Project 2804: J. D. Beech, J. C. Lamsdell, J. Álvaro. 2021. Phylogeny, disparity and mass extinction response in the trilobite order Harpetida. Papers in Palaeontology. 7 (4):2205-2225.
Specimen: † Bohemoharpes ungula Ebach and McNamara, 2002 (YPM/IP:559330)
View: dorsal thorax

Abstract

The trilobite order Harpetida has long been easily recognized but many unanswered evolutionary questions about the group remain. This work explores the phylogenetic relationships within Harpetida, and studies the harpetid response to the Late Ordovician mass extinction to better understand the relationship between extinction events and disparity. A discrete morphological character matrix was assembled from published descriptions and refined through first-hand observations. This matrix is the first attempt of its kind to characterize the overall morphology of Harpetida, rather than focusing on individual harpetid genera. Phylogenetic analyses under both maximum parsimony and Bayesian inference optimality criteria retrieve tree topologies that support harpetid monophyly but throw doubt onto previous hypotheses of the internal relationships of the order. Harpetid disparity proves remarkably stable over time. A modest peak in the Ordovician is followed by a slow decline throughout the Silurian and Devonian. After the Ordovician, harpetids demonstrate little or no ability to colonize new areas of morphospace. This may represent a fundamental failure to recover, where the lasting impacts of Late Ordovician mass extinction continue to suppress morphological innovation. These findings demonstrate that mass extinction events may have complex impacts that play out over many millions of years.


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Article DOI: 10.1002/spp2.1399

Project DOI: 10.7934/P2804, http://dx.doi.org/10.7934/P2804
This project containsMatrices
  • 45 Media
  • 3 Matrices
  • 2 Documents
  • 190 Taxa
  • 39 Specimens
  • 108 Characters
Total size of project's media files: 257.77M

Download Project SDD File
Total scored cells: 13218
Total media associated with cells: 0
Total labels associated with cell media: 0
Characters
Total characters: 255
Total characters with associated media: 0
Total characters with media with labels: 0
Total character states: 239
Total character states with associated media: 0
Total character states with media with labels:0
Total unordered/ordered characters:255/0
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MorphoBank Project 2804

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    Authors' Institutions

    • West Virginia University



    Members

    member name taxa specimens media media
    notes
    chars character
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    notes  media
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    cell scorings
    (scored, NPA, "-")
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    James Beech
    Project Administrator
    188384542440132008066
    (7135, 5, 926)
    01270040
    James Lamsdell
    Full membership
    21001106005152
    (4263, 0, 889)
    082000


    Taxonomic Overview for Matrix 'M24940' (47 Taxa)

    taxon unscored
    cells
    scored
    cells
    no cell
    support
    NPA
    cells
    "-" cellscell images labels on
    cell images
    member
    access
    [1] † Eoredlichia intermediata
    Taxon name last Modified on 06/28/19
    05655020002
    [2] † Baikadamaspis jikdongensis
    Taxon name last Modified on 06/28/19
    35352020002
    [3] † Bohemoharpes naumanni
    Taxon name last Modified on 06/28/19
    7636206002
    [4] † Bohemoharpes ungula
    Taxon name last Modified on 06/28/19
    2666608002
    [5] † Bowmania lassieae
    Taxon name last Modified on 06/28/19
    35353020002
    [6] † Brachyhipposiderus secundus
    Taxon name last Modified on 06/28/19
    8636115002
    [7] † Conococheaguea ovata
    Taxon name last Modified on 06/28/19
    114645019002
    [8] † Dolichoharpes dentoni
    Taxon name last Modified on 06/28/19
    1717004002
    [9] † Dubhglasina parvula
    Taxon name last Modified on 06/28/19
    8595809002
    [10] † Entomaspis radiatus
    Taxon name last Modified on 06/28/19
    36160012002
    [11] † Eoharpes benignensis
    Taxon name last Modified on 06/28/19
    1706405002
    [12] † Eoharpes primus
    Taxon name last Modified on 06/28/19
    2696905002
    [13] † Eskoharpes boltoni
    Taxon name last Modified on 06/28/19
    8625906002
    [14] † Eskoharpes guthae
    Taxon name last Modified on 06/28/19
    12585806002
    [15] † Eskoharpes palanasus
    Taxon name last Modified on 06/28/19
    2686606002
    [16] † Eskoharpes wandjina
    Taxon name last Modified on 06/28/19
    9616106002
    [17] † Globoharpes friendi
    Taxon name last Modified on 06/28/19
    8625906002
    [18] † Globoharpes teicherti
    Taxon name last Modified on 06/28/19
    8626106002
    [19] † Harpes gracilis
    Taxon name last Modified on 06/28/19
    32424002002
    [20] † Harpes macrocephalus
    Taxon name last Modified on 10/31/19
    0716605002
    [21] † Harpes neogracilis
    Taxon name last Modified on 06/28/19
    11595906002
    [22] † Harpes whidbournei
    Taxon name last Modified on 06/28/19
    10616005002
    [23] † Harpides grimmi
    Taxon name last Modified on 06/28/19
    54444225002
    [24] † Helioharpes perradiatus
    Taxon name last Modified on 06/28/19
    2686806002
    [25] † Helioharpes radians
    Taxon name last Modified on 06/28/19
    12585706002
    [26] † Heterocaryon vargum
    Taxon name last Modified on 06/28/19
    55151020002
    [27] † Hibbertia balclatchiensis
    Taxon name last Modified on 06/28/19
    9626205002
    [28] † Hibbertia wegelini
    Taxon name last Modified on 06/28/19
    9616106002
    [29] † Kielania dorbignyana
    Taxon name last Modified on 06/28/19
    11575708002
    [30] † Lioharpes montagnei
    Taxon name last Modified on 06/28/19
    12565108002
    [31] † Lioharpes venulosus
    Taxon name last Modified on 06/28/19
    8606008002
    [32] † Loganopeltis depressa
    Taxon name last Modified on 06/28/19
    54646025002
    [33] † Loganopeltoides zenkeri
    Taxon name last Modified on 06/28/19
    64747023002
    [34] † Notchpeakia milleri
    Taxon name last Modified on 01/22/19
    25454020002
    [35] † Scotoharpes consuetus
    Taxon name last Modified on 06/28/19
    10595907002
    [36] † Scotoharpes domina
    Taxon name last Modified on 06/28/19
    11595906002
    [37] † Scotoharpes latior
    Taxon name last Modified on 06/28/19
    11585507002
    [38] † Scotoharpes loma
    Taxon name last Modified on 06/28/19
    8626106002
    [39] † Scotoharpes raaschi
    Taxon name last Modified on 06/28/19
    8626106002
    [40] † Scotoharpes spasskii
    Taxon name last Modified on 06/28/19
    8615807002
    [41] † Scotoharpes tatouyangensis
    Taxon name last Modified on 06/28/19
    16535016002
    [42] † Cedaria minor
    Taxon name last Modified on 06/28/19
    05555021002
    [43] † Coosina kindlei
    Taxon name last Modified on 06/28/19
    104746019002
    [44] † Crepicephalus iowensis
    Taxon name last Modified on 06/28/19
    94545022002
    [45] † Modocia whiteleyi
    Taxon name last Modified on 06/28/19
    15454021002
    [46] † Ptychoparia striata
    Taxon name last Modified on 06/28/19
    15454120002
    [47] † Tricrepicephalus coria
    Taxon name last Modified on 06/28/19
    44948023002


    Taxonomic Overview for Matrix 'M25898' (53 Taxa)

    taxon unscored
    cells
    scored
    cells
    no cell
    support
    NPA
    cells
    "-" cellscell images labels on
    cell images
    member
    access
    [1] † Eoredlichia intermediata
    Taxon name last Modified on 06/28/19
    97371022002
    [2] † Baikadamaspis jikdongensis
    Taxon name last Modified on 06/28/19
    147067020002
    [3] † Bohemoharpes naumanni
    Taxon name last Modified on 06/28/19
    345755012002
    [4] † Bohemoharpes ungula
    Taxon name last Modified on 06/28/19
    78682012002
    [5] † Bowmania lassieae
    Taxon name last Modified on 06/28/19
    464545012002
    [6] † Brachyhipposiderus secundus
    Taxon name last Modified on 06/28/19
    474645010002
    [7] † Chencunia tripleura
    Taxon name last Modified on 02/22/19
    967700002
    [8] † Conococheaguea ovata
    Taxon name last Modified on 06/28/19
    255554023002
    [9] † Dolichoharpes dentoni
    Taxon name last Modified on 06/28/19
    21787704002
    [10] † Dubhglasina parvula
    Taxon name last Modified on 06/28/19
    524140011002
    [11] † Entomaspis radiatus
    Taxon name last Modified on 06/28/19
    206462020002
    [12] † Eoharpes antiquatus
    Taxon name last Modified on 02/22/19
    21766607002
    [13] † Eoharpes benignensis
    Taxon name last Modified on 06/28/19
    247065010002
    [14] † Eoharpes primus
    Taxon name last Modified on 06/28/19
    38565609002
    [15] † Eskoharpes boltoni
    Taxon name last Modified on 06/28/19
    28716407002
    [16] † Eskoharpes guthae
    Taxon name last Modified on 06/28/19
    35605809002
    [17] † Eskoharpes palanasus
    Taxon name last Modified on 06/28/19
    19787309002
    [18] † Eskoharpes wandjina
    Taxon name last Modified on 06/28/19
    226260021002
    [19] † Globoharpes friendi
    Taxon name last Modified on 06/28/19
    27676509002
    [20] † Globoharpes teicherti
    Taxon name last Modified on 06/28/19
    30696606002
    [21] † Harpes gracilis
    Taxon name last Modified on 06/28/19
    563229015002
    [22] † Harpes macrocephalus
    Taxon name last Modified on 10/31/19
    26716406002
    [23] † Harpes neogracilis
    Taxon name last Modified on 06/28/19
    266159018002
    [24] † Harpes whidbournei
    Taxon name last Modified on 06/28/19
    275756019002
    [25] † Harpides grimmi
    Taxon name last Modified on 06/28/19
    72282703002
    [26] † Helioharpes perradiatus
    Taxon name last Modified on 06/28/19
    286161014002
    [27] † Helioharpes radians
    Taxon name last Modified on 06/28/19
    295655018002
    [28] † Heterocaryon vargum
    Taxon name last Modified on 06/28/19
    186260025002
    [29] † Hibbertia balclatchiensis
    Taxon name last Modified on 06/28/19
    46515106002
    [30] † Hibbertia ottawaensis
    Taxon name last Modified on 02/22/19
    84171702002
    [31] † Hibbertia wegelini
    Taxon name last Modified on 06/28/19
    28686807002
    [32] † Kielania dorbignyana
    Taxon name last Modified on 06/28/19
    375555011002
    [33] † Lioharpes montagnei
    Taxon name last Modified on 06/28/19
    226861013002
    [34] † Lioharpes venulosus
    Taxon name last Modified on 06/28/19
    21747408002
    [35] † Loganopeltis depressa
    Taxon name last Modified on 06/28/19
    20777508002
    [36] † Loganopeltoides zenkeri
    Taxon name last Modified on 06/28/19
    414946015002
    [37] Notchpeakia milleri
    Taxon name last Modified on 06/28/19
    157271016002
    [38] † Scotoharpes consuetus
    Taxon name last Modified on 06/28/19
    444646013002
    [39] † Scotoharpes domina
    Taxon name last Modified on 06/28/19
    395252012002
    [40] † Scotoharpes judex
    Taxon name last Modified on 02/22/19
    23727208002
    [41] † Scotoharpes latior
    Taxon name last Modified on 06/28/19
    31646009002
    [42] † Scotoharpes loma
    Taxon name last Modified on 06/28/19
    22757506002
    [43] † Scotoharpes raaschi
    Taxon name last Modified on 06/28/19
    38616004002
    [44] † Scotoharpes singularis
    Taxon name last Modified on 02/22/19
    474040016002
    [45] † Scotoharpes spasskii
    Taxon name last Modified on 06/28/19
    494441011002
    [46] † Scotoharpes tatouyangensis
    Taxon name last Modified on 06/28/19
    43535107002
    [47] † Scotoharpes telleri
    Taxon name last Modified on 02/22/19
    44505009002
    [48] † Cedaria minor
    Taxon name last Modified on 06/28/19
    146363026002
    [49] † Coosina kindlei
    Taxon name last Modified on 06/28/19
    275653021002
    [50] † Crepicephalus iowensis
    Taxon name last Modified on 06/28/19
    265352025002
    [51] † Modocia whiteleyi
    Taxon name last Modified on 06/28/19
    96464030002
    [52] † Ptychoparia striata
    Taxon name last Modified on 06/28/19
    97773018002
    [53] † Tricrepicephalus coria
    Taxon name last Modified on 06/28/19
    374442023002


    Taxonomic Overview for Matrix 'M26129' (86 Taxa)

    taxon unscored
    cells
    scored
    cells
    no cell
    support
    NPA
    cells
    "-" cellscell images labels on
    cell images
    member
    access
    [1] † Baikadamaspis jikdongensis
    Taxon name last Modified on 06/28/19
    35352020002
    [2] † Bohemoharpes naumanni
    Taxon name last Modified on 06/28/19
    7636206002
    [3] † Bohemoharpes ungula
    Taxon name last Modified on 06/28/19
    2666608002
    [4] † Bowmania lassieae
    Taxon name last Modified on 06/28/19
    35353020002
    [5] † Brachyhipposiderus secundus
    Taxon name last Modified on 06/28/19
    9636105002
    [6] † Conococheaguea ovata
    Taxon name last Modified on 06/28/19
    114645019002
    [7] † Dolichoharpes dentoni
    Taxon name last Modified on 06/28/19
    1717004002
    [8] † Dubhglasina parvula
    Taxon name last Modified on 06/28/19
    8595809002
    [9] † Entomaspis radiatus
    Taxon name last Modified on 06/28/19
    36160012002
    [10] † Eoharpes benignensis
    Taxon name last Modified on 06/28/19
    1706405002
    [11] † Eoharpes primus
    Taxon name last Modified on 06/28/19
    2696905002
    [12] † Eskoharpes boltoni
    Taxon name last Modified on 06/28/19
    8625906002
    [13] † Eskoharpes guthae
    Taxon name last Modified on 06/28/19
    12585806002
    [14] † Eskoharpes palanasus
    Taxon name last Modified on 06/28/19
    2686606002
    [15] † Eskoharpes wandjina
    Taxon name last Modified on 06/28/19
    9616106002
    [16] † Globoharpes friendi
    Taxon name last Modified on 06/28/19
    8625906002
    [17] † Globoharpes teicherti
    Taxon name last Modified on 06/28/19
    8626106002
    [18] † Harpes gracilis
    Taxon name last Modified on 06/28/19
    32424002002
    [19] † Harpes macrocephalus
    Taxon name last Modified on 10/31/19
    0716605002
    [20] † Harpes neogracilis
    Taxon name last Modified on 06/28/19
    11595906002
    [21] † Harpes whidbournei
    Taxon name last Modified on 06/28/19
    10616005002
    [22] † Harpides grimmi
    Taxon name last Modified on 06/28/19
    74444025002
    [23] † Helioharpes perradiatus
    Taxon name last Modified on 06/28/19
    2686806002
    [24] † Helioharpes radians
    Taxon name last Modified on 06/28/19
    12585706002
    [25] † Heterocaryon vargum
    Taxon name last Modified on 06/28/19
    55151020002
    [26] † Hibbertia balclatchiensis
    Taxon name last Modified on 06/28/19
    9626205002
    [27] † Hibbertia wegelini
    Taxon name last Modified on 06/28/19
    9616106002
    [28] † Kielania dorbignyana
    Taxon name last Modified on 06/28/19
    11575708002
    [29] † Lioharpes montagnei
    Taxon name last Modified on 06/28/19
    12565108002
    [30] † Lioharpes venulosus
    Taxon name last Modified on 06/28/19
    8606008002
    [31] † Loganopeltis depressa
    Taxon name last Modified on 06/28/19
    54646025002
    [32] † Loganopeltoides zenkeri
    Taxon name last Modified on 06/28/19
    64747023002
    [33] Notchpeakia milleri
    Taxon name last Modified on 06/28/19
    25454020002
    [34] † Scotoharpes consuetus
    Taxon name last Modified on 06/28/19
    10595907002
    [35] † Scotoharpes domina
    Taxon name last Modified on 06/28/19
    11595906002
    [36] † Scotoharpes latior
    Taxon name last Modified on 06/28/19
    11585507002
    [37] † Scotoharpes loma
    Taxon name last Modified on 06/28/19
    8626106002
    [38] † Scotoharpes raaschi
    Taxon name last Modified on 06/28/19
    8626106002
    [39] † Scotoharpes spasskii
    Taxon name last Modified on 06/28/19
    8615807002
    [40] † Scotoharpes tatouyangensis
    Taxon name last Modified on 06/28/19
    17535006002
    [41] † Cedaria minor
    Taxon name last Modified on 06/28/19
    05555021002
    [42] † Coosina kindlei
    Taxon name last Modified on 06/28/19
    104746019002
    [43] † Crepicephalus iowensis
    Taxon name last Modified on 06/28/19
    94545022002
    [44] † Modocia whiteleyi
    Taxon name last Modified on 06/28/19
    15454021002
    [45] † Ptychoparia striata
    Taxon name last Modified on 06/28/19
    25454020002
    [46] † Tricrepicephalus coria
    Taxon name last Modified on 06/28/19
    44948023002
    [47] † Ancestor_1
    Taxon name last Modified on 06/28/19
    0686808002
    [48] † Ancestor_2
    Taxon name last Modified on 06/28/19
    0717105002
    [49] † Ancestor_3
    Taxon name last Modified on 06/28/19
    0707006002
    [50] Ancestor_4
    Taxon name last Modified on 06/28/19
    0727204002
    [51] † Ancestor_5
    Taxon name last Modified on 06/28/19
    0727204002
    [52] † Ancestor_6
    Taxon name last Modified on 06/28/19
    0717105002
    [53] † Ancestor_7
    Taxon name last Modified on 06/28/19
    0737303002
    [54] † Ancestor_8
    Taxon name last Modified on 06/28/19
    0747402002
    [55] † Ancestor_9
    Taxon name last Modified on 06/28/19
    0737303002
    [56] † Ancestor_10
    Taxon name last Modified on 06/28/19
    0747402002
    [57] † Ancestor_11
    Taxon name last Modified on 06/28/19
    0737303002
    [58] † Ancestor_12
    Taxon name last Modified on 06/28/19
    0747402002
    [59] † Ancestor_13
    Taxon name last Modified on 06/28/19
    0737303002
    [60] † Ancestor_14
    Taxon name last Modified on 06/28/19
    0747402002
    [61] † Ancestor_15
    Taxon name last Modified on 06/28/19
    0737303002
    [62] † Ancestor_16
    Taxon name last Modified on 06/28/19
    0737303002
    [63] † Ancestor_17
    Taxon name last Modified on 06/28/19
    0747402002
    [64] † Ancestor_18
    Taxon name last Modified on 06/28/19
    0737303002
    [65] † Ancestor_19
    Taxon name last Modified on 06/28/19
    0747402002
    [66] † Ancestor_20
    Taxon name last Modified on 06/28/19
    0757501002
    [67] † Ancestor_21
    Taxon name last Modified on 06/28/19
    0757501002
    [68] † Ancestor_22
    Taxon name last Modified on 06/28/19
    0757501002
    [69] † Ancestor_23
    Taxon name last Modified on 06/28/19
    0757501002
    [70] † Ancestor_24
    Taxon name last Modified on 06/28/19
    0757501002
    [71] † Ancestor_25
    Taxon name last Modified on 06/28/19
    0737303002
    [72] † Ancestor_26
    Taxon name last Modified on 06/28/19
    0747402002
    [73] † Ancestor_27
    Taxon name last Modified on 06/28/19
    0757501002
    [74] † Ancestor_28
    Taxon name last Modified on 06/28/19
    0757501002
    [75] † Ancestor_29
    Taxon name last Modified on 06/28/19
    0757501002
    [76] † Ancestor_30
    Taxon name last Modified on 06/28/19
    0767600002
    [77] † Ancestor_31
    Taxon name last Modified on 06/28/19
    0767600002
    [78] † Ancestor_32
    Taxon name last Modified on 06/28/19
    0767600002
    [79] † Ancestor_33
    Taxon name last Modified on 06/28/19
    0727204002
    [80] † Ancestor_34
    Taxon name last Modified on 06/28/19
    0727204002
    [81] † Ancestor_35
    Taxon name last Modified on 06/28/19
    0727204002
    [82] † Ancestor_36
    Taxon name last Modified on 06/28/19
    0727204002
    [83] † Ancestor_37
    Taxon name last Modified on 06/28/19
    1727203002
    [84] † Ancestor_38
    Taxon name last Modified on 06/28/19
    0737303002
    [85] † Ancestor_39
    Taxon name last Modified on 06/28/19
    1717104002
    [86] † Eoredlichia intermediata
    Taxon name last Modified on 06/28/19
    05655020002


    Project downloads

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    Total downloads from project141
    Project downloads58
    Document downloads14Appendix S2. Illustrated character list. (12 downloads); Appendix S1: Characters for Phylogenetic Analysis (2 downloads);
    Matrix downloads26Appendix S3. Character-taxon matrix. (11 downloads); Character Matrix 1 (11 downloads); Backup matrix (4 downloads);
    Media downloads43M565519 (1 download); M565520 (1 download); M565521 (3 downloads); M565523 (3 downloads); M565524 (3 downloads); M565525 (1 download); M565528 (2 downloads); M565527 (1 download); M565530 (1 download); M565531 (1 download); M565533 (1 download); M565534 (1 download); M595169 (2 downloads); M595186 (1 download); M595172 (1 download); M595173 (1 download); M595188 (3 downloads); M595174 (1 download); M565506 (1 download); M565508 (2 downloads); M565509 (1 download); M565510 (1 download); M565514 (1 download); M565515 (1 download); M565522 (1 download); M565526 (1 download); M586603 (1 download); M595166 (1 download); M595167 (1 download); M595170 (1 download); M651735 (2 downloads);