Morphological Evidence for the Phylogeny of Cetacea

A cladistic analysis of 54 extant and extinct cetacean taxa scored for 304 morphological characters supports a monophyletic Odontoceti, Mysticeti, Autoceta, and Cetacea. Forcing a sister-group relationship between Mysticeti and Physeteridae, as suggested by some, but not all, molecular studies, requires an additional 72 steps. In agreement with recent molecular studies, morphological data divide extant mysticetes into two clades: Balaenopteroidea (Eschrichtiidae + Balaenopteridae) and Balaenoidea (Balaenidae + Neobalaenidae). Cetotheriopsinae is removed from Cetotheriidae, elevated to Family Cetotheriopsidae, and placed within the Superfamily Eomysticetoidea. All extant mysticetes and all cetotheriids are placed in a new Parvorder Balaenomorpha, which is diagnosed by many morphological characters, including fusion of the anterior and posterior processes of petrosal to ectotympanic bulla, pronounced median keel on palate, and absence of ventral margin of sigmoid process of bulla. Many of the clades within Odontoceti in the most parsimonious trees of this study are at odds with recent phylogenetic analyses. For example, Platanistidae is not closely related to the extinct odontocete families Squalodontidae and Squalodelphinidae. Instead, it is more closely related to extant river-dwelling odontocetes (i.e., Lipotes, Inia), suggesting a single dispersal of odontocetes into freshwater habitats. We found several characters to support Physeteroidea (Physeteridae + Ziphiidae), a taxon considered paraphyletic by several molecular and some morphological analyses. Lack of agreement on the phylogeny within Odontoceti indicates that additional analyses, which include molecular and anatomical data as well as extant and extinct taxa, are needed.     

Conservation of highly repetitive DNA in cetaceans

It is controversial whether odontocetes (toothed whales) and mysticetes (whalebone whales) have a common ancestry. Cetacean karyological uniformity, which is unique among mammalian orders, suggests a monophyletic origin; however, several anatomical authorities have maintained that odontocetes and mysticetes are diphyletic. We investigated the issue using Southern blot hybridization. Two labelled restriction fragment probes from the DNA of the sei whale (a mysticete) were hybridized to restricted DNA of cetacean species representing all extant families except the Eschrichtiidae, the gray whales. The probes hybridized to specific restriction fragments in all odontocete and mysticete materials. Hybridizations showed preservation of hybridization homologies and a striking conservation of the length of highly repeated DNA sequences. The results are compatible with a common ancestry of odontocetes and mysticetes.     

Assessing mandibular shape variation within Gigantopithecus using a geometric morphometric approach

This study provides a survey of mandibular shape in a sample of extant hominoids (Pan, Gorilla, Pongo, and Hylobates), as well as extinct Asian and Eurasian taxa (Ouranopithecus, Sivapithecus, and Gigantopithecus) in order to compare overall shape similarity. Results presented call into question differences in mandible shape recently used to distinguish Gigantopithecus giganteus from Gigantopithecus blacki and to justify resurrecting a different generic designation, "Indopithecus," for the former. It is concluded that while the two large-bodied Asian taxa may have been adapted to slightly different dietary niches with different geographic and temporal ranges, the unique mandibular/dental characters that the two taxa share should not be viewed as independent evolutionary developments.     

THE EVOLUTIONARY HISTORY OF CETACEAN BRAIN AND BODY SIZE

Cetaceans rival primates in brain size relative to body size and include species with the largest brains and biggest bodies to have ever evolved. Cetaceans are remarkably diverse, varying in both phenotypes by several orders of magnitude, with notable differences between the two extant suborders, Mysticeti and Odontoceti. We analyzed the evolutionary history of brain and body mass, and relative brain size measured by the encephalization quotient (EQ), using a data set of extinct and extant taxa to capture temporal variation in the mode and direction of evolution. Our results suggest that cetacean brain and body mass evolved under strong directional trends to increase through time, but decreases in EQ were widespread. Mysticetes have significantly lower EQs than odontocetes due to a shift in brain:body allometry following the divergence of the suborders, caused by rapid increases in body mass in Mysticeti and a period of body mass reduction in Odontoceti. The pattern in Cetacea contrasts with that in primates, which experienced strong trends to increase brain mass and relative brain size, but not body mass. We discuss what these analyses reveal about the convergent evolution of large brains, and highlight that until recently the most encephalized mammals were odontocetes, not primates.     

Comparative anatomy and evolutionary history of suction feeding in cetaceans

Some odontocetes possess unique features of the hyolingual apparatus that are involved in suction feeding. The hyoid bone and associated musculature generates rapid, piston‐like retraction, and depression of the hyoid and tongue. “Capture” suction feeders (e.g., Globicephala) use suction for capturing and swallowing prey. “Combination” feeders (i.e., Lagenorhynchus) use both raptorial feeding (to capture prey) and suction (to ingest prey). In “capture” suction feeders, features of the hyoid and skull have been attributed to creating suction (i.e., large surface area and mandibular bluntness). In addition to odontocetes, a mysticete, the gray whale (Eschrichtius robustus), is considered a benthic suction feeder. However, anatomical studies of purported suction‐feeding structures of the gray whale are lacking. In addition, few studies have utilized evolutionary approaches to understand the history of suction feeding in cetaceans. This study incorporates quantitative and qualitative hyoid and cranial data from 35 extant and 14 extinct cetacean species into a multivariate principal component analysis and comparative phylogenetic analyses. Conclusions from these analyses are that some commonly attributed features (i.e., ventral throat grooves and mandibular bluntness) and one principal component are significantly correlated with suction feeding. Finally, ancestral state reconstructions indicate that suction feeding likely evolved once, early in cetacean evolutionary history.     

Comparative anatomy of the bony labyrinth of extant and extinct porpoises (Cetacea: Phocoenidae)

The inner ear anatomy of cetaceans, now more readily accessible by means of nondestructive high‐resolution X‐ray computed tomographic (CT) scanning, provides a window into their acoustic abilities and ecological preferences. Inner ear labyrinths also may be a source for additional morphological characters for phylogenetic analyses. In this study, we explore digital endocasts of the inner ear labyrinths of representative species of extinct and extant porpoises (Mammalia: Cetacea: Phocoenidae), a clade of some of the smallest odontocete cetaceans, which produce some of the highest‐frequency clicks for biosonar and communication. Metrics used to infer hearing ranges based on cochlear morphology indicate that all taxa considered could hear high‐frequency sounds, thus the group had already acquired high‐frequency hearing capabilities by the Miocene (9–11 Mya) at the latest. Vestibular morphology indicates that extant species with pelagic preferences have similarly low semicircular canal deviations from 90°, values indicating more sensitivity to head rotations. Species with near‐shore preferences have higher canal deviation values, indicating less sensitivity to head rotations. Extending these analyses to the extinct species, we demonstrate a good match between those predicted to have coastal (such as Semirostrum cerutti) preferences and high canal deviation values. We establish new body length relationships based on correlations with inner ear labyrinth volume, which can be further explored among other aquatic mammals to infer body size of specimens consisting of fragmentary material.     

Elbow joint adductor moment arm as an indicator of forelimb posture in extinct quadrupedal tetrapods

Forelimb posture has been a controversial aspect of reconstructing locomotor behaviour in extinct quadrupedal tetrapods. This is partly owing to the qualitative and subjective nature of typical methods, which focus on bony articulations that are often ambiguous and unvalidated postural indicators. Here we outline a new, quantitatively based forelimb posture index that is applicable to a majority of extant tetrapods. By determining the degree of elbow joint adduction/abduction mobility in several tetrapods, the carpal flexor muscles were determined to also play a role as elbow adductors. Such adduction may play a major role during the stance phase in sprawling postures. This role is different from those of upright/sagittal and sloth-like creeping postures, which, respectively, depend more on elbow extensors and flexors. Our measurements of elbow muscle moment arms in 318 extant tetrapod skeletons (Lissamphibia, Synapsida and Reptilia: 33 major clades and 263 genera) revealed that sprawling, sagittal and creeping tetrapods, respectively, emphasize elbow adductor, extensor and flexor muscles. Furthermore, scansorial and non-scansorial taxa, respectively, emphasize flexors and extensors. Thus, forelimb postures of extinct tetrapods can be qualitatively classified based on our quantitative index. Using this method, we find that Triceratops (Ceratopsidae), Anhanguera (Pterosauria) and desmostylian mammals are categorized as upright/sagittally locomoting taxa.     

Histological variability in fossil and recent alligatoroid osteoderms: Systematic and functional implications

Statements about morphological variation in extinct taxa often suffer from insufficient sampling that can be remedied by taking advantage of larger sample sizes provided by related, extant taxa. This analysis quantitatively and qualitatively examines histological and morphological variation of osteoderms from extant and extinct alligatoroid specimens. Statistically significant differences were correlated with changes in osteoderm size and shape. These differences are independent of position on the body, taxonomy, or evolution. Histological variation in alligatoroid osteoderms is due to morphological constraints on the elements themselves, and not taxonomic differences. This has implications for the recognition of histological characters in the osteoderms of extinct archosaur groups that lack extant representatives. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

The anatomy of Dolichocebus gaimanensis, a stem platyrrhine monkey from Argentina

Dolichocebus is known from the type skull encased in a concretion, numerous isolated teeth, parts of two mandibles, and a talus. The specimens come from the Trelew Member (early Miocene, Colhuehuapian South American Land Mammal Age) of the Sarmiento Formation near the village of Gaiman, Chubut Province, Argentina, dated to about 20Ma. We describe all Dolichocebus fossil material using conventional surface anatomy and micro-CT data from the cranium. The new material and newly imaged internal anatomy of the skull demonstrate that anatomical characters hitherto supposed to support a phyletic link between Dolichocebus and either callitrichines (marmosets, tamarins, and Callimico) or Saimiri (squirrel monkeys) are either indeterminate or absent. To more fully explore the phyletic position of Dolichocebus, we undertook a comprehensive phylogenetic analysis. We examined 268 characters of the cranium and dentition of 16 living platyrrhine genera, some late Oligocene and early Miocene platyrrhines, Tarsius, some Eocene and Oligocene stem anthropoids, and several extant catarrhines. These analyses consistently indicate that Dolichocebus is a stem platyrrhine, as are late Oligocene Branisella and early Miocene Tremacebus, Soriacebus, and Carlocebus. Platyrrhine evolution often is conceived of as a single ancient adaptive radiation. Review of all available phyolgenetic data suggests a more layered evolutionary pattern, with several independent extinct clades filling modern platyrrhine niche space, and modern platyrrhine families and subfamilies appearing over a nine-million-year interval in the Miocene. The outcome of these analyses highlights the pervasiveness of homoplasy in dental and cranial characters. Homoplasy is a real evolutionary phenomenon that is present at all levels of biological analysis, from amino-acid sequences to aspects of adult bony morphology, behavior, and adaptation.     

The Postcranial Musculoskeletal System of Xenarthrans: Insights from over Two Centuries of Research and Future Directions

Xenarthrans stand out among mammals for various reasons, one of them being their musculoskeletal postcranial specializations. Extant armadillos, anteaters, and sloths feature archetypical adaptations to digging and/or diverse arboreal lifestyles. Numerous extinct xenarthrans dramatically depart in size and morphology from their extant relatives, which has sparked functional interpretations since the end of the eighteenth century. Here, we review the diverse methodological approaches that have been used to investigate functional aspects of the postcranial musculoskeletal system in extant and extinct xenarthrans. Specifically, we address qualitative and quantitative bone morphology (including geometric morphometrics), body size and allometry, bone inner structure, myology, as well as in vivo, ex vivo, and in silico experimentation. Finally, a short account is given on those analyses that included xenarthrans to gain insight into primate anatomy. This review helped to identify potential future directions for the functional analysis of the xenarthran anatomy, a tradition over two centuries old.     

Specialize or risk disappearance – empirical evidence of anisomerism based on comparative and developmental studies of gnathostome head and limb musculature

William K. Gregory was one of the most influential authors defending the existence of an evolutionary trend in vertebrates from a higher degree of polyisomerism (more polyisomeric or ‘serial’ anatomical structures arranged along any body axis) to cases of anisomerism (specialization or loss of at least some original polyisomeric structures). Anisomerism was the subject of much interest during the 19th and the beginning of the 20th centuries, particularly due to the influence of the Romantic German School and the notion of ‘primitive archetype’ and because it was conceptually linked to other crucial biological issues (e.g. complexity, scala naturae, progress, modularity or phenotypic integration). However, discussions on anisomerism and related issues (e.g. Williston's law) have been almost exclusively based on hard tissues. Here we provide the first detailed empirical test, and discussion, of anisomerism based on quantitative data obtained from phylogenetic and comparative analyses of the head and forelimb muscles of gnathostomes. Our results strongly support the existence of such a trend in both forelimb and head musculature. For instance, the last common ancestor (LCA) of extant tetrapods likely had 38 polyisomeric muscles (PMs) out of a total of 70 forelimb muscles (i.e. 54%), whereas in the LCAs of extant amniotes and of mammals these numbers were 38/73 (52%) and 21/67 (31%), and in humans are 11/59 (19%). Interestingly, the number of PMs that became specialized during the forelimb evolutionary transition from the LCA of extant tetrapods to humans (13) is very similar to the number of PMs that became lost (14), indicating that both specialization and loss contributed equally to the trend towards anisomerism. By contrast, during the evolution of the head musculature from the LCA of gnathostomes to humans a total of 27 PMs were lost whereas only one muscle became specialized. Importantly, the evolutionary trend towards anisomerism is not related to a general trend leading to the presence of fewer muscles in derived taxa, because for instance humans have more head muscles in total, but many less head polyisomeric muscles than early gnathostomes and extant fish such as sharks, and than early tetrapods and amphibians such as salamanders. This is because new muscles have also been acquired during gnathostome evolution (e.g. facial muscles of mammals). Interestingly, many new PMs have also been acquired during head evolution (but subsequently lost during the transitions towards humans), whereas only a few new PMs were acquired during forelimb evolution. Our comparisons and review of the literature indicate that there is also a trend towards anisomerism during development, thus providing a further example of a parallel between ontogeny and phylogeny, e.g. some forelimb PMs (e.g. contrahentes, intermetacarpales) become specialized or lost (re‐absorbed) during human ontogeny and some head PMs (e.g. constrictores branchiales) become lost during salamander ontogeny. This review will inform future discussions on modularity, complexity, body plans, phenotypic integration and macroevolution, which should ideally include soft tissues and the use of new tools (e.g. anatomical networks) in order to provide a broader and more integrative understanding of these relevant subjects.     

Morphological variation among the inner ears of extinct and extant baleen whales (Cetacea: Mysticeti)

Living mysticetes (baleen whales) and odontocetes (toothed whales) differ significantly in auditory function in that toothed whales are sensitive to high‐frequency and ultrasonic sound vibrations and mysticetes to low‐frequency and infrasonic noises. Our knowledge of the evolution and phylogeny of cetaceans, and mysticetes in particular, is at a point at which we can explore morphological and physiological changes within the baleen whale inner ear. Traditional comparative anatomy and landmark‐based 3D‐geometric morphometric analyses were performed to investigate the anatomical diversity of the inner ears of extinct and extant mysticetes in comparison with other cetaceans. Principal component analyses (PCAs) show that the cochlear morphospace of odontocetes is tangential to that of mysticetes, but odontocetes are completely separated from mysticetes when semicircular canal landmarks are combined with the cochlear data. The cochlea of the archaeocete Zygorhiza kochii and early diverging extinct mysticetes plot within the morphospace of crown mysticetes, suggesting that mysticetes possess ancestral cochlear morphology and physiology. The PCA results indicate variation among mysticete species, although no major patterns are recovered to suggest separate hearing or locomotor regimes. Phylogenetic signal was detected for several clades, including crown Cetacea and crown Mysticeti, with the most clades expressing phylogenetic signal in the semicircular canal dataset. Brownian motion could not be excluded as an explanation for the signal, except for analyses combining cochlea and semicircular canal datasets for Balaenopteridae. J. Morphol. 277:1599–1615, 2016. © 2016 Wiley Periodicals, Inc.     

The position of Cetacea within mammalia: phylogenetic analysis of morphological data from extinct and extant taxa

Knowledge of the phylogenetic position of the order Cetacea (whales, dolphins, and porpoises) within Mammalia is of central importance to evolutionary biologists studying the transformations of biological form and function that accompanied the shift from fully terrestrial to fully aquatic life in this clade. Phylogenies based on molecular data and those based on morphological data both place cetaceans among ungulates but are incongruent in other respects. Morphologists argue that cetaceans are most closely related to mesonychians, an extinct group of terrestrial ungulates. They have disagreed, however, as to whether Perissodactyla (odd-toed ungulates) or Artiodactyla (even-toed ungulates) is the extant clade most closely related to Cetacea, and have long maintained that each of these orders is monophyletic. The great majority of molecule-based phylogenies show, by contrast, not only that artiodactyls are the closest extant relatives of Cetacea, but also that Artiodactyla is paraphyletic unless cetaceans are nested within it, often as the sister group of hippopotamids. We tested morphological evidence for several hypotheses concerning the sister taxon relationships of Cetacea in a maximum parsimony analysis of 123 morphological characters from 10 extant and 30 extinct taxa. We advocate treating certain multistate characters as ordered because such a procedure incorporates information about hierarchical morphological transformation. In all most-parsimonious trees, whether multistate characters are ordered or unordered, Artiodactyla is the extant sister taxon of Cetacea. With certain multistate characters ordered, the extinct clade Mesonychia (Mesonychidae + Hapalodectidae) is the sister taxon of Cetacea, and Artiodactyla is monophyletic. When all fossils are removed from the analysis, Artiodactyla is paraphyletic with Cetacea nested inside, indicating that inclusion of mesonychians and other extinct stem taxa in a phylogenetic analysis of the ungulate clade is integral to the recovery of artiodactyl monophyly. Phylogenies derived from molecular data alone may risk recovering inconsistent branches because of an inability to sample extinct clades, which by a conservative estimate, amount to 89% of the ingroup. Addition of data from recently described astragali attributed to cetaceans does not overturn artiodactyl monophyly.     

Fossil Dolphin Otekaikea marplesi (Latest Oligocene,New Zealand) Expands the Morphological and Taxonomic Diversity of Oligocene Cetaceans

The Oligocene Epoch was a time of major radiation of the Odontoceti (echolocating toothed whales, dolphins). Fossils reveal many odontocete lineages and considerable structural diversity, but whether the clades include some crown taxa or only archaic groups is contentious. The New Zealand fossil dolphin “Prosqualodon” marplesi (latest Oligocene, ≥23.9 Ma) is here identified as a crown odontocete that represents a new genus, Otekaikea, and adds to the generic diversity of Oligocene odontocetes. Otekaikea marplesi is known only from the holotype, which comprises a partial skeleton from the marine Otekaike Limestone of the Waitaki Valley. Otekaikea marplesi was about 2.5 m long; it had procumbent anterior teeth, and a broad dished face for the nasofacial muscles implicated in production of echolocation sounds. The prominent condyles and unfused cervical vertebrae suggest a flexible neck. A phylogenetic analysis based on morphological features places Otekaikea marplesi in the extinct group Waipatiidae, within the clade Platanistoidea. The phylogeny implies an Oligocene origin for the lineage now represented by the endangered Ganges River dolphin (Platanista gangetica), supporting an Oligocene history for the crown Odontoceti.     

Influence of Evolutionary Allometry on Rates of Morphological Evolution and Disparity in strictly Subterranean Moles (Talpinae,Talpidae, Lipotyphla,Mammalia)

The adaptation to a particular function could directly influence the morphological evolution of an anatomical structure as well as its rates. The humeral morphology of moles (subfamily Talpinae) is highly modified in response to intense burrowing and fully fossorial lifestyle. However, little is known of the evolutionary pathways that marked its diversification in the two highly fossorial moles tribes Talpini and Scalopini. We used two-dimensional landmark-based geometric morphometrics and comparative methods to understand which factors influenced the rates and patterns of the morphological evolution of the humerus in 53 extant and extinct species of the Talpini (22 extant plus 12 extinct) and Scalopini (six extant plus 13 extinct) tribes, for a total of 623 humeri. We first built a synthetic phylogeny of extinct and extant taxa of the subfamily Talpinae based on all the available information from known phylogenies, molecular data, and age ranges of fossil records. We tested for evolutionary allometry by means of multivariate regression of shape on size variables. Evolutionary allometric trajectories exhibited convergence of humeral shape between the two tribes, even when controlling for phylogeny, though a significant differences in the evolutionary rates was found between the two tribes. Talpini, unlike Scalopini, seem to have reached a robust fossorial morphology early during their evolution, and their shape disparity did not change, if it did not decrease, through time. Furthermore, the basal Geotrypus spp. clearly set apart from the other highly fossorial moles, exhibiting a significant acceleration of evolutionary shifts toward higher degree of fossorial adaptation. Our observations support the hypothesis that the evolution of allometry may reflect a biological demand (in this case functional) that constrains the rates of evolution of anatomical structures.     

Tooth morphology of Echimyidae (Rodentia,Caviomorpha): homology assessments,fossils, and evolution

Echimyidae constitute the most important radiation of caviomorph rodents in the Neotropical region, represented by 20 extant genera and several extinct species. Both in extant and fossil forms, this diversity is reflected by a significant morphological variation found in crown structures of the cheek teeth. Different hypotheses of primary homology have been proposed for these structures, which, in turn, support diverse dental evolutionary hypotheses. In this contribution we inspect the main structures (cusps and lophids) of the lower deciduous teeth and molars in extinct and extant Echimyidae, and establish their topological correspondences. Comparisons with cusps and lophids of Erethizontidae are emphasized. We explore the testing of alternative primary hypotheses of lophid correspondences in a cladistic context. Following a ‘dynamic’ approach, we select the hypothesis of primary homology, which produced the more parsimonious results, and evaluate the evolutionary transformations of the dental characters analysed. In this context, the phylogenetic relationships of living Myocastor coypus (Molina, 1782) with the extinct Tramyocastor and Paramyocastor are tested. Our results indicate that pentalophodonty is the derived condition for the lower molars in Echimyidae, that trilophodonty evolved independently at least three times during the evolutionary history of these rodents, and that tetralophodonty represents the plesiomorphic condition. This study shows that dental evolution in echimyids can be better understood when occlusal structures are expressed as reliably comparable characters, and when fossils are taken into account. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 451–480.     

Pakicetus inachus and the origin of whales and dolphins (Mammalia: Cetacea)

The present paper is concerned with the comparative morphology of the archeocete and odontocete skull. Among the archeocetes, the recently described lower Eocene Pakicetus inachus obviously represents an early stage of adaptation to aquatic life. The morphology of the incomplete cranial remains, however, gives no evidence that Pakicetus was an amphibious intermediate stage. The evolution of advanced archeocetes and odontocetes is characterized by the successive acquirement of new morphological devices related to the emission and perception of ultrasound under water. The formation of a sonar system in odontocetes obviously not only helped to compensate for the loss of the peripheral olfactory system but moreover was a substantial factor in the evolution of the exceptional dolphin brain.     

Morphometric variation in the hominoid orbital aperture: a case study with implications for the use of variable characters in Miocene catarrhine systematics

Variable characters are ubiquitous in hominoid systematics and present a number of unique problems for phylogenetic analyses that include extinct taxa. As yet, however, few studies have quantified ranges of variation in complex morphometric characters within extant taxa and then used those data to assess the consistency with which discrete character states can be applied to poorly represented fossil species. In this study, ranges of intrageneric morphometric variation in the shape of the hominoid orbital aperture are estimated using exact randomization of average pairwise taxonomic distances (ATDs) derived from size-adjusted centroid, height-width, and elliptic Fourier (EF) variables. Using both centroid and height-width variables, 19 of the 21 possible ATDs between individuals representing seven extinct catarrhine taxa (Aegyptopithecus, Afropithecus, Ankarapithecus, Ouranopithecus, Paranthropus, Sivapithecus and Turkanapithecus) can be observed within a single extant hominoid subspecies, although generally with low probabilities. A resampling study is employed as a means for gauging the effect that this intrataxonomic variation may have on the consistency with which discrete orbital shape character states can be delimited given the small sample sizes available for most Miocene catarrhine taxa preserving this feature (i.e., n=1). For each type of morphometric variable, 100 cluster (UPGMA) analyses of pairwise ATDs are performed in which a single individual is randomly selected from each hominoid genus and analyzed alongside known extinct taxa; consensus trees are computed in order to obtain the frequencies with which different shape clusters appeared in each of the three analyses. The two major clusters appearing most frequently in all three consensus trees are found in only 57% (centroid variables), 49% (height-width variables), and 36% (EF variables) of these trees. If ranges of variation within represented extinct taxa could also be estimated, these frequencies would certainly be far lower. Hominoids clearly exhibit considerable intrageneric, intraspecific, and even intrasubspecific variation in orbit shape, and substantial morphometric overlap exists between taxa; consequently, discrete character states delimiting these patterns of continuous variation are likely to be highly unreliable in phylogenetic analyses of living and extinct species, particularly as the number of terminal taxa increases. Morphological phylogenetic studies of extant catarrhines that assess the effect of different methods (e.g., use of objective a priori weighting or frequency coding of variable characters, inclusion vs. exclusion of variable characters, use of specific vs. supraspecific terminal taxa) on phylogenetic accuracy may help to improve the techniques that systematists employ to make phylogenetic inferences about extinct taxa.