Torosaurus Is Not Triceratops: Ontogeny in Chasmosaurine Ceratopsids as a Case Study in Dinosaur Taxonomy
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{"title"=>"Torosaurus is not triceratops: Ontogeny in chasmosaurine ceratopsids as a case study in dinosaur taxonomy", "type"=>"journal", "authors"=>[{"first_name"=>"Nicholas R.", "last_name"=>"Longrich", "scopus_author_id"=>"13806457100"}, {"first_name"=>"Daniel J.", "last_name"=>"Field", "scopus_author_id"=>"36170492900"}], "year"=>2012, "source"=>"PLoS ONE", "identifiers"=>{"pui"=>"364355321", "sgr"=>"84857667102", "issn"=>"19326203", "pmid"=>"22393425", "scopus"=>"2-s2.0-84857667102", "doi"=>"10.1371/journal.pone.0032623", "isbn"=>"1932-6203"}, "id"=>"c27b1213-0023-3f82-87c0-cf3274572593", "abstract"=>"BACKGROUND: In horned dinosaurs, taxonomy is complicated by the fact that the cranial ornament that distinguishes species changes with age. Based on this observation, it has been proposed that the genera Triceratops and Torosaurus are in fact synonymous, with specimens identified as Torosaurus representing the adult form of Triceratops. The hypothesis of synonymy makes three testable predictions: 1) the species in question should have similar geographic and stratigraphic distributions, 2) specimens assigned to Torosaurus should be more mature than those assigned to Triceratops, and 3) intermediates should exist that combine features of Triceratops and Torosaurus. The first condition appears to be met, but it remains unclear whether the other predictions are borne out by the fossil evidence.\\n\\nMETHODOLOGY/PRINCIPAL FINDINGS: We assessed the relative maturity of Torosaurus and Triceratops specimens by coding skulls for characters that vary with maturity, and then using a clustering analysis to arrange them into a growth series. We found that a well-defined sequence of changes exists in horned dinosaurs: development of cranial ornament occurs in juveniles, followed by fusion of the skull roof in subadults, and finally, the epoccipitals, epijugals, and rostral fuse to the skull in adults. Using this scheme, we identified mature and immature individuals of both Torosaurus and Triceratops. Furthermore, we describe the ventral depressions on the frill of Triceratops, and show that they differ in shape and position from the parietal fenestrae of Torosaurus. Thus, we conclude that these structures are not intermediates between the solid frill of Triceratops and the fenestrated frill of Torosaurus.\\n\\nCONCLUSIONS/SIGNIFICANCE: Torosaurus is a distinct genus of horned dinosaur, not the adult of Triceratops. Our method provides a framework for assessing the hypothesis of synonymy through ontogeny in the fossil record.", "link"=>"http://www.mendeley.com/research/torosaurus-not-triceratops-ontogeny-chasmosaurine-ceratopsids-case-study-dinosaur-taxonomy", "reader_count"=>84, "reader_count_by_academic_status"=>{"Unspecified"=>1, "Professor > Associate Professor"=>1, "Librarian"=>1, "Researcher"=>19, "Student > Doctoral Student"=>5, "Student > Ph. D. Student"=>14, "Student > Postgraduate"=>2, "Other"=>7, "Student > Master"=>16, "Student > Bachelor"=>15, "Lecturer"=>1, "Lecturer > Senior Lecturer"=>1}, "reader_count_by_user_role"=>{"Unspecified"=>1, "Professor > Associate Professor"=>1, "Librarian"=>1, "Researcher"=>19, "Student > Doctoral Student"=>5, "Student > Ph. D. Student"=>14, "Student > Postgraduate"=>2, "Other"=>7, "Student > Master"=>16, "Student > Bachelor"=>15, "Lecturer"=>1, "Lecturer > Senior Lecturer"=>1}, "reader_count_by_subject_area"=>{"Unspecified"=>1, "Medicine and Dentistry"=>1, "Agricultural and Biological Sciences"=>38, "Philosophy"=>1, "Physics and Astronomy"=>1, "Social Sciences"=>2, "Computer Science"=>2, "Immunology and Microbiology"=>1, "Earth and Planetary Sciences"=>35, "Economics, Econometrics and Finance"=>1}, "reader_count_by_subdiscipline"=>{"Medicine and Dentistry"=>{"Medicine and Dentistry"=>1}, "Social Sciences"=>{"Social Sciences"=>2}, "Physics and Astronomy"=>{"Physics and Astronomy"=>1}, "Immunology and Microbiology"=>{"Immunology and Microbiology"=>1}, "Earth and Planetary Sciences"=>{"Earth and Planetary Sciences"=>35}, "Economics, Econometrics and Finance"=>{"Economics, Econometrics and Finance"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>38}, "Computer Science"=>{"Computer Science"=>2}, "Unspecified"=>{"Unspecified"=>1}, "Philosophy"=>{"Philosophy"=>1}}, "reader_count_by_country"=>{"Netherlands"=>1, "United States"=>2, "Japan"=>1, "Italy"=>2, "United Kingdom"=>2, "Chile"=>1, "Germany"=>1, "Spain"=>2}, "group_count"=>1}

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/344915", "https://ndownloader.figshare.com/files/344962", "https://ndownloader.figshare.com/files/345033"], "description"=>"<div><h3>Background</h3><p>In horned dinosaurs, taxonomy is complicated by the fact that the cranial ornament that distinguishes species changes with age. Based on this observation, it has been proposed that the genera <em>Triceratops</em> and <em>Torosaurus</em> are in fact synonymous, with specimens identified as <em>Torosaurus</em> representing the adult form of <em>Triceratops</em>. The hypothesis of synonymy makes three testable predictions: 1) the species in question should have similar geographic and stratigraphic distributions, 2) specimens assigned to <em>Torosaurus</em> should be more mature than those assigned to <em>Triceratops</em>, and 3) intermediates should exist that combine features of <em>Triceratops</em> and <em>Torosaurus</em>. The first condition appears to be met, but it remains unclear whether the other predictions are borne out by the fossil evidence.</p> <h3>Methodology/Principal Findings</h3><p>We assessed the relative maturity of <em>Torosaurus</em> and <em>Triceratops</em> specimens by coding skulls for characters that vary with maturity, and then using a clustering analysis to arrange them into a growth series. We found that a well-defined sequence of changes exists in horned dinosaurs: development of cranial ornament occurs in juveniles, followed by fusion of the skull roof in subadults, and finally, the epoccipitals, epijugals, and rostral fuse to the skull in adults. Using this scheme, we identified mature and immature individuals of both <em>Torosaurus</em> and <em>Triceratops</em>. Furthermore, we describe the ventral depressions on the frill of <em>Triceratops</em>, and show that they differ in shape and position from the parietal fenestrae of <em>Torosaurus</em>. Thus, we conclude that these structures are not intermediates between the solid frill of <em>Triceratops</em> and the fenestrated frill of <em>Torosaurus</em>.</p> <h3>Conclusions/Significance</h3><p><em>Torosaurus</em> is a distinct genus of horned dinosaur, not the adult of <em>Triceratops</em>. Our method provides a framework for assessing the hypothesis of synonymy through ontogeny in the fossil record.</p> </div>", "links"=>[], "tags"=>["ontogeny", "chasmosaurine", "ceratopsids", "dinosaur", "taxonomy"], "article_id"=>128193, "categories"=>["Physiology", "Evolutionary Biology"], "users"=>["Nicholas R. Longrich", "Daniel J. Field"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0032623.s001", "https://dx.doi.org/10.1371/journal.pone.0032623.s002", "https://dx.doi.org/10.1371/journal.pone.0032623.s003"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Torosaurus_Is_Not_Triceratops_Ontogeny_in_Chasmosaurine_Ceratopsids_as_a_Case_Study_in_Dinosaur_Taxonomy/128193", "title"=>"<em>Torosaurus</em> Is Not <em>Triceratops</em>: Ontogeny in Chasmosaurine Ceratopsids as a Case Study in Dinosaur Taxonomy", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2012-02-29 02:16:33"}
  • {"files"=>["https://ndownloader.figshare.com/files/674152"], "description"=>"<p>A, <i>Triceratops prorsus</i> YPM 1822 and B, <i>Torosaurus latus</i> ANSP 15192. <i>Triceratops</i> is characterized by a short frill with a flat squamosal, an upturned caudal margin of the frill, the absence of fenestrae, and a midline epiparietal. <i>Torosaurus</i> is characterized by an elongate frill with a straighter edge, a concave squamosal, and lack of upturning of the frill.</p>", "links"=>[], "tags"=>["physiology", "Evolutionary biology"], "article_id"=>344647, "categories"=>["Physiology", "Evolutionary Biology"], "users"=>["Nicholas R. Longrich", "Daniel J. Field"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0032623.g001"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Torosaurus_and_Triceratops_compared_/344647", "title"=>"<i>Torosaurus</i> and <i>Triceratops</i> compared.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-02-29 01:17:27"}
  • {"files"=>["https://ndownloader.figshare.com/files/674214"], "description"=>"<p>1, Scollard Formation, Alberta; 2, Frenchman Formation, Saskatchewan, 3, Hell Creek Formation, Montana; 4, Hell Creek Formation, North Dakota; 5, Hell Creek Formation, South Dakota; 6, Lance Formation, Wyoming; 7, Denver Formation, Colorado; 8, North Horn Formation, Utah; 9, Javelina Formation, Texas.</p>", "links"=>[], "tags"=>["physiology", "Evolutionary biology"], "article_id"=>344708, "categories"=>["Physiology", "Evolutionary Biology"], "users"=>["Nicholas R. Longrich", "Daniel J. Field"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0032623.g002"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Distribution_of_Torosaurus_and_Triceratops_/344708", "title"=>"Distribution of <i>Torosaurus</i> and <i>Triceratops</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-02-29 01:18:28"}
  • {"files"=>["https://ndownloader.figshare.com/files/674314"], "description"=>"<p>A, <i>Triceratops prorsus</i> YPM 1823; B, <i>Torosaurus latus</i> YPM 1830. Abbreviations: fr, frontal; lac, lacrimal; nas, nasal; pos, postorbital; pre, prefrontal.</p>", "links"=>[], "tags"=>["physiology", "Evolutionary biology"], "article_id"=>344808, "categories"=>["Physiology", "Evolutionary Biology"], "users"=>["Nicholas R. Longrich", "Daniel J. Field"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0032623.g003"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Fusion_of_the_skull_roof_in_Chasmosaurinae_/344808", "title"=>"Fusion of the skull roof in Chasmosaurinae.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-02-29 01:20:08"}
  • {"files"=>["https://ndownloader.figshare.com/files/674441"], "description"=>"<p><b>A</b>, <i>Triceratops horridus</i> USNM 1201; B, <i>Triceratops prorsus</i> YPM 1822. Abbreviations: epn, epinasal; nas, nasal; pmx, premaxilla; ros, rostral.</p>", "links"=>[], "tags"=>["rostrum"], "article_id"=>344936, "categories"=>["Physiology", "Evolutionary Biology"], "users"=>["Nicholas R. Longrich", "Daniel J. Field"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0032623.g004"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Fusion_of_the_rostrum_in_Chasmosaurinae_/344936", "title"=>"Fusion of the rostrum in Chasmosaurinae.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-02-29 01:22:16"}
  • {"files"=>["https://ndownloader.figshare.com/files/674514"], "description"=>"<p>Inferred sequence of ontogenetic character changes is mapped onto the diagram. An initial run found that MNHN 1912.20, YPM 1830, YPM 1828 and USNM 5740 are adults and that USNM 15583 was immature. Due to missing data these specimens caused a lack of resolution and a second run of the analysis was conducted excluding these specimens. Asterisks indicate character change mappings that are provisional owing to missing or conflicting data.</p>", "links"=>[], "tags"=>["diagram", "specimens", "arranged"], "article_id"=>345009, "categories"=>["Physiology", "Evolutionary Biology"], "users"=>["Nicholas R. Longrich", "Daniel J. Field"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0032623.g005"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Clustering_diagram_with_Triceratops_and_Torosaurus_specimens_arranged_in_developmental_sequence_/345009", "title"=>"Clustering diagram with <i>Triceratops</i> and <i>Torosaurus</i> specimens arranged in developmental sequence.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-02-29 01:23:29"}
  • {"files"=>["https://ndownloader.figshare.com/files/674656"], "description"=>"<p>A, lateral view; B, dorsal view. Abbreviations: epn, epinasal; max, maxilla; nas, nasal; pmx, premaxilla; pos, postorbital horncore, rec, reconstruction; rgr, rostral groove.</p>", "links"=>[], "tags"=>["ansp"], "article_id"=>345155, "categories"=>["Physiology", "Evolutionary Biology"], "users"=>["Nicholas R. Longrich", "Daniel J. Field"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0032623.g006"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Immature_features_in_Torosaurus_latus_ANSP_15192_a_young_adult_/345155", "title"=>"Immature features in <i>Torosaurus latus</i> ANSP 15192, a young adult.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-02-29 01:25:55"}
  • {"files"=>["https://ndownloader.figshare.com/files/674760"], "description"=>"<p>A, unfused epijugal. A1, anterior view, A2, lateral view; A3, medial view. B, unfused rostral. B1, dorsal view; B2, ventral view. C, occipital condyle formed of unfused exoccipitals and basioccipital. D, caudal margin of parietal showing rounded margin where unfused epoccipitals attach. E, dorsal surface of parietal showing striated surface texture. Scales = 50 mm for A and B, 20 mm for E.</p>", "links"=>[], "tags"=>["ypm"], "article_id"=>345256, "categories"=>["Physiology", "Evolutionary Biology"], "users"=>["Nicholas R. Longrich", "Daniel J. Field"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0032623.g007"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Immature_features_in_Torosaurus_latus_YPM_1831_/345256", "title"=>"Immature features in <i>Torosaurus latus</i> YPM 1831.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-02-29 01:27:36"}
  • {"files"=>["https://ndownloader.figshare.com/files/674844"], "description"=>"<p>A, YPM 1831, a subadult, skull length >2.6 m, and B, ANSP 15192, an early adult, skull length 1.8 m. The rostrum of YPM 1831 is reconstructed and would probably have been slightly longer, as in ANSP 15192.</p>", "links"=>[], "tags"=>["physiology", "Evolutionary biology"], "article_id"=>345339, "categories"=>["Physiology", "Evolutionary Biology"], "users"=>["Nicholas R. Longrich", "Daniel J. Field"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0032623.g008"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Comparison_of_the_size_of_Torosaurus_latus_specimens_/345339", "title"=>"Comparison of the size of <i>Torosaurus latus</i> specimens.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-02-29 01:28:59"}
  • {"files"=>["https://ndownloader.figshare.com/files/674904"], "description"=>"<p>A, <i>Triceratops</i> YPM 1823; B, <i>Torosaurus</i> ANSP 15192. Abbreviations: fen, fenestra; fos, parietal-squamosal fossa; par, parietal; sq, squamosal. P0, midline epiparietal; p1–p5 epiparietals 1–5; eps, epoccipital crossing the parietal-squamosal suture.</p>", "links"=>[], "tags"=>["frills", "parietal", "fenestrae", "fossae"], "article_id"=>345400, "categories"=>["Physiology", "Evolutionary Biology"], "users"=>["Nicholas R. Longrich", "Daniel J. Field"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0032623.g009"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Comparison_of_frills_in_Torosaurus_and_Triceratops_showing_the_different_position_of_the_parietal_fenestrae_of_Torosaurus_and_parietal_fossae_of_Triceratops_/345400", "title"=>"Comparison of frills in <i>Torosaurus</i> and <i>Triceratops</i>, showing the different position of the parietal fenestrae of <i>Torosaurus</i> and parietal fossae of <i>Triceratops</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-02-29 01:30:00"}

PMC Usage Stats | Further Information

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Relative Metric

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