean diversification and the evolution of characters shared by "river dolphins," a non-monophyletic set of species according to all of our phylogenetic analyses. Conclusions The parsimony analysis of the supermatrix and the analysis of morphology constrained to fit the ML/Bayesian molecular tree yielded broadly congruent phylogenetic hypotheses. In trees from both analyses, all Oligocene taxa included in our study fell outside crown Mysticeti and crown Odontoceti, suggesting that these two clades radiated in the late Oligocene or later, contra some recent molecular clock studies. Our trees also imply that many character states shared by river dolphins evolved in their oceanic ancestors, contradicting the hypothesis that these characters are convergent adaptations to fluvial habitats.
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                Author and article information

                Journal
                DIVEC6
                Diversity
                Diversity
                MDPI AG
                1424-2818
                March 2022
                March 18 2022
                : 14
                : 3
                : 221
                Article
                10.3390/d14030221
                b43bf1ff-4b42-4728-8fb5-216fd132287d
                © 2022

                https://creativecommons.org/licenses/by/4.0/

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