New Protocetid Whale from the Middle Eocene of Pakistan: Birth on Land, Precocial Development, and Sexual Dimorphism
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{"title"=>"New protocetid whale from the middle Eocene of Pakistan: Birth on land, precocial development, and sexual dimorphism", "type"=>"journal", "authors"=>[{"first_name"=>"Philip D.", "last_name"=>"Gingerich", "scopus_author_id"=>"7006477387"}, {"first_name"=>"Munir", "last_name"=>"Ul-Haq", "scopus_author_id"=>"26424844600"}, {"first_name"=>"Wighart", "last_name"=>"Von Koenigswald", "scopus_author_id"=>"6603287914"}, {"first_name"=>"William J.", "last_name"=>"Sanders", "scopus_author_id"=>"7202691637"}, {"first_name"=>"B. Holly", "last_name"=>"Smith", "scopus_author_id"=>"56379121300"}, {"first_name"=>"Iyad S.", "last_name"=>"Zalmout", "scopus_author_id"=>"6507215182"}], "year"=>2009, "source"=>"PLoS ONE", "identifiers"=>{"issn"=>"19326203", "isbn"=>"1932-6203 (Electronic)", "pmid"=>"19194487", "scopus"=>"2-s2.0-84892938337", "doi"=>"10.1371/journal.pone.0004366", "sgr"=>"84892938337", "pui"=>"618784158"}, "id"=>"b2bdf416-b6d0-3f33-91c6-ab900d77875a", "abstract"=>"BACKGROUND: Protocetidae are middle Eocene (49-37 Ma) archaeocete predators ancestral to later whales. They are found in marine sedimentary rocks, but retain four legs and were not yet fully aquatic. Protocetids have been interpreted as amphibious, feeding in the sea but returning to land to rest. METHODOLOGY/PRINCIPAL FINDINGS: Two adult skeletons of a new 2.6 meter long protocetid, Maiacetus inuus, are described from the early middle Eocene Habib Rahi Formation of Pakistan. M. inuus differs from contemporary archaic whales in having a fused mandibular symphysis, distinctive astragalus bones in the ankle, and a less hind-limb dominated postcranial skeleton. One adult skeleton is female and bears the skull and partial skeleton of a single large near-term fetus. The fetal skeleton is positioned for head-first delivery, which typifies land mammals but not extant whales, evidence that birth took place on land. The fetal skeleton has permanent first molars well mineralized, which indicates precocial development at birth. Precocial development, with attendant size and mobility, were as critical for survival of a neonate at the land-sea interface in the Eocene as they are today. The second adult skeleton is the most complete known for a protocetid. The vertebral column, preserved in articulation, has 7 cervicals, 13 thoracics, 6 lumbars, 4 sacrals, and 21 caudals. All four limbs are preserved with hands and feet. This adult is 12% larger in linear dimensions than the female skeleton, on average, has canine teeth that are 20% larger, and is interpreted as male. Moderate sexual dimorphism indicates limited male-male competition during breeding, which in turn suggests little aggregation of food or shelter in the environment inhabited by protocetids. CONCLUSIONS/SIGNIFICANCE: Discovery of a near-term fetus positioned for head-first delivery provides important evidence that early protocetid whales gave birth on land. This is consistent with skeletal morphology enabling Maiacetus to support its weight on land and corroborates previous ideas that protocetids were amphibious. Specimens this complete are virtual 'Rosetta stones' providing insight into functional capabilities and life history of extinct animals that cannot be gained any other way.", "link"=>"http://www.mendeley.com/research/new-protocetid-whale-middle-eocene-pakistan-birth-land-precocial-development-sexual-dimorphism", "reader_count"=>215, "reader_count_by_academic_status"=>{"Unspecified"=>3, "Professor > Associate Professor"=>8, "Student > Doctoral Student"=>16, "Researcher"=>48, "Student > Ph. D. Student"=>45, "Student > Postgraduate"=>13, "Other"=>18, "Student > Master"=>27, "Student > Bachelor"=>17, "Lecturer"=>3, "Lecturer > Senior Lecturer"=>2, "Professor"=>15}, "reader_count_by_user_role"=>{"Unspecified"=>3, "Professor > Associate Professor"=>8, "Student > Doctoral Student"=>16, "Researcher"=>48, "Student > Ph. D. Student"=>45, "Student > Postgraduate"=>13, "Other"=>18, "Student > Master"=>27, "Student > Bachelor"=>17, "Lecturer"=>3, "Lecturer > Senior Lecturer"=>2, "Professor"=>15}, "reader_count_by_subject_area"=>{"Unspecified"=>5, "Agricultural and Biological Sciences"=>122, "Arts and Humanities"=>1, "Business, Management and Accounting"=>1, "Veterinary Science and Veterinary Medicine"=>1, "Earth and Planetary Sciences"=>60, "Energy"=>1, "Environmental Science"=>11, "Biochemistry, Genetics and Molecular Biology"=>3, "Nursing and Health Professions"=>1, "Medicine and Dentistry"=>2, "Psychology"=>1, "Social Sciences"=>5, "Philosophy"=>1}, "reader_count_by_subdiscipline"=>{"Medicine and Dentistry"=>{"Medicine and Dentistry"=>2}, "Social Sciences"=>{"Social Sciences"=>5}, "Psychology"=>{"Psychology"=>1}, "Unspecified"=>{"Unspecified"=>5}, "Environmental Science"=>{"Environmental Science"=>11}, "Arts and Humanities"=>{"Arts and Humanities"=>1}, "Energy"=>{"Energy"=>1}, "Earth and Planetary Sciences"=>{"Earth and Planetary Sciences"=>60}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>122}, "Business, Management and Accounting"=>{"Business, Management and Accounting"=>1}, "Nursing and Health Professions"=>{"Nursing and Health Professions"=>1}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>3}, "Veterinary Science and Veterinary Medicine"=>{"Veterinary Science and Veterinary Medicine"=>1}, "Philosophy"=>{"Philosophy"=>1}}, "reader_count_by_country"=>{"Colombia"=>1, "Ecuador"=>1, "Argentina"=>8, "Romania"=>1, "United States"=>11, "Japan"=>1, "United Kingdom"=>3, "Malaysia"=>1, "Thailand"=>1, "Canada"=>2, "Czech Republic"=>1, "Brazil"=>4, "Poland"=>1, "France"=>2, "Chile"=>2, "Germany"=>4, "Indonesia"=>1}, "group_count"=>6}

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  • {"month"=>"7", "year"=>"2019", "pdf_views"=>"17", "xml_views"=>"0", "html_views"=>"29"}
  • {"month"=>"8", "year"=>"2019", "pdf_views"=>"21", "xml_views"=>"0", "html_views"=>"38"}
  • {"month"=>"9", "year"=>"2019", "pdf_views"=>"19", "xml_views"=>"1", "html_views"=>"72"}
  • {"month"=>"10", "year"=>"2019", "pdf_views"=>"30", "xml_views"=>"0", "html_views"=>"74"}
  • {"month"=>"11", "year"=>"2019", "pdf_views"=>"22", "xml_views"=>"1", "html_views"=>"85"}
  • {"month"=>"12", "year"=>"2019", "pdf_views"=>"20", "xml_views"=>"1", "html_views"=>"131"}
  • {"month"=>"1", "year"=>"2020", "pdf_views"=>"27", "xml_views"=>"0", "html_views"=>"73"}
  • {"month"=>"2", "year"=>"2020", "pdf_views"=>"11", "xml_views"=>"0", "html_views"=>"115"}
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  • {"month"=>"8", "year"=>"2020", "pdf_views"=>"11", "xml_views"=>"0", "html_views"=>"21"}
  • {"month"=>"9", "year"=>"2020", "pdf_views"=>"7", "xml_views"=>"0", "html_views"=>"38"}

Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/910545"], "description"=>"<p>Pelves are viewed ventrally. Sexes differ in the shape of the notch separating inferior rami of left and right pubes. (A)– Female <i>Dama dama</i> has the more U-shaped pubic notch labeled in red. (B)– Male <i>Dama dama</i> has the more V-shaped pubic notch labeled in red. (C)– Male <i>Maiacetus inuus</i>, GSP-UM 3551, has the V-shaped pubic notch labeled in red. Drawings of <i>Dama dama</i> are from <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Lemppenau1\" target=\"_blank\">[29]</a>. Part of the right pubic ramus of GSP-UM 3551 is restored from the left side.</p>", "links"=>[], "tags"=>["morphology", "extant", "sexually-dimorphic", "fallow", "compared"], "article_id"=>580996, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g010", "stats"=>{"downloads"=>8, "page_views"=>248, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Pelvic_morphology_in_the_extant_sexually_dimorphic_fallow_deer_Dama_dama_compared_to_that_of_male_Maiacetus_inuus_/580996", "title"=>"Pelvic morphology in the extant sexually-dimorphic fallow deer <i>Dama dama</i> compared to that of male <i>Maiacetus inuus</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 07:05:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/910226"], "description"=>"<p>(A)–Photograph showing bones (shaded white) and teeth (shaded brown) in lateral view. (B)–CT image overlain on tooth-bearing part of fetal skull in lateral veiw. <i>Abbreviations</i>: left frontal (<i>L. Front.</i>), left jugal (<i>L Jug.</i>), occipital (<i>Occip</i>.) with left and right condyles flanking the foramen magnum, and left and right dentaries (<i>L dent.</i>, <i>R dent.</i>). Teeth are shaded brown, with darker brown representing enamel, and lighter brown representing roots and exposed dentine. White and black labels identify teeth of the left and right sides, respectively. Dotted lines trace outlines of fully formed crowns of left dP<sup>3</sup> and dP<sup>4</sup>. These crowns are visible on the surface (A) where thin bone of the maxilla is pressed over more rigid underlying crowns, and as denser masses in the CT scan (B). Remaining teeth are identified by size and position relative to dP<sup>3</sup> and dP<sup>4</sup>. Note presence of the developing crown of permanent left M<sup>1</sup> posterodorsal to the crown of left dP<sup>4</sup> (dorsal to the left jugal and posterior to the left frontal). Partial crown of right M<sup>1</sup> (unlabeled) is visible just below and posterior to left M<sup>1</sup>.</p>", "links"=>[], "tags"=>["Evolutionary biology", "evolutionary biology/animal behavior", "evolutionary biology/paleontology"], "article_id"=>580681, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g008", "stats"=>{"downloads"=>1, "page_views"=>13, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Fetal_skull_of_Maiacetus_inuus_GSP_UM_3475b_/580681", "title"=>"Fetal skull of <i>Maiacetus inuus</i> (GSP-UM 3475b).", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 07:03:35"}
  • {"files"=>["https://ndownloader.figshare.com/files/909203"], "description"=>"<p>Skull of the adult female (GSP-UM 3475a) is colored beige with brown teeth; her postcranial skeleton is colored red; the fetal skeleton (GSP-UM 3475b) is colored blue with red teeth. Blue dashed lines indicate the contours of the three field jackets and the red dashed line marks the edge of erosion.</p>", "links"=>[], "tags"=>["fetal", "skeletons", "protocetid"], "article_id"=>579658, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g002", "stats"=>{"downloads"=>0, "page_views"=>15, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Adult_female_and_fetal_skeletons_type_of_the_protocetid_Maiacetus_inuus_/579658", "title"=>"Adult female and fetal skeletons (type) of the protocetid <i>Maiacetus inuus</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 06:57:22"}
  • {"files"=>["https://ndownloader.figshare.com/files/911136"], "description"=>"<p>Living whales are not included because none have deciduous teeth and many have few or no teeth. Because early whales evolved from artiodactyls <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Gingerich2\" target=\"_blank\">[3]</a>, <i>Ovis</i>, <i>Dama</i>, <i>Hexaprotodon</i>, and <i>Giraffa</i> are better analogs than primates.</p>1<p>Presence of mineralization of the crown of M<sup>1</sup> in the fetal skull of <i>Maiacetus inuus</i> indicates precocial development and the probability that newborns were open-eyed, hairy and nidifugous.</p>", "links"=>[], "tags"=>["maturity", "categories", "langer", "mineralization", "molar", "teeth", "newborn", "terrestrial", "mammals", "compared"], "article_id"=>581580, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.t004", "stats"=>{"downloads"=>8, "page_views"=>12, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Developmental_maturity_categories_of_Langer_21_and_mineralization_of_first_molar_M_1_and_other_permanent_teeth_in_newborn_terrestrial_mammals_compared_to_the_early_whale_Maiacetus_inuus_/581580", "title"=>"Developmental maturity categories of Langer [21] and mineralization of first molar M<sup>1</sup> and other permanent teeth in newborn terrestrial mammals compared to the early whale <i>Maiacetus inuus</i>.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-02-21 07:08:45"}
  • {"files"=>["https://ndownloader.figshare.com/files/909527"], "description"=>"<p>Red rectangle highlights the platy limestones and marls of the upper part of the marine Habib Rahi Formation. Nannoplankton <i>Sphenolithus spininger</i> and <i>Helicosphaera bramlettei</i> indicative of NP zones 14–15 have been reported from the Habib Rahi Formation <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Kthe1\" target=\"_blank\">[13]</a>. The age of <i>Maiacetus inuus</i> is about 47.5 Ma. Figure from <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Gingerich6\" target=\"_blank\">[14]</a>.</p>", "links"=>[], "tags"=>["constraints"], "article_id"=>579990, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g004", "stats"=>{"downloads"=>1, "page_views"=>9, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Temporal_constraints_on_the_age_of_Maiacetus_inuus_/579990", "title"=>"Temporal constraints on the age of <i>Maiacetus inuus</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 06:59:21"}
  • {"files"=>["https://ndownloader.figshare.com/files/910755"], "description"=>"<p>(A)—<i>Maiacetus inuus</i>, a semiaquatic foot-powered swimmer from the Middle Eocene. (B)—<i>Dorudon atrox</i>, a fully-aquatic tail-powered swimmer from the Late Eocene. Baseline is mean length of anterior thoracic vertebrae (stippled box). Sacral vertebrae are enclosed in a second stippled box where sacrals can be identified (e.g., by co-ossification). <i>Maiacetus</i> has a profile more like that of mammals capable of supporting their weight on land, whereas <i>Dorudon</i> has the profile of a modern whale. Interpretation of profiles and the method of median serial-multiple-regression estimation of body weights is explained in <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Gingerich8\" target=\"_blank\">[50]</a>. <i>Abbreviations</i>: <i>TCNDSFB</i>, longest tooth length, condylobasal cranium length, narial position, dentary length, symphysis position, mandibular foramen height; and bulla length, respectively; <i>SHRC</i>, scapula, humerus, radius, and Mc-III lengths, respectively; <i>IFTT</i>, innominate, femur, tibia, and Mt-III lengths, respectively.</p>", "links"=>[], "tags"=>["skeletal", "profiles", "eocene", "archaeocete", "whales"], "article_id"=>581207, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g012", "stats"=>{"downloads"=>1, "page_views"=>19, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Proportion_adjusted_skeletal_profiles_of_middle_and_late_Eocene_archaeocete_whales_Maiacetus_inuus_and_Dorudon_atrox_respectively_/581207", "title"=>"Proportion-adjusted skeletal profiles of middle and late Eocene archaeocete whales <i>Maiacetus inuus</i> and <i>Dorudon atrox</i>, respectively.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 07:06:39"}
  • {"files"=>["https://ndownloader.figshare.com/files/910886"], "description"=>"<p><i>Maiacetus inuus</i> is similar to <i>Rodhocetus</i> species, but lacks elongation of the manus, hind limb, and pes characteristic of the latter. <i>Maiacetus</i> is about equally aquatic (PC-II) to <i>Rodhocetus</i> but less specialized as a hind-limb swimmer (PC-III). It falls closest to the giant otter (<i>Pteronura brasiliensis</i>) among extant mammals. Background and comparative data for this analysis are documented and explained in <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Gingerich3\" target=\"_blank\">[4]</a>.</p>", "links"=>[], "tags"=>["components", "skeletal", "proportions", "50", "extant", "semiaquatic"], "article_id"=>581336, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g013", "stats"=>{"downloads"=>1, "page_views"=>20, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Principal_components_plot_of_trunk_and_limb_skeletal_proportions_for_Maiacetus_inuus_and_a_representative_sample_of_50_extant_semiaquatic_mammals_/581336", "title"=>"Principal components plot of trunk and limb skeletal proportions for <i>Maiacetus inuus</i> and a representative sample of 50 extant semiaquatic mammals.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 07:07:22"}
  • {"files"=>["https://ndownloader.figshare.com/files/911202"], "description"=>"<p><i>Abbreviations</i>: <i>C</i>, cervical vertebra; <i>Ca</i>, caudal vertebra; <i>L</i>, lumbar vertebra; <i>S</i>, sacral vertebra; <i>T</i>, thoracic vertebra.</p>*<p>Asterisk indicates estimated measurement.</p><p>( ) indicate interpolated measurement.</p><p>— Ca3 through Ca12 are bilaterally compressed, precluding measurement of their width.</p>", "links"=>[], "tags"=>["centrum"], "article_id"=>581660, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.t003", "stats"=>{"downloads"=>1, "page_views"=>11, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Vertebral_centrum_measurements_mm_for_adult_male_Maiacetus_inuus_GSP_UM_3551_Figure_9_/581660", "title"=>"Vertebral centrum measurements (mm) for adult male <i>Maiacetus inuus</i> (GSP-UM 3551; Figure 9).", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-02-21 07:09:12"}
  • {"files"=>["https://ndownloader.figshare.com/files/910629"], "description"=>"<p>(A)–Astragalus and cuboid of <i>Artiocetus clavis</i> (GSP-UM 3458, type, reversed). (B)–Ankle of <i>Maiacetus inuus</i> (GSP-UM 3551). (C)–Ankle of <i>Rodhocetus balochistanensis</i> (GSP-UM 3485, type). Specimens are drawn to approximately the same astragalus+cuboid length, and viewed dorsally. Blue arrow points to distinctive indentation in lateral margin of astragalus of <i>M. inuus</i>; red arrows point to the broad and deep, narrow and intermediate, and narrow and shallow indentations in the calcaneum and lateral margins of cuboids. <i>Abbreviations</i>: <i>Ast.</i>, astragalus; <i>Cal.</i>, calcaneum; <i>Cub.</i>, cuboid; <i>Ecc.</i>, ectocuneiform; <i>Enc.</i>, entocuneiform; <i>Mec.</i>, mesocuneiform; <i>Nav.</i>, navicular.</p>", "links"=>[], "tags"=>["bones"], "article_id"=>581086, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g011", "stats"=>{"downloads"=>8, "page_views"=>169, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Comparison_of_right_ankle_bones_of_Artiocetus_Maiacetus_and_Rodhocetus_/581086", "title"=>"Comparison of right ankle bones of <i>Artiocetus</i>, <i>Maiacetus</i>, and <i>Rodhocetus</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 07:05:55"}
  • {"files"=>["https://ndownloader.figshare.com/files/909102"], "description"=>"<p>(A, B)– <i>Dorudon atrox</i> (5.0 m; 36.5 Ma) based on UM 101222 and 101215 <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Uhen1\" target=\"_blank\">[11]</a> in lateral and dorsal views, respectively. (C, D)– <i>Maiacetus inuus</i> (2.6 m; 47.5 Ma) based on male specimen GSP-UM 3551 in lateral and dorsal views, respectively.</p>", "links"=>[], "tags"=>["eocene", "archaeocete", "whales"], "article_id"=>579554, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g001", "stats"=>{"downloads"=>4, "page_views"=>25, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Skeletons_of_the_Eocene_archaeocete_whales_Dorudon_atrox_and_Maiacetus_inuus_in_swimming_pose_/579554", "title"=>"Skeletons of the Eocene archaeocete whales <i>Dorudon atrox</i> and <i>Maiacetus inuus</i> in swimming pose.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 06:56:43"}
  • {"files"=>["https://ndownloader.figshare.com/files/909872"], "description"=>"<p>Skull in dorsal view (A) with interpretive shading and labels (B). Shaded components include skull bones (grey), teeth (brown), and postcranial bones (red). <i>Osteological abbreviations</i>: <i>Dent.</i>, dentary; <i>Front.</i>, frontal; <i>Jug.</i>, jugal; <i>L</i>, left; <i>Lac.</i>, lacrimal; <i>Max.</i>, maxilla; <i>Nas.</i>, nasal; <i>Par.</i>, parietal; <i>Pmax.</i>, premaxilla; R, right, rib; <i>Scap.</i>, scapula; <i>Squ.</i>, squamosal.</p>", "links"=>[], "tags"=>["Evolutionary biology", "evolutionary biology/animal behavior", "evolutionary biology/paleontology"], "article_id"=>580328, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g006", "stats"=>{"downloads"=>1, "page_views"=>12, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Skull_of_Maiacetus_inuus_GSP_UM_3475a_female_/580328", "title"=>"Skull of <i>Maiacetus inuus</i> (GSP-UM 3475a; female).", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 07:01:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/909364"], "description"=>"<p>Map showing the Kunvit area in the southwestern part of Lakha Kach syncline, northwest of the town of Rakhni. Red diamonds mark type localities for <i>Artiocetus clavis</i> (GSP-UM 3458), <i>Rodhocetus balochistanensis</i> (GSP-UM 3485) <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Gingerich2\" target=\"_blank\">[3]</a>, <i>Protosiren eothene</i> (GSP-UM 3487) <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Zalmout1\" target=\"_blank\">[49]</a>, <i>Makaracetus bidens</i> (GSP-UM 3570; from younger beds of the overlying Domanda Formation) <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Gingerich1\" target=\"_blank\">[2]</a>, and <i>Maiacetus inuus</i> (GSP-UM 3475a, b, female, fetal skeleton). Localities are in the upper part of the Habib Rahi Formation of early Lutetian age <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Haq1\" target=\"_blank\">[12]</a>. The red circle marks the locality of the referred specimen of <i>Maiacetus inuus</i> (GSP 3551, male), yellow circles show localities of other archaeocete specimens, and white circles show localities of other vertebrate specimens.</p>", "links"=>[], "tags"=>["localities", "eocene", "whales", "balochistan"], "article_id"=>579817, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g003", "stats"=>{"downloads"=>4, "page_views"=>22, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Map_showing_localities_of_some_Eocene_whales_in_eastern_Balochistan_Pakistan_/579817", "title"=>"Map showing localities of some Eocene whales in eastern Balochistan (Pakistan).", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 06:58:19"}
  • {"files"=>["https://ndownloader.figshare.com/files/911059"], "description"=>"<p>Sexual dimorphism of males and females is quantified for 105 species of cetaceans, pinnipeds, sirenians, and the sea otter, expressed as male-minus-female length in natural log units <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Boness1\" target=\"_blank\">[17]</a>, <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Trites1\" target=\"_blank\">[30]</a>. Territorial and harem systems that involve intense male-male competition and spatial aggregation of females have male-female dimorphism greater than about 0.15 (dashed line) <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Weckerly1\" target=\"_blank\">[31]</a>. <i>Maiacetus inuus</i> falls in the group of ‘dispersed’ mating systems with limited aggregation and limited male control of mating. Both cetaceans and pinnipeds span the entire range shown here, although mating systems are known for relatively few species <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Connor1\" target=\"_blank\">[32]</a>.</p>", "links"=>[], "tags"=>["systems", "dimorphism"], "article_id"=>581516, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g015", "stats"=>{"downloads"=>5, "page_views"=>51, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Mating_systems_and_sexual_dimorphism_in_marine_mammals_/581516", "title"=>"Mating systems and sexual dimorphism in marine mammals.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 07:08:22"}
  • {"files"=>["https://ndownloader.figshare.com/files/909688"], "description"=>"<p>Skull in ventral view (A) with interpretive shading and labels (B). Shaded components include skull bones (grey), teeth (brown), hyoids (orange), and postcranial bones (red). <i>Osteological abbreviations</i>: <i>Ba.</i>, basihyal; <i>Basicr.</i>, basicranium; <i>C</i>, cervical vertebra; <i>Dent.</i>, dentary; <i>Exoc.</i>, exoccipital; <i>Jug.</i>, jugal; <i>L</i>, left; <i>Max.</i>, maxilla; <i>R</i>, right, rib; <i>Scap.</i>, scapula; <i>Squ.</i>, squamosal; <i>St.</i>, stylohyal; <i>Stern.</i>, sternebrum; <i>T</i>, thoracic vertebra; <i>Th.</i>, thyrohyal; <i>Tym.</i>, tympanic.</p>", "links"=>[], "tags"=>["Evolutionary biology", "evolutionary biology/animal behavior", "evolutionary biology/paleontology"], "article_id"=>580143, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g005", "stats"=>{"downloads"=>1, "page_views"=>20, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Skull_of_Maiacetus_inuus_GSP_UM_3475a_female_/580143", "title"=>"Skull of <i>Maiacetus inuus</i> (GSP-UM 3475a; female).", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 07:00:21"}
  • {"files"=>["https://ndownloader.figshare.com/files/910976"], "description"=>"<p>Domestic cow (<i>Bos taurus</i>) calf (blue) in (A) ninth and final month of gestation before ‘turning’ (axially rotating), (B) turned as the birth process begins, and (C) with forelimbs and head partially extruded in the initial stage of birth (from <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Stoss1\" target=\"_blank\">[51]</a>, <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Baier1\" target=\"_blank\">[52]</a>). Harbor porpoise (<i>Phocaena phocaena</i>) (D) with full-term calf (blue) with tail partially extruded in the initial stage of birth (from <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone.0004366-Slijper1\" target=\"_blank\">[18]</a>).</p>", "links"=>[], "tags"=>["near-", "full-term", "calves", "harbor"], "article_id"=>581426, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g014", "stats"=>{"downloads"=>3, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Head_versus_tail_presentation_of_near_and_full_term_calves_in_a_domestic_cow_and_harbor_porpoise_/581426", "title"=>"Head versus tail presentation of near- and full-term calves in a domestic cow and harbor porpoise.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 07:07:52"}
  • {"files"=>["https://ndownloader.figshare.com/files/910017"], "description"=>"<p>Open areas are reconstructed. <i>Abbreviations</i>: <i>C</i>, cuneiform; <i>L</i>, lunar; <i>M</i>, magnum; <i>P</i>, pisiform; <i>S</i>, scaphoid; <i>Tr</i>, trapezium; <i>Trd</i>, trapezoid; <i>U</i>, unciform.</p>", "links"=>[], "tags"=>["carpus", "manus"], "article_id"=>580471, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g007", "stats"=>{"downloads"=>1, "page_views"=>25, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Left_radius_ulna_carpus_and_manus_of_Maiacetus_inuus_GSP_UM_3475a_female_/580471", "title"=>"Left radius, ulna, carpus and manus of <i>Maiacetus inuus</i> (GSP-UM 3475a, female).", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 07:02:17"}
  • {"files"=>["https://ndownloader.figshare.com/files/911173"], "description"=>"<p>Sexes of the <i>A. clavis</i> and <i>R. balochistanensis</i> specimens are not known.</p>*<p>Asterisk indicates estimated measurement.</p>1<p>Body weight is estimated from the centrum size of available thoracic vertebrae, using multiple regression and a reference set of 13 marine mammal skeletons of known weight (8 cetaceans and 5 pinnipeds) spanning a range of 85 to 23,500 kg (see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone-0004366-g013\" target=\"_blank\">Figure 13</a>). Weight for female <i>Maiacetus inuus</i> is estimated from male <i>M. inuus</i> in two ways: using the 12% average difference in comparable linear dimensions or the 4% average difference found for premolars and molars (<a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone-0004366-t002\" target=\"_blank\">Table 2</a>).</p>2<p>Ratio based on measurements of male and female adult specimens in <i>Maiacetus inuus</i>.</p>", "links"=>[], "tags"=>["compared", "protocetids", "eocene", "pakistan"], "article_id"=>581624, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.t001", "stats"=>{"downloads"=>6, "page_views"=>12, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Skeletal_measurements_cm_of_Maiacetus_inuus_compared_to_those_of_the_previously_known_protocetids_Artiocetus_clavis_and_Rodhocetus_balochistanensis_which_are_also_from_the_early_middle_Eocene_of_Pakistan_3_/581624", "title"=>"Skeletal measurements (cm) of <i>Maiacetus inuus</i> compared to those of the previously known protocetids <i>Artiocetus clavis</i> and <i>Rodhocetus balochistanensis</i>, which are also from the early middle Eocene of Pakistan [3].", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-02-21 07:09:00"}
  • {"files"=>["https://ndownloader.figshare.com/files/911230"], "description"=>"*<p>Asterisk indicates estimated measurement.</p>1<p>Ratios quantifying male-to-female tooth size dimorphism are provided when teeth at the same position can be compared. Median dimorphism in canine size, based on upper canines, is 1.20, meaning males have canines on the order of 20% larger than those of females. Median dimorphism in premolar and molar size is 1.04, meaning that males have premolars and molars only about 4% larger than those of females.</p>", "links"=>[], "tags"=>["teeth", "specimens"], "article_id"=>581689, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.t002", "stats"=>{"downloads"=>7, "page_views"=>16, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Measurements_mm_of_teeth_in_female_and_male_specimens_of_Maiacetus_inuus_/581689", "title"=>"Measurements (mm) of teeth in female and male specimens of <i>Maiacetus inuus</i>.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-02-21 07:09:23"}
  • {"files"=>["https://ndownloader.figshare.com/files/910408"], "description"=>"<p>Skeleton as preserved in left lateral view. Gray shading is marl matrix (see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004366#pone-0004366-g001\" target=\"_blank\">Figure 1C, D</a> for skeletal restoration).</p>", "links"=>[], "tags"=>["Evolutionary biology", "evolutionary biology/animal behavior", "evolutionary biology/paleontology"], "article_id"=>580862, "categories"=>["Evolutionary Biology", "Infectious Diseases"], "users"=>["Philip D. Gingerich", "Munir ul-Haq", "Wighart von Koenigswald", "William J. Sanders", "B. Holly Smith", "Iyad S. Zalmout"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004366.g009", "stats"=>{"downloads"=>2, "page_views"=>28, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Skeleton_of_Maiacetus_inuus_GSP_UM_3551_male_/580862", "title"=>"Skeleton of <i>Maiacetus inuus</i> (GSP-UM 3551, male).", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 07:04:38"}

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Relative Metric

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