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A morphometric analysis of pliopithecoid maxillary molar cusp peripheralization Andrew C. Holmes, Jacqueline Garnett, David R. Begun Department of Anthropology, The University of Toronto Introduction Results The Pliopithecoidea is a superfamily of catarrhine primates known from Miocene fossil sites across Eurasia from 18-8 Ma. The most well known and best documented pliopithecoid A B clades are the Pliopithecidae and the Crouzelidae. Despite extensive study, most diagnostic features used to differentiate these groups have remained qualitative.1,2,3,4 This analysis uses geometric measurements to quantify upper molar cusp peripheralization. Our results confirm previous qualitative descriptions and show that crouzeliids have more peripheralized upper molar cusps compared with pliopithecids. We conclude that increased upper molar cusp peripheralization is a derived trait unique to the Crouzeliidae. Materials M1 and M2 were selected because they provide a comprehensive comparative sample of pliopithecoid upper dentition and are known to be phylogenetically informative (M1 n=28, M2 n=30)5 . Pliopithecidae Crouzeliidae Outgroup Pliopithecidae Crouzeliidae Outgroup Figure 3. ROPA measurements for M1 (A) and M2 (B). 2 A B Table 2. M angle Mann-Whitney U results M2 (Angle A) Pliopithecidae Crouzeliidae Crouzeliidae 0.86 Crouzeliidae Outgroup 0.05 0.09 M2 (Angle B) Pliopithecidae Crouzeliidae Crouzeliidae Crouzeliidae 0.42 a b Pliopithecidae Outgroup Outgroup 0.82 0.61 M2 (Angle C) Pliopithecidae Crouzeliidae 2 Figure 1. M a) Pliopithecus platyodon (Göriach 1453) b) Anapithecus hernyaki (SMNS 6089) Pliopithecidae Crouzeliidae 0.04 Our sample represents seven species currently assigned to three pliopithecoid clades: Outgroup Outgroup 0.08 0.004 Crouzeliidae, Pliopithecidae, and Dionysopithecidae (Table 1). Dionysopithecidae (D. M2 (Angle D) Pliopithecidae Crouzeliidae shuangoensis, P. jianghuaiensis) are the earliest, most primitive branch of the Crouzeliidae 0.27 pliopithecoids6. We combined these taxa with two earlier African species, Outgroup 0.76 0.19 Propliopithecus haeckeli and Lomorupithecus harrisoni, to create an outgroup useful for Figure 4. Principal Components Analysis of Length-Breadth (MD/BL) determining whether cusp peripheralization is a primitive or a derived trait for the and ROPA indices for M1 (A) and M2 (B). Pliopithecoidea. Table 1. Taxa used in this analysis. Discussion Pliopithecidae Crouzeliidae Outgroup Our results show that the Crouzeliidae have significantly higher ROPA values for M1 and M2 when compared with the Pliopithecus piveteaui Anapithecus hernyaki Dionysopithecus shuangouensis Pliopithecidae and the outgroup (<0.05). Pliopithecids and the outgroup did not significantly differ (Figure 3). Among the Pliopithecus canmatensis Laccopithecus. robustus Platodontopithecus jianghuaiensis Crouzeliidae, Laccopithecus has the most peripheralized cusps, particularly so in the M2. Consequently, crouzeliid ROPA Pliopithecus zhanxiangi Barberapithecus huezeleri Propliopithecus zeuxis values show a greater range of variation compared with the pliopithecids or the outgroup (Figure 3B, Figure 4B). Even Pliopithecus platyodon Lomorupithecus harrisoni with this large range of variation, pliopithecid and outgroup M2 values are markedly lower and do not overlap Epipliopithecus vindobonensis substantially with those of the Crouzeliidae. Few differences were found in angular cusp measurements among Methods taxonomic categories. The one exception is the M2 cusp angle C (metacone), which is significantly smaller for crouzeliids compared with pliopithecids and the outgroup (<0.05, Table 2). In crouzeliids, the metacone is buccally displaced and We collected a series of linear, closer to the perimeter of the occlusal surface, which in turn causes the metacone angle to be smaller. angular, and area measurements of We used PCA to investigate group specific molar shape patterns by analyzing MD/BL and ROPA (Figure 4). In M1 the molar specimens using ImageJ 1.8.0 B majority of variance is accounted for in PC1 (84%), but in M2 the variance is more evenly split (PC1 = 54%). Because PC1 7 imaging software (Figure 2). High A is loaded primarily by MD/BL and PC2 by ROPA this means that peripheralization can be seen more strongly in M2 than resolution photographs were used M1. Crouzeliids differ significantly from pliopithecids and the outgroup in molar cusp peripheralization, suggesting that (Nikon D7100 with a 60mm Micro OPA increased peripheralization is a crouzeliid synapomorphy. lens). Total Crown Base Area (TCBA) MD is measurement of the total occlusal surface area, and Occlusal Polygon Conclusion C D Area (OPA) is a measurement of a Crouzeliids have more peripheralized upper molar cusps than other pliopithecoids, Propliopithecus and Lomorupithecus, polygon that is created by suggesting that this is a synapomorphy. As both crouzeliids and pliopithecids have been described as primarily connecting the major upper molar frugivorous, the functional significance of cusp peripheralization is not clear and requires further analysis. cusps. When dentin was exposed on a cusp, we estimated the cusp apex BL as being in the center of the exposed Acknowledgments Figure 2. Barberapithecus huerzeleri IPS 1724c dentin8,9. We used the index of OPA Thank you to Museum National d’Histoire Naturelle, Institut Català de Paleontologia Miquel Measurements: Mesiodistal length (MD), Buccolingual relative to TCBA (Relative Occlusal Crusafont, Hungarian Institute of Geology and Geophysics, Naturhistoriches Museum Wien, breadth (BL), TCBA (Total Crown Base Area), OPA Universalmuseum Joanneum, Universität Tübingen, Naturkundemuseum Stuttgart Museum Polygon Area or ROPA) to quantify (Occlusal Polygon Area), Cusp A (Protocone), Cusp B cusp peripheralization (figure 2). (Paracone), Cusp C (Metacone), Cusp D (Hypocone). am Löwentor, Christine Argot, Madelaine Böhme, Ursula Gölich, Dominique Gommery, Martin Angle measurements for OPA were Mesial is up. Groß, Terry Harrison, László Makádi, Klára Palotás, Salvador Moyà-Solà and Reinhard Ziegler. recorded, with each angle named REFERENCES: 1. L. Ginsburg, P. Mein. Bull. Mus. Natl. Hist. Nat. 4: 57–85 (1980). 2. D.M. Alba, S. Moya-Sola. Am. J. Phys. Anthropol. 147(1): 88–112 (2012). 3. T. Relative Occlusal Polygon Area = Harrison in A Companion to Paleoanthropology (Ed. D.R. Begun), p.376-396 (2013). 4. D.R. Begun in in The International Encyclopedia of Primatology. (Ed. A. Fuentes), after the cusp which served as the Occlusal Polygon Area p.1-4. (2017) 5. B. Gamarra, M. Nova Delgado, A. Romero, J. Galbany, A. Pérez-Pérez, Phylogenetic signal in molar dental shape of extant and fossil catarrhine vertex. Total Crown Base Area primates. J. Hum. Evol. 94: 13–27 (2016). 6. T. Harrison, Y. Gu, Y. J. Hum. Evol. 37: 225–277 (1999). 7. W.S. Rasband, W.S. ImageJ (1997-2018). 8. S.E. Bailey. J. Hum. Evol. 183-198 (2004) 9. S.E. Bailey, M. Glantz, T.D. Weaver, B. Viola. J. Hum. Evol. 55: 238-248 (2008)..
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  • Kingdom =Animalia Phylum =Chordata Subphylum =Vertebrata Class =Mammalia Order =Primates I. PARTIAL

    Kingdom =Animalia Phylum =Chordata Subphylum =Vertebrata Class =Mammalia Order =Primates I. PARTIAL

    Kingdom =Animalia '' Phylum =Chordata '' Subphylum =Vertebrata '' Class =Mammalia '' Order =Primates 130-132 I. PARTIAL TAXONOMIC CLASSIFICATION OF PREHISTORIC AND CONTEMPORARY PRIMATES 176-187 197-198/Lab 7.II. Suborder Infraorder Superfamily Family Genus Species Common Name Prosimii [tree shrew = insectivore] (Strepsirhini) lemur 132-134 aye-aye 188-191 loris and bush baby tarsier Anthropoidea Platyrrhini *Parapithecus (basal anthropoid) (Haplorhini) 134-136 *Apidium (basal anthropoid) 134-138 Ceboidea New World monkey 188-191 Catarrhini *Propliopithecus (basal catarrhine) 136-138 *Aegyptopithecus (basal catarrhine) Cercopithecoidea Cercopithecidae Old World monkey 124-126 Macaca macaque Papio baboon Colobidae Colobus colobus monkey Presbytis langur Hominoidea *Proconsulidae *Proconsul 138-143 191-193 Lab 7. II. D. *Oreopithecidea *Oreopithecus Hylobatidae Hylobates gibbon *Pliopithecidae *Pliopithecus Pongidae *Dryopithecus *dryopithecus *Sivapithecus *ramapithecus *kenyapithecus *ouranopithecus *Gigantopithecus Pongo orangutan Panidae Pan traglodytes chimpanzee Pan paniscus bonobo (“pygmy chimpanzee”) Pan ? Gorilla gorilla Mountain gorilla Gorilla Western lowland g. 202-204 Hominidae *Ardipithecus *ramidus 213-218 1 228-237 *Australopithecus *anamensis 241-245 *afarensis Lucy / First Family Lab 8 *africanus southern ape *garhi *[aka Paranthropus]1 *aethiopicus *boisei Zinj *robustus *Kenyanthropus *platyops 1 237-238 Homo *rudolfensis ER-1470 245-247 *habilis human Ch. 11 *erectus Java / Peking “Man” Lab 10 sapiens Mary / John Chs. 12-13 Lab 12 An * marks groups known only through fossils. Compare: FIGURE 6-7 Primate taxonomic classification, p. 131 FIGURE 6-8 Revised partial classification of the primates, p. 132 FIGURE 8-1 Classification chart, modified from Linnaeus p. 177 FIGURE 8-15 Major events in early primate evolution, p. 191 Virtual Lab 1, section II, parts A-D Primate Classification 1(Note: Australopithecus and Paranthropus make up a “Subfamily” called Australopithecinae, more commonly known as Australopithecines.