Hadrosauroidea

Skeletal mount of Probactrosaurus gobiensis, copyright Greg Goebel.


Belongs within: Iguanodontia.
Contains: Hadrosauridae.

The name Hadrosauroidea has been used for a clade uniting the Hadrosauridae and Ouranosaurus; it is here used for the smallest clade uniting these two taxa as found in the phylogenetic analysis by Norman (2004). Supporting characters for this clade were not explicitly listed.

<==Hadrosauroidea
    |  i. s.: Jinzhousaurus Wang & Xu 2001 N04
    |           `--J. yangi Wang & Xu 2001 N04
    |--Ouranosaurus Taquet 1976 N04
    |    `--O. nigeriensis Taquet 1976 N04
    `--+--+--Eolambia Kirkland 1998 N04
       |  |    `--E. caroljonesa Kirkland 1998 N04
       |  `--Altirhinus Norman 1998 N04
       |       `--A. kurzanovi Norman 1998 N04 [incl. Iguanodon orientalis D07]
       `--+--Protohadros Head 1998 N04
          |    `--P. byrdi Head 1998 N04
          `--+--+--Hadrosauridae N04
             |  `--Nanyangosaurus Xu, Zhao et al. 2000 XZ00
             |       `--*N. zhugeii Xu, Zhao et al. 2000 XZ00
             `--Probactrosaurus Rozhdestvensky 1966 N04
                  |--P. alashanicus Rozhdestvensky 1966 N04
                  |--P. gobiensis Rozhdestvensky 1966 N04
                  `--P. mazongshanensis Lü 1997 N04

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[N04] Norman, D. B. 2004. Basal Iguanodontia. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2nd ed. pp. 413–437. University of California Press: Berkeley.

[XZ00] Xu X., Zhao X.-J., Lü J.-C., Huang W.-B., Li Z.-Y. & Dong Z.-M. 2000. A new iguanodontian from Sangping Formation of Neixiang, Henan and its stratigraphical implication. Vertebrata PalAsiatica 38 (3): 176–191.

Iguanodontia

Reconstruction of Iguanodon atherfieldensis, copyright Steveoc 86.


Belongs within: Ornithopoda.
Contains: Hadrosauroidea.

The Iguanodontia are a diverse group of bipedal and facultatively quadrupedal herbivorous dinosaurs known from the Late Jurassic and Cretaceous, recognised by Norman (2004) as the clade of all taxa closer to Edmontosaurus than Thescelosaurus.

Characters (from Norman 2004): Premaxilla with transversely expanded and edentulous margin; predentary with smoothly convex rostral margin in dorsal aspect; dentary ramus deep, with parallel dorsal and ventral borders; sternal rib ossification absent; one phalanx lost on manus digit III; prepubic process compressed, blade-shaped.

Iguanodontia [Iguanodontidae]
    |  i. s.: Albisaurus scutifer Fritsch 1905 (n. d.) N04
    |         Loncosaurus argentinus Ameghino 1898 (n. d.) N04
    |         Tichosteus (n. d.) N04
    |           |--T. aequifacies Cope 1877 (n. d.) N04
    |           `--T. lucasanus Cope 1877 (n. d.) N04
    |--Tenontosaurus Ostrom 1970 N04
    |    |--T. dossi Winkler, Murry & Jacobs 1997 N04
    |    `--T. tilletti Ostrom 1970 N04
    `--+--Muttaburrasaurus Bartholomai & Molnar 1981 N04
       |    `--M. langdoni Bartholomai & Molnar 1981 N04
       `--Dryomorpha N04
            |  i. s.: Callovosaurus Galton 1980 (n. d.) N85, N04
            |           `--*C. leedsi (Lydekker 1889) [=Camptosaurus leedsi] N04
            |         Kangnasaurus coetzeei Haughton 1915 (n. d.) N85, N04
            |--Dryosauridae N04
            |    |--Planicoxa DiCroce & Carpenter 2001 N04
            |    |    `--P. venenica DiCroce & Carpenter 2001 N04
            |    |--Dryosaurus Marsh 1894 [incl. Dysalotosaurus Virchow 1919] N04
            |    |    |--D. altus (Marsh 1878) [=Laosaurus altus] N04
            |    |    `--D. lettowvorbecki (Virchow 1919) [=Dysalotosaurus lettowvorbecki] N04
            |    `--Valdosaurus Galton 1977 N04
            |         |--V. canaliculatus (Galton 1975) (see below for synonymy) N04
            |         `--V. nigeriensis Galton & Taquet 1982 N04
            `--Ankylopollexia N04
                 |  i. s.: Draconyx Mateus & Antunes 2001 N04
                 |           `--D. loureiroi Mateus & Antunes 2001 N04
                 |--Camptosaurus Marsh 1885 (see below for synonymy) N04
                 |    |--C. amplus (Marsh 1879) [=Camptonotus amplus] N04
                 |    |--C. depressus Gilmore 1909 N04
                 |    |--C. dispar (Marsh 1879) (see below for synonymy) N04
                 |    |--C. hoggi (Owen 1874) [=Iguanodon hoggi] N04
                 |    `--C. prestwichii (Hulke 1880) [=Iguanodon prestwichii] N04
                 `--Styracosterna N04
                      |--Lurdusaurus Taquet & Russell 1999 [incl. Gravisaurus Chabli 1988] N04
                      |    `--L. arenatus Taquet & Russell 1999 [incl. Gravisaurus tenerensis Chabli 1988] N04
                      |--Equijubus You, Luo et al. 2003 N04
                      |    `--E. normani You, Luo et al. 2003 N04
                      `--Iguanodontoidea [Hadrosauriformes] N04
                           |  i. s.: Craspedodon Dollo 1883 N04
                           |           `--C. lonzeensis Dollo 1883 N04
                           |         Fukuisaurus Kobayashi & Azuma 2003 N04
                           |           `--F. tetoriensius Kobayashi & Azuma 2003 N04
                           |         Shuangmiaosaurus You, Ji et al. 2003 N04
                           |           `--S. gilmorei You, Ji et al. 2003 N04
                           |--Hadrosauroidea N04
                           `--Iguanodon Mantell 1825 [incl. Heterosaurus Cornuel 1850, Iguanosaurus Ritgen 1828] N04
                                |--*I. anglicus Holl 1829 WU03, N04 (see below for synonymy)
                                |--I. atherfieldensis Hooley 1924 (see below for synonymy) N04
                                |--I. bernissartensis Boulenger 1881 [incl. I. seelyi Hulke 1882] N04
                                |--I. dawsoni Lydekker 1888 N04
                                |--I. exogirarus Fritsch 1878 (n. d.) N04
                                |--I. fittoni Lydekker 1889 [incl. I. hollingtoniensis Lydekker 1889] N04
                                |--I. lakotaensis Weishampel & Bjork 1989 N04
                                `--I. ottingeri Galton & Jensen 1979 (n. d.) N04

Camptosaurus Marsh 1885 [incl. Brachyrophus Cope 1878, Cumnoria Seeley 1999, Symphyrophus Cope 1878; Camptosauridae] N04

Camptosaurus dispar (Marsh 1879) [=Camptonotus dispar; incl. Brachyrophus altarkansanus Cope 1878, Camptosaurus browni Gilmore 1909, Camptosaurus medius Marsh 1894, Symphyrophus musculosus Cope 1878, Camptosaurus nanus Marsh 1894] N04

*Iguanodon anglicus Holl 1829 WU03, N04 [incl. Cetiosaurus brevis Owen 1842 N04, Streptospondylus major Owen 1842 N04]

Iguanodon atherfieldensis Hooley 1924 [incl. Sphenospondylus gracilis Lydekker 1888, I. mantelli Mayer 1832, Heterosaurus neocomiensis Cornuel 1850, Vectisaurus valdensis Hulke 1879] N04

Valdosaurus canaliculatus (Galton 1975) [=Dryosaurus canaliculatus; incl. Camptosaurus valdensis Lydekker 1889] N04

*Type species of generic name indicated

REFERENCES

[N85] Norman, D. 1985 (reprinted 2000). The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.

[N04] Norman, D. B. 2004. Basal Iguanodontia. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2nd ed. pp. 413–437. University of California Press: Berkeley.

[WU03] Wilson, J. A., & P. Upchurch. 2003. A revision of Titanosaurus Lydekker (Dinosauria—Sauropoda), the first dinosaur genus with a ‘Gondwanan’ distribution. Journal of Systematic Palaeontology 1 (3): 125–160.

Hadrosaurinae

Skeletal mount of Edmontosaurus annectens, copyright Andytang20.


Belongs within: Hadrosauridae.

The Hadrosaurinae are a lineage of hadrosaurid dinosaurs that are mostly crestless, or possess a solid crest formed from an extension of the nasal bones. The clade as given below is supported by the presence of a caudal margin on the circumnarial fossa (Horner et al. 2004).

<==Hadrosaurinae
    |--Lophorhothon Langston 1960 HWF04
    |    `--L. atopus Langston 1960 HWF04
    `--+--+--Naashoibitosaurus Hunt & Lucas 1993 HWF04
       |  |    `--N. ostromi Hunt & Lucas 1993 HWF04
       |  |--Kritosaurus Brown 1910 HWF04, D07
       |  |    |--K. australis Bonaparte, Franchi et al. 1984 HWF04
       |  |    `--K. navajovius Brown 1910 (n. d.) HWF04
       |  `--Saurolophus Brown 1912 HWF04
       |       |--S. angustirostris Rozhdestvensky 1952 HWF04
       |       |--S. kryschtofovici Riabinin 1930 (n. d.) HWF04
       |       `--S. osborni Brown 1912 HWF04
       `--+--Prosaurolophus Brown 1916 HWF04
          |    |--P. blackfeetensis Horner 1992 HWF04
          |    `--P. maximus Brown 1916 HWF04
          `--+--Gryposaurus Lambe 1914 HWF04
             |    |--G. incurvimanus (Parks 1920) [=Kritosaurus incurvimanus] HWF04
             |    |--G. latidens Horner 1992 HWF04
             |    `--G. notabilis Lambe 1914 HWF04 [=Kritosaurus notabilis D07]
             `--+--Edmontosaurus Lambe 1920 (see below for synonymy) HWF04
                |    |--E. annectens (Marsh 1892) HWF04 (see below for synonymy)
                |    |--E. regalis Lambe 1917 [incl. Trachodon atavus Cope 1871, Agathaumas milo Cope 1874] HWF04
                |    `--E. saskatchewanensis (Sternberg 1926) [=Thespesius saskatchewanensis] HWF04
                `--+--Brachylophosaurus Sternberg 1953 HWF04
                   |    `--B. canadensis Sternberg 1953 [incl. B. goodwini Horner 1988] HWF04
                   `--Maiasaura Horner & Makela 1979 HWF04
                        `--M. peeblesorum Horner & Makela 1979 HWF04

Hadrosaurinae incertae sedis:
  Anasazisaurus Hunt & Lucas 1993 HWF04
    `--A. horneri Hunt & Lucas 1993 HWF04
  Aralosaurus Rozhdestvensky 1968 HWF04
    `--A. tuberiferus Rozhdestvensky 1968 HWF04
  Shantungosaurus Hu 1973 HWF04
    `--S. giganteus Hu 1973 HWF04
  Kerberosaurus Bolotsky & Godefroit 2004 D07
    `--K. manakini D07
  Hadrosaurus Leidy 1858 HWF04
    |--H. breviceps Marsh 1889 (n. d.) HWF04
    |--H. cavatus Cope 1871 (n. d.) HWF04
    |--H. foulkii Leidy 1858 [incl. Ornithotarsus immanis Cope 1869] HWF04
    |--H. minor Marsh 1870 (n. d.) HWF04
    `--H. paucidens Marsh 1889 (n. d.) HWF04

Edmontosaurus Lambe 1920 [incl. Anatosaurus Lull & Wright 1942, Anatotitan Brett-Surman in Chapman & Brett-Surman 1990] HWF04

Edmontosaurus annectens (Marsh 1892) HWF04 [=Claosaurus annectens HWF04; incl. Anatosaurus copei Lull & Wright 1942 HWF04, Anatotitan copei D07, Thespesius edmontonensis Gilmore 1924 HWF04, Trachodon longiceps Marsh 1890 HWF04, Anatotitan longiceps D07]

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[HWF04] Horner, J. R., D. B. Weishampel & C. A. Forster. 2004. Hadrosauridae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2nd ed. pp. 438–463. University of California Press: Berkeley.

Hadrosauridae

Skeletal mount of Hypacrosaurus altispinus, copyright Roland Tanglao.


Belongs within: Hadrosauroidea.
Contains: Hadrosaurinae.

The Hadrosauridae are a group of large herbivorous dinosaurs known from the Late Cretaceous of Eurasia and North and South America, recognised by Horner et al. (2004) as the least inclusive clade containing Telmatosaurus and Parasaurolophus.

Characters (from Horner et al. 2004): Three or more replacement teeth present per tooth family; dentary tooth row extending distally to terminate caudal to apex of coronoid process, caudalmost termination of dentary well behind coronoid process; surangular foramen absent; ectopterygoid and jugal not in contact; supraorbital absent or fused to orbit rim; neural arches with cervical zygapophyseal peduncles elevated to extend well above level of neural canal; postzygapophyses long, dorsally arched; coracoid bearing long cranioventral process extending well below glenoid; proximal scapula dorsoventrally narrow, with acromion process projecting horizontally, notched cranioventral corner, reduced area for coracoid articulation; femur bearing deep intercondylar extensor groove, with edges meeting or nearly meeting cranially to enclose extensor tunnel.

<==Hadrosauridae
    |--Telmatosaurus Nopcsa 1903 [incl. Hecatasaurus Brown 1910, Limnosaurus Nopcsa 1900] HWF04
    |    |--T. cantabrigiensis D07
    |    `--T. transsylvanicus Nopcsa 1900 HWF04 [=Limnosaurus transsylvanicus HWF04, Orthomerus transsylvanicus PST99]
    `--Euhadrosauria HWF04
         |--Hadrosaurinae HWF04
         `--Lambeosaurinae HWF04
              |  i. s.: Pararhabdodon isonesis WJ03
              |         Barsboldia Maryańska & Osmólska 1981 HWF04
              |           `--B. sicinskii Maryańska & Osmólska 1981 HWF04
              |         Nipponosaurus Nagao 1936 HWF04
              |           `--N. sachalinensis Nagao 1936 HWF04
              |         Bactrosaurus Gilmore 1933 D07
              |           |--B. johnsoni Gilmore 1933 HWF04
              |           |--B. kyzylkumensis D07
              |           `--B. prynadai Riabinin 1939 (n. d.) HWF04
              |--Tsintaosaurus Young 1958 HWF04
              |    `--T. spinorhinus Young 1958 [incl. Tanius laiyangensis Zhen 1976] HWF04
              `--+--Jaxartosaurus Riabinin 1939 GBA03, HWF04
                 |    |--J. aralensis Riabinin 1939 [incl. Procheneosaurus convincens Rozhdestvensky 1968] HWF04
                 |    `--J. fuyunensis Wu 1984 (n. d.) HWF04
                 `--+--Amurosaurus Bolotsky & Kurzanov 1991 GBA03, HWF04
                    |    `--A. riabinini Bolotsky & Kurzanov 1991 HWF04
                    `--+--+--Charonosaurus Godefroit, Zan & Jin 2000 GBA03, HWF04
                       |  |    `--C. jiayinensis Godefroit, Zan & Jin 2000 HWF04
                       |  `--Parasaurolophus Parks 1922 GBA03, HWF04
                       |       |--P. cyrtocristatus Ostrom 1961 HWF04
                       |       |--P. tubicen Wiman 1931 HWF04
                       |       `--P. walkeri Parks 1922 HWF04
                       `--+--Lambeosaurus Parks 1923 GBA03, HWF04 [incl. Didanodon Osborn 1902 HWF04]
                          |    |--L. lambei Parks 1923 HWF04 (see below for synonymy)
                          |    |--L. laticaudus Morris 1981 HWF04
                          |    `--L. magnicristatus Sternberg 1935 HWF04
                          `--+--Corythosaurus Brown 1914 GBA03, HWF04 [incl. Procheneosaurus Matthew 1920 HWF04]
                             |    `--C. casuarius Brown 1914 HWF04 (see below for synonymy)
                             |--Olorotitan Godefroit, Boltsky & Alifanov 2003 GBA03
                             |    `--*O. arharensis Godefroit, Bolotsky & Alifanov 2003 GBA03
                             `--Hypacrosaurus Brown 1913 GBA03, HWF04
                                  |--H. altispinus Brown 1913 [incl. Cheneosaurus tolmanensis Lambe 1917] HWF04
                                  |--H. casuarius P88
                                  `--H. stebingeri Horner & Currie 1994 HWF04

Hadrosauridae incertae sedis:
  Gilmoreosaurus Brett-Surman 1979 HWF04
    |--G. arkhangelskyi D07
    |--G. atavus D07
    `--G. mongoliensis (Gilmore 1933) [=Mandschurosaurus mongoliensis] HWF04
  Tanius Wiman 1929 HWF04
    `--T. sinensis Wiman 1929 [incl. T. chingkankouensis Young 1958] HWF04
  Claosaurus Marsh 1890 HWF04
    |--C. affinis Wieland 1903 (n. d.) HWF04
    `--C. agilis (Marsh 1872) [=Hadrosaurus agilis] HWF04
  Secernosaurus Brett-Surman 1979 HWF04
    `--S. koerneri Brett-Surman 1979 HWF04
  Arstanosaurus akkurganensis Suslov & Shilin 1982 (n. d.) HWF04
  Cionodon HWF04
    |--C. arctatus Cope 1874 (n. d.) HWF04
    |--C. kysylkumensis Riabinin 1931 (n. d.) HWF04
    `--C. stenopsis Cope 1875 (n. d.) HWF04
  Claorhynchus trihedrus Cope 1892 (n. d.) HWF04
  Diclonius HWF04
    |--D. calamarius Cope 1876 (n. d.) HWF04
    |--D. pentagonus Cope 1876 (n. d.) HWF04
    `--D. perangulatus Cope 1876 (n. d.) HWF04
  Hypsibema crassicauda Cope 1869 (n. d.) HWF04
  ‘Iguanodon’ hilli Newton 1892 (n. d.) HWF04
  Mandschurosaurus Riabinin 1930 (n. d.) HWF04
    |--M. amurensis (Riabinin 1925) [=Trachodon amurensis] HWF04
    `--M. laosensis Hoffet 1943 (n. d.) HWF04
  Microhadrosaurus nanshiungensis Dong 1979 (n. d.) HWF04
  Orthomerus HWF04
    |--O. dolloi Seeley 1883 (n. d.) HWF04
    `--O. weberi Riabinin 1945 (n. d.) HWF04
  Parrosaurus Gilmore 1945 (n. d.) HWF04
    `--*P. missouriensis (Gilmore & Stewart 1945) [=Neosaurus missouriensis] HWF04
  Pteropelyx grallipes Cope 1889 (n. d.) HWF04
  Thespesius occidentalis Leidy 1856 (n. d.) HWF04
  Trachodon HWF04
    |--T. altidens Lambe 1902 (n. d.) HWF04
    |--T. cantabrigiensis Lydekker 1888 (n. d.) HWF04
    |--T. marginatus Lambe 1902 (n. d.) HWF04
    |--T. mirabilis Leidy 1856 (n. d.) HWF04
    `--T. selwyni Lambe 1902 (n. d.) HWF04

Corythosaurus casuarius Brown 1914 HWF04 [incl. C. bicristatus Parks 1935 HWF04, C. brevicristatus Parks 1935 HWF04, Tetragonosaurus cranibrevis Sternberg 1935 HWF04, Procheneosaurus cranibrevis D75, T. erectofrons Parks 1931 HWF04, P. erectofrons D75, Corythosaurus excavatus Gilmore 1923 HWF04, C. intermedius Parks 1923 HWF04]

Lambeosaurus lambei Parks 1923 HWF04 [incl. L. clavinitialis Sternberg 1935 HWF04, Corythosaurus frontalis Parks 1935 HWF04, Tetragonosaurus praeceps Parks 1931 HWF04, Procheneosaurus praeceps D75]

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[D75] Dodson, P. 1975. Taxonomic implications of relative growth in lambeosaurine hadrosaurs. Systematic Zoology 24 (1): 37–54.

[GBA03] Godefroit, P., Y. Bolotsjy & V. Alifanov. 2003. A remarkable hollow-crested hadrosaur from Russia: an Asian origin for lambeosaurines. Comptes Rendus Palevol 2: 143–151.

[HWF04] Horner, J. R., D. B. Weishampel & C. A. Forster. 2004. Hadrosauridae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2nd ed. pp. 438–463. University of California Press: Berkeley.

[P88] Paul, G. S. 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide. Simon & Schuster: New York.

[PST99] Pereda Suberbiola, X., & P. Taquet. 1999. Restes de Rhabdodon (dinosaure ornithopode) de Transylvanie donnés par Nopcsa au Muséum national d'Histoire naturelle de Paris. Geodiversitas 21 (2): 157–166.

[WJ03] Weishampel, D. B., C.-M. Jianu, Z. Csiki & D. B. Norman. 2003. Osteology and phylogeny of Zalmoxes (n. g.), an unusual euornithopod dinosaur from the latest Cretaceous of Romania. Journal of Systematic Palaeontology 1: 65–123.

Ceratopsia

Skeletal mounts of Psittacosaurus mongoliensis, copyright Ninjatacoshell.


Belongs within: Genasauria.
Contains: Ceratopsidae.

The Ceratopsia are a group of herbivorous dinosaurs known from the Late Jurassic to Cretaceous of Asia and North America (with some less definite records from other parts of the world). Members of this group are characterised by a novel bone, the rostral bone, at the front of the upper jaw (which forms a narrow beak), and a tendency to the develop of a frill on the rear margin of the skull (You & Dodson 2004).

See also: Big horned lizards; Ceratopsids: a Cretaceous flash in the pan.

Synapomorphies (from You & Dodson 2004): External naris high, separated from ventral border of premaxilla by flat area; rostral bone present; premaxilla enlarged; jugal with well-developed lateral flaring; wide dorsoventral length of infraorbital ramus of jugal; palatal extensions of maxillae contacting rostral to choana.

Ceratopsia
    |  i. s.: Micropachycephalosaurus Dong 1978 ZY09, MCW04
    |           `--M. hongtuyanensis Dong 1978 MCW04
    |--Psittacosaurus Osborn 1923 [incl. Protiguanodon Osborn 1923] YD04
    |    |--P. guyangensis Cheng 1983 YD04
    |    |--P. mazongshanensis Xu 1997 YD04
    |    |--P. meileyingensis Sereno, Chao et al. 1988 YD04
    |    |--P. mongoliensis Osborn 1923 (see below for synonymy) YD04
    |    |--P. neimongoliensis Russell & Zhao 1996 YD04
    |    |--P. ordosensis Russell & Zhao 1996 YD04
    |    |--P. osborni Young 1931 [incl. P. tingi Young 1931] YD04
    |    |--P. sattayaraki Buffetaut & Suteethorn 1992 (n. d.) YD04
    |    |--P. sinensis Young 1958 YD04
    |    |--P. xinjiangensis Sereno & Chao 1988 YD04
    |    `--P. youngi Chao 1962 YD04
    `--Neoceratopsia [Protoceratopidae, Protoceratopoidea] YD04
         |  i. s.: Microceratops YD04
         |           |--M. gobiensis Bohlin 1953 (n. d.) YD04
         |           `--M. sulcidens Bohlin 1953 (n. d.) YD04
         |         Kulceratops kulensis Nessov 1995 (n. d.) YD04
         |         Yamaceratops dorngobiensis BN08
         |         Bainoceratops efremovi T08
         |--Yinlong downsi BSN07
         `--+--Chaoyangsaurus Zhao, Cheng & Xu 1999 YD04
            |    `--C. youngi Zhao, Cheng & Xu 1999 YD04
            `--+--Liaoceratops Xu, Makovicky et al. 2002 XM02
               |    `--*L. yanzigouensis Xu, Makovicky et al. 2002 XM02
               `--+--Archaeoceratops Dong & Azuma 1997 [Archaeoceratopidae] YD04
                  |    `--A. oshimai Dong & Azuma 1997 YD04
                  `--Coronosauria YD04
                       |--Ceratopsoidea YD04
                       |    |--+--Ceratopsidae YD04
                       |    |  `--Zuniceratops Wolfe & Kirkland 1998 YD04
                       |    |       `--Z. christopheri Wolfe & Kirkland 1998 YD04
                       |    `--+--Asiaceratops salsopaludalis Nessov, Kaznyshkina & Cherepanov 1989 (n. d.) XM02, YD04
                       |       `--Leptoceratopsidae YD04
                       |            |--Montanoceratops Sternberg 1951 YD04
                       |            |    `--M. cerorhynchus (Brown & Schlaikjer 1942) [=Leptoceratops cerorhynchus] YD04
                       |            `--+--Leptoceratops Brown 1914 YD04
                       |               |    `--L. gracilis Brown 1914 YD04
                       |               `--Udanoceratops Kurzanov 1992 XM02, YD04
                       |                    `--U. tschizhovi Kurzanov 1992 YD04
                       `--Protoceratopsidae YD04
                            |--Graciliceratops Sereno 2000 YD04
                            |    `--G. mongoliensis Sereno 2000 YD04
                            |--Protoceratops Granger & Gregory 1923 YD04
                            |    |--P. andrewsi Granger & Gregory 1923 YD04
                            |    `--P. hellenikorhinus Lambert, Godefroit et al. 2001 YD04
                            `--Bagaceratopidae A03
                                 |--Bagaceratops Maryańska & Osmólska 1975 [incl. Breviceratops Kurzanov 1990] YD04
                                 |    `--B. rozhdestvenskyi Maryańska & Osmólska 1975 (see below for synonymy) YD04
                                 |--Lamaceratops Alifanov 2003 A03
                                 |    `--*L. tereschenkoi Alifanov 2003 A03
                                 `--Platyceratops Alifanov 2003 A03
                                      `--*P. tatarinovi Alifanov 2003 A03

Bagaceratops rozhdestvenskyi Maryańska & Osmólska 1975 [incl. Protoceratops kozlowskii Maryańska & Osmólska 1975, Breviceratops kozlowskii] YD04

Psittacosaurus mongoliensis Osborn 1923 [incl. Protiguanodon mongoliensis Osborn 1923, Ps. protiguanodonensis Young 1958] YD04

*Type species of generic name indicated

REFERENCES

[A03] Alifanov, V. R. 2003. Two new dinosaurs of the infraorder Neoceratopsia (Ornithischia) from the Upper Cretaceous of the Nemegt Depression, Mongolian People’s Republic. Paleontologicheskii Zhurnal 2003 (5): 77–88 (transl. Paleontological Journal 37 (5): 524–534).

[BN08] Balanoff, A. M., M. A. Norell, G. Grellet-Tinner & M. R. Lewin. 2008. Digital preparation of a probable neoceratopsian preserved within an egg, with comments on microstructural anatomy of ornithischian eggshells. Naturwissenschaften 95: 493–500.

[BSN07] Butler, R. J., R. M. H. Smith & D. B. Norman. 2007. A primitive ornithischian dinosaur from the Late Triassic of South Africa, and the early evolution and diversification of Ornithischia. Proceedings of the Royal Society of London Series B—Biological Sciences 274 (1621): 2041–2046.

[MCW04] Maryańska, T., R. E. Chapman & D. B. Weishampel. 2004. Pachycephalosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2nd ed. pp. 464–477. University of California Press: Berkeley.

[T08] Tereschenko, V. S. 2008. Adaptive features of protoceratopoids (Ornithischia: Neoceratopsia). Paleontologicheskii Zhurnal 2008 (3): 50–64.

[XM02] Xu, X., P. J. Makovicky, X.-L. Wang, M. A. Norell & H.-L. You. 2002. A ceratopsian dinosaur from China and the early evolution of Ceratopsia. Nature 416: 314–317.

[YD04] You H. & P. Dodson. 2004. Basal Ceratopsia. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2nd ed. pp. 478–493. University of California Press: Berkeley.

[ZY09] Zheng, X.-T., H.-L. You, X. Xu & Z.-M. Dong. 2009. An Early Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures. Nature 458: 333–336.

Pachycephalosauria

Cranium of Sphaerotholus buchholtzae in dorsal view (arrows indicate lesions), from Peterson et al. (2013).


Belongs within: Genasauria.

The Pachycephalosauria are a group of bipedal herbivorous dinosaurs with a thickened skull roof known from the Cretaceous of Eurasia and North America (Maryanska et al. 2004).

Characters (from Maryanska et al. 2004): Skull roof thickened; frontal excluded from orbital margin; postorbital bar with dorsal surface flat, broad; parietal without caudolateral wings; squamosal extended onto occiput, bearing tubercles on caudolateral margin; basal tubera thin, platelike; prootic-basisphenoid plate contacting pterygoid-prootic plate; jugal and quadrate eliminating free ventral margin of quadratojugal; dorsal vertebrae with double ridge-and-groove articulations; sacral ribs elongate; caudal basket of fusiform ossified tendons present; illium with sigmoidal border; medial process present on iliac blade; pubis nearly excluded from acetabulum.

Pachycephalosauria
    |  i. s.: Yaverlandia Galton 1971 MCW04
    |           `--Y. bitholus Galton 1971 MCW04
    |         Heishansaurus Bohlin 1953 MCW04, D07
    |           `--H. pachycephalus Bohlin 1953 (n. d.) MCW04
    |--Stenopelix Mayer 1857 MCW04
    |    `--S. valdensis Meyer 1857 MCW04
    `--+--Wannanosaurus Hou 1977 MCW04
       |    `--W. yansiensis Hou 1977 MCW04
       `--Goyocephala MCW04
            |--Goyocephale Perle, Maryańska & Osmólska 1982 MCW04
            |    `--G. lattimorei Perle et al. 1982 MCW04
            `--Homalocephaloidea MCW04
                |--Homalocephale Maryańska & Osmólska 1974 MCW04
                |    `--H. calathocercos Maryańska & Osmólska 1974 MCW04
                `--+--Ornatotholus Galton & Sues 1983 MCW04
                   |    `--*O. browni (Wall & Galton 1979) S03 [=Stegoceras browni MCW04]
                   `--Pachycephalosauridae [Pachycephalosaurinae, Pachycephalosaurini, Tholocephalidae] MCW04
                        |  i. s.: ‘Troodon’ bexelli Bohlin 1953 MCW04 [=Stegoceras bexelli S03]
                        |         Gravitholus Wall & Galton 1979 MCW04
                        |           `--*G. albertae Wall & Galton 1979 (n. d.) S03
                        |         Dracorex hogwartsia D07
                        |--Stygimoloch Galton & Sues 1983 [incl. Stenotholus Giffin, Gabriel & Johnson 1987] MCW04
                        |    `--*S. spinifer Galton & Sues 1983 S03 (see below for synonymy)
                        `--+--Stegoceras Lambe 1902 [incl. Colepiocephale Sullivan 2003, Hanssuesia Sullivan 2003] MCW04
                           |    |--*S. validum Lambe 1918 S03, MCW04 (see below for synonymy)
                           |    `--S. edmontonense (Brown & Schlaikjer 1943) MCW04 (see below for synonymy)
                           `--+--Pachycephalosaurus Brown & Schlaikjer 1943 (see below for synonymy) MCW04
                              |    `--*P. wyomingensis (Gilmore 1931) S03 (see below for synonymy)
                              `--+--Tylocephale Maryańska & Osmólska 1974 MCW04
                                 |    |--T. bexelli D07
                                 |    `--T. gilmorei Maryańska & Osmólska 1974 S03
                                 |--Prenocephale Maryańska & Osmólska 1974 S03
                                 |    `--*P. prenes Maryańska & Osmólska 1974 S03
                                 `--Sphaerotholus Williamson & Carr 2002 MCW04
                                      |--S. buchholtzae Williamson & Carr 2002 MCW04
                                      |--S. edmontonense D07
                                      `--S. goodwini Williamson & Carr 2002 MCW04 [=Prenocephale goodwini S03]

Pachycephalosaurus Brown & Schlaikjer 1943 (nom. cons.) [incl. Tylosteus Leidy 1872] MCW04

*Pachycephalosaurus wyomingensis (Gilmore 1931) S03 [=Troodon wyomingensis MCW04; incl. P. grangeri Brown & Schlaikjer 1943 MCW04, Tylosteus ornatus Leidy 1872 MCW04, P. reinheimeri Brown & Schlaikjer 1943 MCW04]

Stegoceras edmontonense (Brown & Schlaikjer 1943) MCW04 [=Troodon edmontonensis MCW04, Prenocephale edmontonensis S03, Stegoceras edmontonenis (l. c.) S03, S. edmontonensis S03, S. edmontonesis (l. c.) S03]

*Stegoceras validum Lambe 1918 S03, MCW04 [=S. validus (l. c.) S03, Troodon validus S03; incl. S. breve Lambe 1918 MCW04, Prenocephale brevis S03, S. brevis (l. c.) S03, S. lambei Sternberg 1945 MCW04, *Colepiocephale lambei S03, Troodon sternbergi Brown & Schlaikjer 1943 MCW04, Gravitholus sternbergi S03, *Hansuessia sternbergi S03, Stegoceras sternbergi S03]

*Sygimoloch spinifer Galton & Sues 1983 S03 [incl. Stenotholus kohleri Giffin, Gabriel & Johnson 1987 MCW04]

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[MCW04] Maryańska, T., R. E. Chapman & D. B. Weishampel. 2004. Pachycephalosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2nd ed. pp. 464–477. University of California Press: Berkeley.

[S03] Sullivan, R. M. 2003. Revision of the dinosaur Stegoceras Lambe (Ornithischia, Pachycephalosauridae). Journal of Vertebrate Paleontology 23 (1): 181–207.

Gonyleptidae

Neopucroliella pertyi, copyright Luis Acosta.


Belongs within: Grassatores.
Contains: Heteropachylinae, Ampycinae, Mitobatinae, Goniosomatinae, Hernandariinae, Caelopyginae, Progonyleptoidellinae, Sodreaninae, Gonyleptes, Geraecormobius, Metagonyleptes, Collonychium, Mischonyx, Sphaerobunus, Discocyrtus, Eusarcus, Sadocus, Pachyloides.

The Gonyleptidae are a diverse group of harvestmen found in Central and South America, many of which are ornately armoured and/or brightly coloured. The family is divided between several subfamilies, though some of these (particularly the larger subfamilies Gonyleptinae and Pachylinae) remain poorly defined.

Characters (from Kury & Pinto-da-Rocha 2007): Carapace usually much narrower than opisthosomal scutum, body rarely subrectangular. Frontal hump usually present; with tubercles short or long. Dorsal scutum usually longer than wide. With one or two ozopores on each side. Body depressed or convex. Common ocularium always present. Three or four areas on dorsal scutum; sulci well marked; area I divided by median groowve; IV rarely divided. Chelicerae usually similar in size and small in both sexes, rarely hypertelic in male. Coxa IV usually dorsally visible in entire extension; with dorsoapical apophysis short to robust, single or bifid, much more developed on males. Trochanter, femur and tibia IV often armed with tubercles/apophyses in males. Scopulae usually present on legs III and IV. Claws usually smooth, sometimes pectinate. Penis with ventral plate well defined, rectangular or pyriform with basal lobes more or less developed; lateral margin of ventral plate with two groups of setae; glans columnar, more or less elongate, rarely with dorsal process, commonly with ventral process.

Gonyleptidae [Ejonyleptidae, Golyleptidae, Gonileptidae, Gonyleptoidae]
    |  i. s.: Heteropachylinae P-R02
    |         Ampycinae KP-R07
    |         Nanophareus Roewer 1929 K03b
    |           `--*N. palpalis Roewer 1929 K03b
    |--Bourguyiinae [Bourguyinae] P-R02
    |    |--Bogdana Mello-Leitão 1940 K03b
    |    |    `--*B. ingenua Mello-Leitão 1940 [=Asarcus ingenuus] K03b
    |    |--Caldasius Roewer 1930 K03b
    |    |    `--*C. trochanteralis Roewer 1930 K03b
    |    |--Cnemoleptes Mello-Leitão 1941 K03b [=Cnemoleptus (l. c.) KA-Z11]
    |    |    `--*C. passarellii Mello-Leitão 1941 K03b
    |    |--Styloleptes Piza 1943 K03b
    |    |    `--*S. conspersus Piza 1943 [=Stylopteptes conspersus] K03b
    |    |--Stylopisthos Roewer 1930 K03b
    |    |    `--*S. laevibunus Roewer 1930 K03b
    |    |--Opisthoplites Sørensen 1884 [=Opisthoplithes (l. c.), Opysthoplytes (l. c.)] K03b
    |    |    |--*O. ypsilon Sørensen 1884 K03b
    |    |    `--O. corallipes (Simon 1879) [=Asarcus corallipes] K03b
    |    |--Bourguyia Mello-Leitão 1923 (see below for synonymy) K03b
    |    |    |--*B. albiornata Mello-Leitão 1923 (see below for synonymy) K03b
    |    |    |--B. amarali (Mello-Leitão 1934) [=*Afranius amarali] K03b
    |    |    |--B. hamata (Roewer 1943) [=*Drastus hamatus] K03b
    |    |    `--B. maculata (Roewer 1930) [=Caldasius maculatus] K03b
    |    `--Asarcus Koch 1839 K03b
    |         |--*A. longipes Kollar in Koch 1839 K03b
    |         |--A. lutescens Sørensen 1884 K03b
    |         |--A. nigriconspersus Soares & Soares 1945 K03b
    |         `--A. pallidus Mello-Leitão 1923 K03b
    |--Pachylospeleus Šilhavý 1974 K03b [Pachylospeleinae P-R02]
    |    `--*P. strinatii Šilhavý 1974 K03b
    |--Cobania Roewer 1913 P-R02, K03b (see below for synonymy)
    |    |--*C. picea (Bertkau 1880) (see below for synonymy) K03b
    |    `--C. validissima (Mello-Leitão 1933) [=*Flangeia validissima] K03b
    |--Gonyassamiinae P-R02
    |    |--Phalangochilus Mello-Leitão 1938 K03b
    |    |    `--*P. luteipes Mello-Leitão 1938 K03b
    |    `--Trichominua Mello-Leitão 1938 [incl. Gonyassamia Soares & Soares 1988] K03b
    |         |--*T. annulipes Mello-Leitão 1938 K03b
    |         `--T. roeweri (Soares & Soares 1954) [=Progyndes roeweri, *Gonyassamia roeweri] K03b
    |--+--Mitobatinae DG09
    |  `--+--+--Goniosomatinae DG09
    |     |  `--+--+--Hernandariinae DG09
    |     |     |  `--Caelopyginae DG09
    |     |     `--+--Progonyleptoidellinae DG09
    |     |        `--Sodreaninae DG09
    |     `--Gonyleptinae DG09
    |          |    i. s.: ‘Mitobates’ albipunctatus Roewer 1943 K03b
    |          |--Gonyleptes P-R02
    |          |--Geraecormobius K03b
    |          |--Metagonyleptes K03b
    |          |--Collonychium K03b
    |          |--Mischonyx K03b
    |          |--Sphaerobunus DG09
    |          |--Haversia Roewer 1913 K03b
    |          |    `--*H. defensa (Butler 1876) [=Gonyleptes defensus] K03b
    |          |--Progonyleptes Roewer 1913 K03b
    |          |    `--*P. borellii (Sørensen 1895) [=Gonyleptes borellii] K03b
    |          |--Eugonyleptes Roewer 1913 K03b
    |          |    `--*E. scaber (Kirby 1819) K03b, KA-Z11 [=Gonyleptes scaber K03b, Discocyrtus scaber K03b]
    |          |--Triaenomeros Roewer 1913 K03b
    |          |    `--*T. olivaceus Roewer 1913 K03b
    |          |--Adhynastes Roewer 1930 K03b
    |          |    `--*A. tenuis Roewer 1930 K03b
    |          |--Cearinus Roewer 1929 [incl. Nemoribalta Mello-Leitão 1941] K03b
    |          |    `--*C. corniger Roewer 1929 [incl. *Nemoribalta plana Mello-Leitão 1941] K03b
    |          |--Centroleptes Roewer 1943 K03b
    |          |    `--*C. flavus Roewer 1943 K03b
    |          |--Deltaspidium Roewer 1927 K03b
    |          |    `--D. asper (Perty 1833) (see below for synonymy) K03b
    |          |--Gonazula Roewer 1930 K03b
    |          |    `--*G. gibbosa Roewer 1930 K03b
    |          |--Guatubesia Soares 1978 K03b
    |          |    `--*G. clarae Soares 1978 K03b
    |          |--Gyndesops Roewer 1943 K03b
    |          |    `--*G. denisi Roewer 1943 [=Temucus denisi] K03b
    |          |--Hoggellula Roewer 1930 K03b
    |          |    `--*H. vallentini (Hogg 1913) [=Sadocus vallentini] K03b
    |          |--Holoversia Mello-Leitão 1940 K03b
    |          |    `--*H. nigra Mello-Leitão 1940 K03b
    |          |--Oxapampeus Roewer 1963 K03b
    |          |    `--*O. weyrauchi Roewer 1963 K03b
    |          |--Proctobunoides Mello-Leitão 1944 K03b
    |          |    `--*P. tuberosus Mello-Leitão 1944 K03b
    |          |--Stefanesia Soares & Soares 1988 K03b
    |          |    `--*S. avramae Soares & Soares 1988 K03b
    |          |--Urodiabunus Mello-Leitão 1935 K03b
    |          |    `--*U. arlei Mello-Leitão 1935 K03b
    |          |--Ypsilonurus Mello-Leitão 1933 K03b
    |          |    `--*Y. mutilatus Mello-Leitão 1933 K03b
    |          |--Progonyleptoides Roewer 1917 [incl. Paranabius Roewer 1943] K03b
    |          |    |--*P. spinifrons Roewer 1917 K03b
    |          |    `--P. castaneus (Roewer 1943) [=*Paranabius castaneus] K03b
    |          |--Currala Roewer 1927 K03b
    |          |    |--*C. spinifrons Roewer 1927 K03b
    |          |    `--C. bahiensis Soares 1972 K03b
    |          |--Schenkelibunus Strand 1932 K03b, KA-Z11 (see below for synonymy)
    |          |    |--*S. perditus (Mello-Leitão 1927) K03b, KA-Z11 (see below for synonymy)
    |          |    `--S. impar (Mello-Leitão 1932) KA-Z11 (see below for synonymy)
    |          |--Gonyleptellus Roewer 1930 [incl. Stephanocranion Mello-Leitão 1931] K03b
    |          |    |--G. cancellatus (Roewer 1917) (see below for synonymy) K03b
    |          |    `--G. bufoninus (Mello-Leitão 1949) [=Stephanocranion bufoninus] K03b
    |          |--Inhuma Piza 1938 [incl. Hugoesia Soares 1968] K03b
    |          |    |--*I. pessoai Piza 1938 K03b
    |          |    `--I. lopesi (Soares 1968) [=*Hugoesia lopesi] K03b
    |          |--Uracantholeptes Mello-Leitão 1926 [incl. Gonypernoides Mello-Leitão 1932] K03b
    |          |    |--*U. anomalus (Mello-Leitão 1922) (see below for synonymy) K03b
    |          |    `--U. damicoi (Soares & Soares 1945) [=Gonypernoides damicoi] K03b
    |          |--Megapachylus Roewer 1913 (see below for synonymy) K03b
    |          |    |--*M. grandis Roewer 1913 (see below for synonymy) K03b
    |          |    |--M. mutilatus (Mello-Leitão 1932) K03b [=Gonyperna mutilata K03b, Paragonyleptes mutilatus SS49]
    |          |    `--M. simoni (Roewer 1930) (see below for synonymy) K03b
    |          |--Tupacarana Mello-Leitão 1939 K03b
    |          |    |--*T. serrina Mello-Leitão 1939 K03b
    |          |    |--T. gofferjei Soares & Soares 1947 K03b
    |          |    `--T. marmorata Mello-Leitão 1939 K03b
    |          |--Neosadocus Mello-Leitão 1926 K-PR07, K03b (see below for synonymy)
    |          |    |--*N. bufo (Mello-Leitão 1923) (see below for synonymy) K03b
    |          |    |--N. maximus (Giltay 1928) [=Neomitobates maximus, *Mitoperna maxima] K03b
    |          |    |--N. misandrus (Mello-Leitão 1934) [=Metagonyleptes misandrus] K03b
    |          |    |--N. robustus (Mello-Leitão 1936) [=*Ilhania robusta] K03b
    |          |    `--N. variabilis (Mello-Leitão 1935) DG09 [=*Bunoweyhia variabilis K03b]
    |          |--Liogonyleptoides Mello-Leitão 1925 (see below for synonymy) K03b
    |          |    |--*L. inermis (Mello-Leitão 1922) (see below for synonymy) K03b
    |          |    |--L. capichaba Soares & Soares 1946 K03b
    |          |    |--L. heliae Kury 2003 [=L. minensis Soares 1966 non Chaquesia minensis Piza 1946] K03b
    |          |    |--L. minensis (Piza 1946) [=Chaquesia minensis] K03b
    |          |    `--L. tetracanthus (Mello-Leitão 1932) [=*Pachyleptes tetracanthus] K03b
    |          `--Parampheres Roewer 1913 KP-R07, K03b (see below for synonymy)
    |               |--*P. pectinatus Roewer 1913 K03b
    |               |--P. bimaculatus (Mello-Leitão 1932) (see below for synonymy) K03b
    |               |--P. boliviensis (Roewer 1931) [=*Callampheres boliviensis] K03b
    |               |--P. lucidus (Mello-Leitão 1940) [=Penygorna lucida, Arleius lucidus, Ilhaia lucida] K03b
    |               |--P. ronae (Mello-Leitão 1927) (see below for synonymy) K03b
    |               `--P. tibialis Roewer 1917 [incl. Philocrates maculatus Sørensen in Roewer 1943 (n. n.)] K03b
    `--Pachylinae DG09
         |  i. s.: ‘Caldasius’ nigripes Mello-Leitão 1942 K03b
         |         ‘Liogyndulus’ bispinifrons Roewer 1943 K03b
         |         ‘Liogyndulus’ opacus Roewer 1952 K03b
         |         ‘Prosontes’ brasiliensis Soares 1966 K03b
         |--Discocyrtus DG09
         |--Eusarcus K03b
         |--Sadocus P-R02
         |--Pachyloides K08
         |--Parapucrolia Roewer 1917 [incl. Goytacazia Mello-Leitão 1942, Goitacazia (l. c.), Goytacazio (l. c.)] K03b
         |    `--*P. ocellata Roewer 1917 (see below for synonymy) K03b
         |--Eubalta Roewer 1923 [=Balta Sørensen 1902 non Tapper 1893, Baltanata Strand 1942] K03b
         |    `--*E. meridionalis (Sørensen 1902) [=*Balta meridionalis, *Baltanata meridionalis] K03b
         |--Meteusarcus Roewer 1913 K03b
         |    `--*M. armatus Roewer 1913 K03b
         |--Paraphalangodus Roewer 1915 K03b
         |    `--*P. synacanthus Roewer 1915 K03b
         |--Acanthopachylus Roewer 1913 (see below for synonymy) K03b
         |    `--*A. aculeatus (Kirby 1818) K03b, KA-Z11 (see below for synonymy)
         |--Acanthopachylopsis Soares & Soares 1949 K03b
         |    `--*A. spectabilis Soares & Soares 1949 K03b
         |--Allogonyleptes Roewer 1917 K03b
         |    `--*A. insignitus Roewer 1917 K03b
         |--Anoplogynopsis Soares 1966 K03b
         |    `--*A. concolor Soares 1966 K03b
         |--Antetriceras Roewer 1949 K03a
         |    `--*A. signatum Roewer 1949 K03b, KA-Z11, K03b [=A. signatus K03b]
         |--Berlaia Mello-Leitão 1940 K03b
         |    `--*B. dissimilis Mello-Leitão 1940 (see below for synonymy) K03b
         |--Biconisoma Roewer 1936 K03b
         |    `--*B. mirabile Roewer 1936 K03b, KA-Z11, K03b [=B. mirabilis K03b]
         |--Bullaepus Roewer 1930 [=Bullaeepus (l. c.)] K03b
         |    `--*B. enoplus (Chamberlin 1916) [=Gonoleptes (l. c. for Gonyleptes) enoplus] K03b
         |--Bunoplus Roewer 1927 K03b
         |    `--*B. pachypalpis Roewer 1927 [=B. pachypalpi (l. c.)] K03b
         |--Calcarogyndes Mello-Leitão 1932 [=Calcogyndes (l. c.), Callcarogyndes (l. c.)] K03b
         |    `--*C. calcar (Roewer 1913) [=Metagyndes calcar] K03b
         |--Caldanatus Roewer 1943 K03b
         |    `--*C. marginalis Roewer 1943 K03b
         |--Camposicoloides Soares 1944 K03b
         |    `--*C. mendax Soares 1944 K03b
         |--Capichabesia Soares 1944 K03b
         |    `--*C. rarissima Soares 1944 K03b
         |--Ceratoleptes Soares & Soares 1979 K03b
         |    `--*C. proboscis Soares & Soares 1979 (see below for synonymy) K03b
         |--Chaquesia Soares 1944 K03b
         |    `--*C. guaranitica (Mello-Leitão 1933) [=Goyazella guaranitica] K03b
         |--Chauveaua Canals 1939 K03b
         |    `--*C. nahuelhapiensis Canals 1939 [=Phalangodus nahuelhapiensis] K03b
         |--Chilebalta Roewer 1961 K03b
         |    `--*C. angulipes Roewer 1961 K03b
         |--Chilegyndes Roewer 1961 K03b
         |    `--*C. phillipsoni Roewer 1961 K03b
         |--Corralia Roewer 1913 K03b
         |    `--*C. depressa (Loman 1899) [=Gonyleptes depressus; incl. Neogonyleptes ignotus Soares 1968] K03b
         |--Eopachyloides Soares 1970 K03b
         |    `--*E. trochanteralis Soares 1970 K03b
         |--Ergastria Mello-Leitão 1941 [=Castellanosia Mello-Leitão 1937 non Kraglievitch 1932] K03b
         |    `--*E. forficula (Mello-Leitão 1937) [=*Castellanosia forficula] K03b
         |--Eugyndopsiella Soares 1972 K03b
         |    `--*E. trochanteroceros Soares 1972 K03b
         |--Fonckia Roewer 1913 K03b
         |    `--*F. processigera (Sørensen 1902) [=Gonyleptes processiger, F. prossigera (l. c.)] K03b
         |--Giupponia Pérez G. & Kury 2002 K03b
         |    `--*G. chagasi Pérez G. & Kury 2002 K03b
         |--Goodnightiella Soares & Soares 1945 [incl. Prosampycoides Soares & Bauab-Vianna 1972] K03b
         |    `--*G. impar Soares & Soares 1945 [incl. *Prosampycoides spinipes Soares & Bauab-Vianna 1972] K03b
         |--Gyndesoides Mello-Leitão 1933 K08 [incl. Klobinia Mello-Leitão 1936 K03b]
         |    `--*G. dispar Mello-Leitão 1933 [incl. *Klobinia infuscata Mello-Leitão 1936] K03b
         |--Harpachylus Roewer 1943 K03b
         |    `--*H. tibialis Roewer 1943 K03b
         |--Huadquina Roewer 1930 K03b
         |    `--*H. huadquinae (Chamberlin 1916) (see below for synonymy) K03b
         |--Hyperpachylus Roewer 1957 K03b
         |    `--*H. mirabilis Roewer 1957 K03b
         |--Iandumoema Pinto-da-Rocha 1997 K03b, KA-Z11
         |    `--*I. uai Pinto-da-Rocha 1997 K03b, KA-Z11
         |--Ibarra Roewer 1925 K03b
         |    `--*I. festae Roewer 1925 K03b
         |--Iguassua Mello-Leitão 1935 K03b
         |    `--*I. humilis Mello-Leitão 1935 K03b
         |--Iguassoides Soares & Soares 1954 K03b
         |    `--*I. lucidus (Soares 1944) [=Iguassua lucida] K03b
         |--Itatiaincola Soares & Soares 1948 K03b
         |    `--*I. nanus Soares & Soares 1948 K03b
         |--Izecksohnopilio Soares 1977 K03b
         |    `--*I. eugenioi Soares 1977 K03b
         |--Junicus Goodnight & Goodnight 1947 [incl. Sokkupia Mello-Leitão 1949] K03b
         |    `--*J. gerhardi Goodnight & Goodnight 1947 (see below for synonymy) K03b
         |--Juticus Roewer 1943 K03b
         |    `--*J. furcidens Roewer 1943 K03b
         |--Maromba Soares & Soares 1954 K03b
         |    `--*M. dandrettai Soares & Soares 1954 K03b
         |--Metadiscocyrtus Roewer 1929 K03b
         |    `--M. fornicatus (Sørensen 1884) [=Discocyrtus fornicatus] K03b
         |--Metagyndoides Mello-Leitão 1931 K03b
         |    `--*M. granulatus Mello-Leitão 1931 [=Metagyndes granulata, Metagyndes granulatus] K03b
         |--Metaphalangodella Roewer 1915 K03b
         |    `--*M. pachyliformis Roewer 1915 K03b
         |--Oliverius Soares & Soares 1945 K03b
         |    `--*O. jordanensis Soares & Soares 1945 K03b
         |--Osornogyndes Maury 1993 K03b
         |    `--*O. tumifrons Maury 1993 K03b
         |--Pachylusius Mello-Leitão 1949 K08
         |    `--*P. incus Mello-Leitão 1949 K03b
         |--Paraprosontes Soares & Soares 1947 K03b
         |    `--*P. inermis Soares & Soares 1947 K03b
         |--Pareusarcus Roewer 1929 K03b
         |    `--*P. corniculatus Roewer 1929 K03b
         |--Passosa Roewer 1928 K03b, KA-Z11
         |    `--*P. instructa Roewer 1928 K03b, KA-Z11
         |--Pichitus Roewer 1959 K03b
         |    `--*P. aniarmatus Roewer 1959 K03b
         |--Pirunipygus Roewer 1936 K03b
         |    `--*P. paradoxus Roewer 1936 K03b
         |--Planiphalangodus Roewer 1929 K03b
         |    `--*P. robustus Roewer 1929 [=Pachyloides robustus] K03b
         |--Platygyndes Roewer 1943 K03b
         |    `--*P. titicaca Roewer 1943 K03b
         |--Polyacanthoprocta Mello-Leitão 1927 K03b
         |    `--*P. orina (Chamberlin 1916) [=Pachylus orinus] K03b
         |--Pseudoacrographinotus Soares 1966 K03b
         |    `--*P. simplex Soares 1966 K03b
         |--Punagraphinotus Soares & Bauab-Vianna 1973 K03b, KA-Z11
         |    `--*P. punae Soares & Bauab-Vianna 1973 K03b, KA-Z11
         |--Riosegundo Canals 1943 K03b
         |    `--*R. birabeni Canals 1943 K03b
         |--Schubartesia Soares 1944 K03b
         |    `--*S. singularis Soares 1944 K03b
         |--Tarmapachylus Roewer 1956 K03b [=Tarmopachylus (l. c.) KA-Z11]
         |    `--*T. koepckei Roewer 1956 K03b
         |--Temucus Roewer 1943 K03b
         |    `--*T. palpiconus Roewer 1943 K03b
         |--Tingomaria Mello-Leitão 1949 K03b, KA-Z11 [incl. Goyazelloides Soares & Bauab-Vianna 1972 K03b]
         |    `--*T. hydrophila Mello-Leitão 1949 K03b, KA-Z11 (see below for synonymy)
         |--Trochanteroceros Canals 1935 K03b
         |    `--*T. misionicus Canals 1935 K03b
         |--Unduavius Roewer 1929 K03b
         |    `--*U. ornatus Roewer 1929 K03b
         |--Victoriaincola Soares & Soares 1946 K03b
         |    `--*V. penhae Soares & Soares 1946 K03b
         |--Spinivunus Roewer 1943 K03b
         |    `--*S. adumbratus Roewer 1943 K03b
         |--Pherania Strand 1942 [=Phera Sørensen 1932 non Stål 1864, Phaera (l. c.)] K03a
         |    |--*P. pygmaea (Sørensen 1932) [=*Phera pygmaea] K03a
         |    `--P. giupponii Kury 2003 K03a
         |--Soaresia Soares 1945 K08
         |    |--*S. uncina Soares 1945 K08
         |    `--S. forficula Kury 2008 K08
         |--Acanthoprocta Loman 1899 (see below for synonymy) K03b
         |    |--*A. pustulata Loman 1899 (see below for synonymy) K03b
         |    `--A. conica Maury 1991 K03b
         |--Parabalta Roewer 1913 KP-R07, K03b
         |    |--*P. reedii (Butler 1876) [=Gonyleptes reedii, Lycomedes reedii] K03b
         |    `--P. cristobalia (Roewer 1943) [=Neopucroliella cristobalia] K03b
         |--Metapachyloides Roewer 1917 K03b
         |    |--*M. rugosus Roewer 1917 K03b
         |    `--M. almeidai Soares & Soares 1946 K03b
         |--Anoplogynus Piza 1938 [incl. Neotrochantoceros Canals 1943] K03b
         |    |--*A. nasutus Piza 1938 K03b
         |    `--A. singularis (Canals 1943) [=*Neotrochantoceros singularis] K03b
         |--Beckeresia Soares 1970 K03b
         |    |--*B. beckeri Soares 1970 K03b
         |    `--B. delicata Soares & Soares 1970 K03b
         |--Bristoweia Mello-Leitão 1924 K08 [=Bristoweria (l. c.) K03b]
         |    |--*B. diamantinae Mello-Leitão 1924 K03b
         |    `--B. zorodes Mello-Leitão 1924 K03b
         |--Camposicola Mello-Leitão 1924 K08
         |    |--*C. altifrons Mello-Leitão 1924 K03b
         |    `--C. sanctateresae (Soares & Soares 1946) [=Camposicoloides sanctateresae] K03b
         |--Carlotta Roewer 1943 (see below for synonymy) K03b
         |    |--*C. serratipes Roewer 1943 (see below for synonymy) K03b
         |    `--C. dubia (Soares 1944) [=*Pseudoneogonyleptoides dubius] K03b
         |--Ceropachylinus Mello-Leitão 1943 K03b
         |    |--*C. granulosus Mello-Leitão 1943 K03b
         |    `--C. peruvianus Roewer 1956 K08
         |--Diconospelta Canals 1934 K03b
         |    |--*D. gallardoi Canals 1934 K03b
         |    `--D. vazferreirae Mello-Leitão 1946 K03b
         |--Discocyrtulus Roewer 1927 K03b
         |    |--*D. bresslaui Roewer 1927 K03b
         |    `--D. marginalis Roewer 1929 K03b
         |--Eopachylus Mello-Leitão 1931 [=Eupachylus (l. c.)] K03b
         |    |--*E. ignotus Mello-Leitão 1931 K03b
         |    `--E. illectus Soares & Bauab-Vianna 1970 K03b
         |--Gyndoides Mello-Leitão 1927 K03b
         |    |--*G. elaphus Mello-Leitão 1927 K03b
         |    `--G. springmanni Soares & Soares 1947 K03b
         |--Gyndulus Roewer 1929 K03b
         |    |--*G. trispinifrons Roewer 1929 [=Progyndes trispinifrons] K03b
         |    `--G. ‘roeweri’ (Soares & Soares 1954b) (see below for synonymy) K03b
         |--Huralvioides Soares 1970 K03b
         |    |--*H. paruensis Soares 1970 K03b
         |    `--H. hoeferi Kury 1995 K03b
         |--Metalycomedes Mello-Leitão 1927 K03b
         |    |--*M. perditus Mello-Leitão 1927 K03b
         |    `--M. secundus Soares & Soares 1947 K03b
         |--Meteusarcoides Mello-Leitão 1922 [incl. Cercopachylus Piza 1940] K03b
         |    |--*M. mutilatus Mello-Leitão 1922 [incl. Uropachylus grovesi Soares 1944] K03b
         |    `--M. caudatus (Piza 1940) (see below for synonymy) K03b
         |--Palcapachylus Roewer 1952 K03b
         |    |--*P. bicalcariventris Roewer 1952 K03b
         |    `--P. peruvianus Roewer 1959 K03b
         |--Paraluederwaldtia Mello-Leitão 1927 K03b
         |    |--*P. bituberculata (Mello-Leitão 1922) (see below for synonymy) K03b
         |    `--P. caramaschii Soares & Soares 1979 K03b
         |--Kuryella Özdikmen 2006 O06 [=Paranaleptes Soares & Soares 1947 non Breuning 1937 K03b, O06]
         |    |--*K. xanthoacantha (Soares & Soares 1947) [=*Paranaleptes xanthoacanthus] O06
         |    `--K. melanoacantha (Soares & Soares 1947) O06 (see below for synonymy)
         |--Pseudogyndes Mello-Leitão 1932 K03b
         |    |--*P. subsimilis (Roewer 1913) [=Metagyndes subsimilis] K03b
         |    `--P. marginatus Roewer 1961 K03b, KA-Z11 [=P. marginata K03b]
         |--Roeweria Mello-Leitão 1923 (see below for synonymy) K03b
         |    |--*R. bittencourti Mello-Leitão 1923 (see below for synonymy) K03b
         |    `--R. virescens (Mello-Leitão 1940) DG09 (see below for synonymy)
         |--Tumbesia Loman 1899 K03b
         |    |--*T. fuliginosa Loman 1899 K03b
         |    `--T. aculeata Roewer 1930 K03b
         |--Rhioxyna Soares & Bauab-Vianna 1970 K03b (see below for synonymy)
         |    |--*R. zoppeii (Soares 1944) [=*Oxyrhina zoppeii] K03b
         |    `--R. unicornis Soares & Bauab-Vianna 1970 K03b
         |--Yraguara Mello-Leitão 1937 [incl. Melloleitanius Piza 1938] K03b
         |    |--*Y. annulipes Mello-Leitão 1937 [incl. *Melloleitanius barretensis Piza 1938, Y. barretensis] K03b
         |    `--Y. fleuryi Soares 1943 K03b
         |--Eugyndes Roewer 1923 [=Gyndes Sørensen 1884 non Stål 1862, Soerenseniopilio Strand 1942] K03b
         |    |--*E. reinhardi (Sørensen 1884) [=*Gyndes reinhardi, *Soerenseniopilio reinhardi] K03b
         |    |--E. flavolimbatus Soares 1946 K03b
         |    `--E. patellaris Soares 1966 K03b
         |--Ceropachylus Mello-Leitão 1942 [incl. Allelochirus Mello-Leitão 1942, Pachylla Roewer 1959] K03b
         |    |--*C. frizzellae Mello-Leitão 1942 K03b
         |    |--C. areolatus (Roewer 1959) [=*Pachylla areolata] K03b
         |    `--C. singularis (Mello-Leitão 1942) [=*Allelochirus singularis] K03b
         |--Huasampillia Roewer 1913 [=Huasampilla (l. c.); incl. Lucma Roewer 1930] K03b
         |    |--*H. terribilis (Butler 1876) [=Gonyleptes terribilis] K03b
         |    |--H. albipustulata Roewer 1943 K03b
         |    `--H. scotia (Chamberlin 1916) [=Gonyleptes scotius, *Lucma scotia] K03b
         |--Hypophyllonomus Giltay 1928 [incl. Parabristoweia Piza 1938] K03b
         |    |--*H. longipes Giltay 1928 K03b
         |    |--H. callidus Soares 1966 K03b
         |    `--H. maculipalpi (Piza 1938) [=*Parabristoweia maculipalpi] K03b
         |--Ogloblinia Canals 1933 (see below for synonymy) K03b
         |    |--*O. loretoensis Canals 1933 (see below for synonymy) K03b
         |    |--O. argenteopilosa (Mello-Leitão 1935) (see below for synonymy) K03b
         |    `--O. pulchra Pinto-da-Rocha & Soares 1995 K03b
         |--Paradiscocyrtus Mello-Leitão 1927 K03b
         |    |--*P. neglectus Mello-Leitão 1927 K03b
         |    |--P. cerayanus Roewer 1929 K03b
         |    `--P. trochanteralis Roewer 1929 K03b
         |--Pseudogyndesoides Soares 1944 K03b
         |    |--*P. latus Soares 1944 K03b
         |    |--P. bariguiensis Soares 1945 K03b
         |    `--P. pallidus Soares & Soares 1945 K03b
         |--Singram Mello-Leitão 1937 K03b
         |    |--*S. simplex Mello-Leitão 1937 K03b
         |    |--S. brasiliensis (Mello-Leitão 1940) [=Oglobinia brasiliensis] K03b
         |    `--S. singularis Soares & Soares 1948 K03b
         |--Ubatubesia Soares 1945 [incl. Ubatubesiopsis Soares & Soares 1954] K03b
         |    |--*U. oliverioi Soares 1945 [incl. U. travassoi Soares & Soares 1947] K03b
         |    |--U. amplicoxae Pinto-da-Rocha 2000 K03b
         |    `--U. rabelloi (Soares & Soares 1954) [=*Ubatubesiopsis rabelloi] K03b
         |--Lacronia Strand 1942 K08 [=Luederwaldtia Mello-Leitão 1923 non Schmidt 1922 KO06]
         |    |--*L. serripes (Mello-Leitão 1923) [=*Luederwaldtia serripes] KO06
         |    |--L. camboriu Kury 2003 KO06
         |    |--L. ceci Kury & Orrico 2006 KO06
         |    `--L. ricardoi Kury 2003 KO06
         |--Pachyloidellus Müller 1917 K03b, KA-Z11 (see below for synonymy)
         |    |--P. butleri (Thorell 1877) (see below for synonymy) K03b
         |    |--P. borellii CM07
         |    |--P. fulvigranulatus (Mello-Leitão 1930) (see below for synonymy) K03b
         |    `--P. goliath Acosta 1993 K03b
         |--Pygophalangodus Mello-Leitão 1931 K03b (see below for synonymy)
         |    |--*P. gemignanii Mello-Leitão 1931 [=P. geminganii (l. c.)] K03b
         |    |    |--P. g. gemignanii K03b
         |    |    `--P. g. uruguayensis Ringuelet 1955 K03b
         |    `--P. canalsi (Mello-Leitão 1931) (see below for synonymy) K03b
         |--Progyndes Roewer 1917 K03b
         |    |--*P. curvitibialis Roewer 1917 K03b
         |    |--P. basiliscus Mello-Leitão 1935 K03b
         |    |--P. iporangae Soares & Soares 1945 K03b
         |    `--P. trochanteralis Soares & Bauab-Vianna 1972 K03b
         |--Parapachyloides Roewer 1913 (see below for synonymy) K03b
         |    |--*P. uncinatus (Sørensen 1879) [=Gonyleptes uncinatus, Pachyloides uncinatus] K03b
         |    |--P. armatus (Mello-Leitão 1931) (see below for synonymy) K03b
         |    |--P. dentipes Roewer 1913 K03b
         |    `--P. fontanensis (Canals 1935) [=*Apophysigerus fontanensis, Goyazella fontanensis] K03b
         |--Pachylibunus Roewer 1913 K03b
         |    |--*P. grandis Roewer 1913 K03b
         |    |--P. armatissimus Roewer 1917 K03b
         |    |--P. gomesianus Mello-Leitão 1922 K03b
         |    `--P. hamatus Roewer 1930 K03b
         |--Punrunata Roewer 1952 K03b
         |    |--*P. tibialis Roewer 1952 K03b
         |    |--P. brasiliensis Soares & Soares 1979 K03b
         |    |--P. femoralis Soares & Bauab-Vianna 1972 K03b
         |    `--P. pulchra Soares & Soares 1979 K03b
         |--Pucrolia Sørensen 1895 K03b
         |    |--*P. minuta (Sørensen 1884) [=Pachylus minutus] K03b
         |    |--P. dubitata Soares & Soares 1945 K03b
         |    |--P. gracilipes (Canestrini 1888) (n. d.) [=Pachylus gracilipes] K03b
         |    |--P. grandis Mello-Leitão 1930 K03b
         |    `--P. pulcherrima Soares 1966 K03b
         |--Metabalta Roewer 1913 K03b
         |    |--*M. tuberculata Roewer 1913 K03b
         |    |--M. albipes Mello-Leitão 1931 [incl. M. porteri Mello-Leitão 1936] K03b
         |    |--M. efformata Roewer 1929 K03b
         |    |--M. geniculata Roewer 1929 K03b
         |    `--M. hostilis Roewer 1913 K03b
         |--Guaraniticus Mello-Leitão 1933 [incl. Guaranilia Mello-Leitão 1937] K03b
         |    |--*G. lesserti Mello-Leitão 1933 K03b
         |    |--G. flavimaculatus Soares & Soares 1947 K03b
         |    |--G. nigrosulcatus (Mello-Leitão 1937) [=*Guaranilia nigrosulcata] K03b
         |    |--G. olivaceus (Roewer 1913) [=Cobania olivacea] K03b
         |    `--G. tetracalcar Soares & Soares 1945 K03b
         |--Neogonyleptes Roewer 1913 [incl. Neogonyleptoides Roewer 1913] K03b
         |    |--*N. docilis (Butler 1876) [=Gonyleptes docilis] K03b
         |    |--N. chilensis (Roewer 1913) [=Neogonyleptoides chilensis] K03b
         |    |--N. frontalis (Sørensen 1902) [=Gonyleptes frontalis] K03b
         |    |--N. hamatus Soares 1968 K03b
         |    `--N. karschii (Sørensen 1902) [=Gonyleptes karschii, *Neogonyleptoides karschii] K03b
         |--Graphinotus Koch 1839 K03b (see below for synonymy)
         |    |--*G. ornatus Kollar in Koch 1839 K03b (see below for synonymy)
         |    |--G. gratiosus (Soares 1974) [=Wygozinaskya (l. c.) gratiosa] K03b
         |    |--G. magnificus (Roewer 1943) (see below for synonymy) K03b
         |    |--G. roseus (Mello-Leitão 1936) [=Therezopolis roseus] K03b
         |    |--G. sawayai (Soares 1946) [=Therezopolis sawayai] K03b
         |    `--G. viridiornatus (Soares & Soares 1945) [=*Wygodzinskyia viridiornata] K03b
         |--Acrographinotus Holmgren 1916 A02, KA-Z11 (see below for synonymy)
         |    |--*A. erectispina Roewer 1929 A02
         |    |--A. ceratopygus (Soares & Bauab-Vianna 1972) [=*Ctatoproceros ceratopygus] K03b
         |    |--A. curvispina Roewer 1929 A02
         |    |--A. mitmaj Acosta 2002 A02
         |    |--A. niawpaq Acosta 2001 A02
         |    `--A. ortizi (Roewer 1957) A02 [=*Liographinotus ortizi K03b, Sokkupia ortizi K03b]
         |--Pachylus Koch 1839 [=Kochipachylus Strand 1942; incl. Acanthopachyloides Piza 1942] K03b
         |    |--P. chilensis (Gray 1833) (see below for synonymy) K03b
         |    |--P. acanthops (Gervais 1849) [=Gonyleptes acanthops, G. acantops (l. c.)] K03b
         |    |--P. crassus (Roewer 1943) [=Pachyloidellus crassus, Progyndes crassus] K03b
         |    |--P. paessleri Roewer 1913 K03b
         |    |--P. quinamavidensis Muñoz-Cuevas 1969 K03b
         |    `--P. vachoni Muñoz-Cuevas 1970 K03b
         |--Metagraphinotus Mello-Leitão 1927 [incl. Jacarepaguana Mello-Leitão 1932] K03b
         |    |--*M. catharinensis Mello-Leitão 1927 K03b
         |    |--M. arlei Mello-Leitão 1935 K03b
         |    |--M. berlae (Mello-Leitão 1932) [=*Jacarepaguana berlae] K03b
         |    |--M. pectinifemur (Piza 1943) [=Jacarepaguana pectinifemur] K03b
         |    |--M. sooretamae Soares & Soares 1946 K03b
         |    `--M. trochanterspinosus Soares & Soares 1947 K03b
         |--Uropachylus Mello-Leitão 1922 (see below for synonymy) K03b
         |    |--*U. striatus Mello-Leitão 1922 K03b
         |    |--U. anthophilus (Mello-Leitão 1926) [=Ypiranga anthophila] K03b
         |    |--U. gratiosus Soares & Soares 1954 K03b
         |    |--U. itatiaia Soares & Soares 1970 K03b
         |    |--U. nasutus Soares & Soares 1947 [=U. nasutis (l. c.)] K03b
         |    `--U. ypiranga (Mello-Leitão 1922) (see below for synonymy) K03b
         |--Triglochinura Mello-Leitão 1924 (see below for synonymy) K03b
         |    |--*T. curvispina Mello-Leitão 1924 K03b (see below for synonymy)
         |    |--T. ancilla (Mello-Leitão 1937) [=*Tayoca ancilla, Piresa ancilla] K03b
         |    |--T. apiaiensis (Soares & Bauab-Vianna 1972) [=Piresa apiaiensis] K03b
         |    |--T. indicta (Soares & Soares 1970) [=Piresa indicta] K03b
         |    |--T. langei (Soares & Soares 1949) [=Piresa langei] K03b
         |    |--T. timida (Mello-Leitão 1940) [=Itaoca timida] K03b
         |    `--T. villosa (Soares 1966) [=Piresa villosa] K03b
         |--Metagyndes Roewer 1913 K03b
         |    |--*M. martensii (Sørensen 1902) [=Pachylus martensii, M. martensis (l. c.)] K03b
         |    |--M. chilensis Roewer 1943 K03b
         |    |--M. innata Roewer 1929 K03b
         |    |--M. intermedia Roewer 1913 K03b
         |    |--M. laeviscutata Roewer 1943 K03b
         |    |--M. longispina Mello-Leitão 1936 K03b
         |    |--M. pulchella (Loman 1899) [=Gyndes pulchella, Pachylus pulchellus] K03b
         |    `--M. trifidus Mello-Leitão 1943 K03b
         |--Neopucroliella Roewer 1931 [incl. Thorellidia Mello-Leitão 1931] K03b
         |    |--N. pertyi (Thorell 1877) (see below for synonymy) K03b
         |    |--N. bruchi Canals 1935 [=Parabalta bruchi] K03b
         |    |--N. calamuchitaensis Canals 1943 [=Parabalta calamuchitaensis] K03b
         |    |--N. extraordinaria Canals 1943 [=Parabalta extraordinaria] K03b
         |    |--N. mesembrina Ringuelet 1961 [=N. borgmeieri mesembrina] K03b
         |    |--N. nonoensis Canals 1943 [=Parabalta nonoensis] K03b
         |    |--N. ochracea Ringuelet 1963 K03b
         |    `--N. sanctiludovici Acosta 1993 K03b
         `--Neopachylus Roewer 1913 (see below for synonymy) K03b
              |--*N. bellicosus Roewer 1913 K03b
              |--N. herteli Soares & Soares 1945 K03b
              |--N. imaguirei Soares & Soares 1947 K03b
              |--N. incertus (Mello-Leitão 1935) [=*Huralvius incertus, Xuraloius (l. c.) incertus] K03b
              |--N. mamillosus Roewer 1915 K03b
              |--N. marginatus (Mello-Leitão 1931) [=*Gephyropachylus marginatus] K03b
              |--N. nebulosus (Mello-Leitão 1936) [=*Nunduavius nebulosus] K03b
              |--N. serrinha Soares & Soares 1947 K03b
              `--N. taioensis Soares 1966 K03b

Acanthopachylus Roewer 1913 [=Acanthpachylus; incl. Guascaia Mello-Leitão 1935, Heteropachyloidellus Mello-Leitão 1927] K03b

*Acanthopachylus aculeatus (Kirby 1819) K03b, KA-Z11 [=Gonyleptes aculeatus K03b, Acanthpachylus (l. c.) aculeatus K03b, Pachylus aculeatus K03b; incl. ‘Faucheur acanthure’ Duméril 1816-1830 K03b, Gonyleptes acanthurus K03b, *Heteropachyloidellus dimorphicus Mello-Leitão 1927 K03b, H. marginatus Mello-Leitão 1932 K03b, Pachylus mesopotamogalis Weyenbergh 1878 K03b, H. paucigranulatus Mello-Leitão 1931 K03b, Gonyleptes robustus Holmberg 1876 K03b, Pachylus robustus K03b, Heteropachyloidellus robustus Roewer 1943 non Gonyleptes robustus Holmberg 1876 K03b, H. crassus (l. c.) K03b, H. serrulatus Mello-Leitão 1931 K03b, Pachyloidellus tricalcaratus Roewer 1923 K03b, Progyndes tricalcaratus K03b, *Guascaia ypsilonota Mello-Leitão 1935 K03b, Guascaia ypsilonata (l. c.) K03b, Guascaya (l. c.) ypsilonota K03b]

Acanthoprocta Loman 1899 [=Acantoprocta (l. c.), Acantrhoprocta (l. c.); incl. Parapachylus Roewer 1913] K03b

*Acanthoprocta pustulata Loman 1899 [=Acanthroprocta (l. c.) pustulata, Pachylus pustulatus; incl. Parapachylus bispinifrons Roewer 1913, Pachyloides glabrio Loman 1899, *Par. glabrio, Pachylus glabrio] K03b

Acrographinotus Holmgren 1916 A02, KA-Z11 [incl. Ctatoproceros Soares & Bauab-Vianna 1973 K03b, KA-Z11, Liographinotus Roewer 1957 A02]

*Berlaia dissimilis Mello-Leitão 1940 [incl. Echinopachylus brevipes Mello-Leitão in Soares & Soares 1987 (n. n.), Joffilyana nasuta Mello-Leitão in Soares & Soares 1987 (n. n.), Berlaia spinulosa Mello-Leitão 1940] K03b

Bourguyia Mello-Leitão 1923 [incl. Afranius Mello-Leitão 1934, Drastus Roewer 1943, Isopucrolia Mello-Leitão 1927, Osasco Piza 1938] K03b

*Bourguyia albiornata Mello-Leitão 1923 [incl. Isopucrolia conspersa Mello-Leitão 1937, B. curvipes Mello-Leitão 1923, I. meleagris Mello-Leitão in Soares 1945 (n. n.), I. tripos Mello-Leitão 1937, *Osasco tuberiproctus Piza 1938, Afranius turberiproctus, *Isopucrolia uniformis Mello-Leitão 1923] K03b

Carlotta Roewer 1943 [incl. Pseudoneogonyleptoides Soares 1944, Quixaba Mello-Leitão 1944, Quixabella Mello-Leitão 1949, Ullia Roewer 1943] K03b

*Carlotta serratipes Roewer 1943 [incl. *Quixaba atrolutea Mello-Leitão 1944, *Ullia furcata Roewer 1943, Carlotta furcata, *Quixabella ornata Mello-Leitão 1949] K03b

*Ceratoleptes proboscis Soares & Soares 1979 [incl. Ancylorhinoleptes assimilis Mello-Leitão in Kury 2003 (n. n.)] K03b

Cobania Roewer 1913 P-R02, K03b [incl. Crypturocytia Mello-Leitão 1932 K03b, Flangeia Mello-Leitão 1933 K03b, Liogyndulus Mello-Leitão 1931 K03b; Cobaniinae]

*Cobania picea (Bertkau 1880) [=Gonyleptes piceus; incl. *Crypturocytia crypturocytia Mello-Leitão 1932, Lyogonyleptoides (l. c.) curvifemur Roewer 1943, Liogonyleptoides curvifemur, *Liogyndulus luteifemur Mello-Leitão 1932] K03b

Deltaspidium asper (Perty 1833) [=Gonyleptes asper, Ampheres asper; incl. *Deltaspidium bresslaui Roewer 1927] K03b

Gonyleptellus cancellatus (Roewer 1917) [=Gonyleptes cancellatus, Megapachylus cancellatus, Paragonyleptes cancellatus; incl. *Stephanocranion bimaculatus Mello-Leitão 1931, *Gonyleptellus multimaculatus Roewer 1930, Progonyleptoides pustulosus Mello-Leitão 1935, Stephanocranion serrulatum Mello-Leitão 1940] K03b

Graphinotus Koch 1839 K03b [incl. Paraorguesia Mello-Leitão 1927 K03b, Therezopolis Mello-Leitão 1923 K03b, Vitiches Roewer 1927 K03b, Wygodzinskyia Soares & Soares 1945 K03b, Wygodzinskya (l. c.) KA-Z11]

Graphinotus magnificus (Roewer 1943) [=Therezopolis magnifica; incl. T. franciscoi Soares & Soares 1945, Graphinotus franciscoi] K03b

*Graphinotus ornatus Kollar in Koch 1839 K03b [incl. *Paraorguesia albiornata Mello-Leitão 1927 K03b, *Therezopolis therezopolis Mello-Leitão 1923 K03b, *Vitiches viridilimbata Roewer 1927 K03b, T. viridilimbatus SS49]

Gyndulus ‘roeweri’ (Soares & Soares 1954b) [=Progyndes roeweri non Soares & Soares 1954a, G. curvitibialis Roewer 1943 (nom. inv.)] K03b

*Huadquina huadquinae (Chamberlin 1916) [=Gonoleptes (l. c. for Gonyleptes) huadquinae, Progonyleptoides huadquinae] K03b

*Junicus gerhardi Goodnight & Goodnight 1947 [incl. Acrographinotus luteipalpis Roewer 1957, *Sokkupia olivacea Mello-Leitão 1949, A. olivaceus] K03b

Kuryella melanoacantha (Soares & Soares 1947) O06 [=Paranaleptes melanoacanthus O06, P. xelanoacanthus (l. c.) K03b]

Liogonyleptoides Mello-Leitão 1925 [=Progonyleptoides Mello-Leitão 1922 non Roewer 1917, Lyogonyleptoides (l. c.); incl. Anomaloleptes Mello-Leitão 1935, Pachyleptes Mello-Leitão 1932] K03b

*Liogonyleptoides inermis (Mello-Leitão 1922) [=*Progonyleptoides inermis, Lyogonyleptoides (l. c.) inermis; incl. Liogonyleptoides calcaratus Piza 1936, Progonyleptoides cimex Mello-Leitão 1923, Li. cimex, Lyogonyleptoides (l. c.) cimex, *Anomaloleptes singularis Mello-Leitão 1935] K03b

Megapachylus Roewer 1913 [incl. Gonyleptilus Roewer 1927, Gonyperna Roewer 1930, Pachylometoides Mello-Leitão 1936, Pegada Roewer 1930 non Moore 1892, Pegadana Strand 1942] K03b

*Megapachylus grandis Roewer 1913 [incl. Gonyperna curvispina Mello-Leitão 1933, Paragonyleptes curvispina, *Gonyperna divaricata Roewer 1930, Paragonyleptes divaricatus, *Pachylometoides pugnax Mello-Leitão 1936, *Gonyleptilus umbonatus Roewer 1927, Paragonyleptes umbonatus] K03b

Megapachylus simoni (Roewer 1930) [=Paragonyleptes simoni; incl. *Pegada oedipoda Roewer 1930, *Pegadana oedipoda] K03b

Meteusarcoides caudatus (Piza 1940) [=*Cercopachylus caudatus, Uropachylus caudatus; incl. Meteusarcoides marombaensis Soares & Soares 1954] K03b

Neopachylus Roewer 1913 [incl. Gephyropachylus Mello-Leitão 1931, Huralvius Mello-Leitão 1935, Xuraloius (l. c.), Nunduavius Mello-Leitão 1936] K03b

Neopucroliella pertyi (Thorell 1877) [=Ostracidium pertyi, *Thorellidia pertyi; incl. Neopucrolia borgmeieri Mello-Leitão 1923, *Neopucroliella borgmeieri, Neopucroliella borgmeyer (l. c.), Parabalta borgmeieri] K03b

Neosadocus Mello-Leitão 1926 KP-R07, K03b [incl. Bunoweyhia Mello-Leitão 1935 (nom. inv.) KA-Z11, Ilhania Mello-Leitão 1936 K03b, Mitoperna Roewer 1931 K03b, Polybunos Piza 1943 K03b]

*Neosadocus bufo (Mello-Leitão 1923) [=Sadocus bufo; incl. Bunoweyhia lata Mello-Leitão 1935, Neosadocus latus, B. minor Mello-Leitão 1935, N. minor, *Polybunos tuberculatus Piza 1943] K03b

Ogloblinia Canals 1933 [=Oglobinia (l. c.), Oglobinius (l. c.); incl. Prosampycus Mello-Leitão 1935, Quitete Mello-Leitão 1936] K03b

Ogloblinia argenteopilosa (Mello-Leitão 1935) [=*Prosampycus argenteopilosus, Oglobinia (l. c.) argenteopilosa; incl. *Quitete marginata Mello-Leitão 1936] K03b

*Ogloblinia loretoensis Canals 1933 [=Oglobinia (l. c.) loretoensis; incl. Ogloblinia calcarata Soares & Bauab-Vianna 1970, Ogloblinia dentei Soares 1945, Oglobinia intermedia Soares 1944] K03b

Pachyloidellus Müller 1917 K03b, KA-Z11 [=Pachiloidellus (l. c.) K03b; incl. Doelloa Mello-Leitão 1930 K03b, Petrocchia Mello-Leitão 1933 K03b, Sphaleropachylus Mello-Leitão 1926 K03b, Sphalepopachylus (l. c.) K03b, Spraleropachylus (l. c.) K03b]

Pachyloidellus butleri (Thorell 1877) [=Pachylus butleri, Acanthopachylus butleri, Acanthpachylus (l. c.) butleri, *Sphaleropachylus butleri, S. buttleri (l. c.); incl. *Pachyloidellus fuscus Müller 1917, Progyndes fuscus, Pachylus gouldii Weyenbergh 1878, *Petrocchia lesserti Mello-Leitão 1933, Sphaleropachylus lesserti, Pe. marginata Mello-Leitão in Kury 2003 (n. n.)] K03b

Pachyloidellus fulvigranulatus (Mello-Leitão 1930) [=*Doelloa fulvigranulata, Sphaleropachylus fulvigranulatus] K03b

Pachylus chilensis (Gray 1833) [=Gonyleptes chilensis, Kochipachylus chilensis; incl. G. curvipes Guérin-Méneville 1838, *Pachylus granulatus Kollar in Koch 1839, *Acanthopachyloides patellaris Piza 1942] K03b

*Paraluederwaldtia bituberculata (Mello-Leitão 1922) [=Neopucrolia bituberculata, Isopucrolia (l. c.) bituberculata, Luederwaldtia bituberculata] K03b

Parampheres Roewer 1913 KP-R07, K03b [incl. Callampheres Roewer 1931 K03b, Cezarella Mello-Leitão 1932 K03b, Pertyana Mello-Leitão 1927 K03b]

Parampheres bimaculatus (Mello-Leitão 1932) [=*Cezarella bimaculata, Cearinus corniger bimaculatus, Ilhaia bimaculata] K03b

Parampheres ronae (Mello-Leitão 1927) [=*Pertyana ronae, Per. ronnae (l. c.); incl. Penygorna bimaculata Mello-Leitão 1937 non Cezarella bimaculata Mello-Leitão 1932, Arleius bimaculatus, Ilhaia bimaculata non (Mello-Leitão 1932), Parampheres bimaculatus Roewer 1943 nec Cezarella bimaculata Mello-Leitão 1932 nec Penygorna bimaculata Mello-Leitão 1937, Pa. nigrimanus Mello-Leitão 1933, Ilhaia ringueleti Capocasale 1973] K03b

Parapachyloides Roewer 1913 [incl. Apophysigerus Canals 1935, Goyazella Mello-Leitão 1931, Tabatinguera Mello-Leitão 1935] K03b

Parapachyloides armatus (Mello-Leitão 1931) [=*Goyazella armata; incl. *Tabatinguera insignis Mello-Leitão 1935, Parapachyloides insignis] K03b

*Parapucrolia ocellata Roewer 1917 [incl. *Goytacazia pulchra Mello-Leitão 1942, Goitacazia (l. c.) pulchra, Goytacazio (l. c.) pulchra] K03b

Pygophalangodus Mello-Leitão 1931 K03b [incl. Bacigalupo Mello-Leitão 1933 K03b, Canestrinia Mello-Leitão 1931 nec Berlese 1881 nec Mégnin & Trouessart 1884 K03b, Melloinia Thor 1933 K03b, Melloinio (l. c.) KA-Z11, Melloleitaoella Strand 1932 K03b]

Pygophalangodus canalsi (Mello-Leitão 1931) [=*Canestrinia canalsi, *Melloinia canalsi, *Melloleitaoella canalsi; incl. *Bacigalupo tenax Mello-Leitão 1933] K03b

Rhioxyna Soares & Bauab-Vianna 1970 K03b [=Oxyrhina Soares 1944 non Agassiz 1835 K03b, Oxyrhyna (l. c.) KA-Z11]

Roeweria Mello-Leitão 1923 [incl. Jupuvura Mello-Leitão 1940, Lyopachylus Mello-Leitão 1926, Liopachylus (l. c.), Paranaincola Soares 1943] K03b

*Roeweria bittencourti Mello-Leitão 1923 [=Discocyrtoides bittencourti; incl. D. catarinae Soares 1946, *Lyopachylus mitobatoides Mello-Leitão 1923] K03b

Roeweria virescens (Mello-Leitão 1940) DG09 [=*Jupuvura virescens K03b, Discocyrtoides virescens K03b; incl. Discocyrtus fortis Soares 1945 K03b, *Paranaincola mourei Soares 1943 K03b, Discocyrtus mourei K03b, Lyopachylus mourei K03b]

Schenkelibunus Strand 1932 K03b, KA-Z11 [incl. Friburgoia Mello-Leitão 1932 KA-Z11, Hanseniella Mello-Leitão 1927 non Bagnall 1913 K03b, Ziltaia Mello-Leitão 1936 K03b, Siltaya (l. c.) K03b]

Schenkelibunus impar (Mello-Leitão 1932) KA-Z11 [=*Friburgoia impar K03b, Hanseniella impar K03b; incl. *Ziltaia nigrifemur Mello-Leitão 1936 K03b, Ilhaia nigrifemur K03b]

*Schenkelibunus perditus (Mello-Leitão 1927) K03b, KA-Z11 [=*Hanseniella perdita K03b, Friburgoia perdita K03b]

*Tingomaria hydrophila Mello-Leitão 1949 K03b, KA-Z11 [=*Goyazelloides peruvicus Soares & Bauab-Vianna 1972 K03b]

Triglochinura Mello-Leitão 1924 [incl. Barbiellinia Mello-Leitão 1944 non Bezzi 1922, Itaoca Mello-Leitão 1935, Piresa Roewer 1927, Tayoca Mello-Leitão 1937] K03b

*Triglochinura curvispina Mello-Leitão 1924 K03b [incl. *Piresa armata Roewer 1927 K03b, T. armata A-NM06, *Barbiellinia hirsuta Mello-Leitão 1944 K03b, Piresa hirsuta K03b, *Itaoca melanacantha Mello-Leitão 1935 K03b, P. melanacantha K03b, P. sanctipauli Soares 1945 K03b]

*Uracantholeptes anomalus (Mello-Leitão 1922) [=Paragonyleptes anomalus; incl. *Gonypernoides fragilis Mello-Leitão 1932] K03b

Uropachylus Mello-Leitão 1922 [=Japyra Mello-Leitão 1934, Japira (l. c.), Ypiranga Mello-Leitão 1922, Ipiranga (l. c.)] K03b

Uropachylus ypiranga (Mello-Leitão 1922) [=*Ypiranga ypiranga; incl. Cercopachylus fragilis Piza 1943, *Japyra regularis Mello-Leitão 1934] K03b

*Type species of generic name indicated

REFERENCES

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[CM07] Curtis, D. J., & G. Machado. 2007. Ecology. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 280–308. Harvard University Press: Cambridge (Massachusetts).

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[KA-Z11] Kury, A. B., & M. A. Alonso-Zarazaga. 2011. Addenda and corrigenda to the "Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones)". Zootaxa 3034: 47–68.

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[KP-R07] Kury, A. B., & R. Pinto-da-Rocha. 2007. Gonyleptidae Sundevall, 1833. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 196–203. Harvard University Press: Cambridge (Massachusetts).

[O06] Özdikmen, H. 2006. Nomenclatural changes for some Laniatores (Opiliones) genera: new substitute names and new combinations. Mun. Ent. Zool. 1 (1): 63–68.

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