Dolichocebus gaimanensis Is Not a Stem Platyrrhine.

The lower molars of the early Miocene New World monkey Dolichocebus gaimanensis from Patagonia, considered by some to be a stem platyrrhine, are for all intents and purposes indistinguishable from the same teeth preserved in the type specimen of the middle Miocene Laventiana annectens from Colombia. Canine morphology further indicates that Dolichocebus is a cebine, closely related to living Saimiri. The phenetic and phyletic continuity of these extinct species, here classified as congeners, means they are the first fossil platyrrhines that link South America's southern and northern Miocene monkey faunas, possibly as vicariant communities or with ancestral-descendant species that evolved by phyletic evolution. These findings further refute the hypothesis that early Miocene platyrrhines are (1) monophyletic and (2) outside the so-called crown platyrrhine radiation. The connection between these species is additional evidence supporting the hypothesis that the pattern of platyrrhine evolution is notably distinct for the predominance of low-level taxonomic clades (genera, tribes, subfamilies) of prolonged ecophylogenetic stability, which is the essence of the Long-Lineage Hypothesis.

Cranial and endocranial diversity in extant and fossil atelids (Platyrrhini: Atelidae): A geometric morphometric study.

OBJECTIVES: Platyrrhines constitute a diverse clade, with the modern Atelidae exhibiting the most variation in cranial and endocast morphology. The processes responsible for this diversification are not well understood. Here, we present a geometric morphometric study describing variation in cranial and endocranial shape of 14 species of Alouatta, Ateles, Brachyteles, and Lagothrix and two extinct taxa, Cartelles and Caipora. METHODS: We examined cranial and endocranial shape variation among species using images reconstructed from CT scans and geometric morphometric techniques based on three-dimensional landmarks and semilandmarks. Principal components analyses were used to explore variation, including the Procrustes shape coordinates, summing the logarithm of the Centroid Size, the common allometric component, and residual shape components. RESULTS: Differences in endocranial shape are related to a relative increase or decrease in the volume of the neocortex region with respect to brainstem and cerebellum regions. The relative position of the brainstem varies from a posterior position in Alouatta to a more ventral position in Ateles. The shape of both the cranium and endocast of Caipora is within the observed variation of Brachyteles. Cartelles occupies the most differentiated position relative to the extant taxa, especially in regards to its endocranial shape. CONCLUSIONS: The pattern of variation in the extant species in endocranial shape is similar to the variation observed in previous cranial studies, with Alouatta as an outlier. The similarities between Caipora and Brachyteles were unexpected and intriguing given the frugivorous adaptations inferred from the fossil's dentition. Our study shows the importance of considering both extant and fossil species when studying diversification of complex traits.

Early evolutionary diversification of mandible morphology in the New World monkeys (Primate, Platyrrhini).

New World monkeys (order Primates) are an example of a major mammalian evolutionary radiation in the Americas, with a contentious fossil record. There is evidence of an early platyrrhine occupation of this continent by the Eocene-Oligocene transition, evolving in isolation from the Old World primates from then on, and developing extensive morphological and size variation. Previous studies postulated that the platyrrhine clade arose as a local version of the Simpsonian ecospace model, with an early phase involving a rapid increase in morphological and ecological diversity driven by selection and ecological opportunity, followed by a diversification rate that slowed due to niche-filling. Under this model, variation in extant platyrrhines, in particular anatomical complexes, may resemble patterns seen among middle-late Miocene (10-14 Ma) platyrrhines as a result of evolutionary stasis. Here we examine the mandible in this regard, which may be informative about the dietary and phylogenetic history of the New World monkeys. Specifically, we test the hypothesis that the Simpsonian ecospace model applies to the platyrrhine mandible through a geometric morphometric analysis of digital images of the jaws of extant and extinct species, and we compare these results to those obtained using a phylogenetic comparative approach based on extant species. The results show a marked phylogenetic structure in the mandibular morphology of platyrrhines. Principal component analyses highlight the morphological diversity among modern forms, and reveal a similar range of variation for the clade when fossil specimens are included. Disparity-Through-Time analysis shows that most of the shape variation between platyrrhines originated early in their evolution (between 20 and 15 Ma). Our results converge with previous studies of body mass, cranial shape, the brain and the basicranium to show that platyrrhine evolution might have been shaped by an early increase in morphological variation followed by a decelerated rate of diversification and evolutionary stasis.

New cranium of the endemic Caribbean platyrrhine, Antillothrix bernensis, from La Altagracia Province, Dominican Republic.

Recent paleontological collection in submerged caves in the eastern Dominican Republic has yielded new specimens of Antillothrix bernensis. Here we describe a complete cranium of an adult individual (MHD 20) and provide phenetic comparisons to other endemic Caribbean taxa and extant mainland platyrrhines using three-dimensional geometric morphometric methods (3DGM). Qualitative and quantitative comparisons support conclusions based on other recently described fossil material: Antillothrix has a dentition lacking clear dietary specialization, an elongated brain case with strong temporal lines, and a vertically oriented nuchal plane. MHD 20 shares a combination of traits with a previously published subadult specimen (MHD 01) including a deep depression at glabella, dorsoventrally elongated orbits, and a relatively large face. This shared morphology reinforces the taxonomic affinity of the two specimens, with differences between the two likely reflecting the younger ontogenetic age of MHD 01. Comparisons to the extant platyrrhines paint a complicated picture as the results of between-group principal components analyses (bgPCA) indicate that Antillothrix does not share a suite of morphological features exclusively with any one genus. Depending on which bgPC axes are visualized, and which subset of landmarks is included (i.e., only those describing the shape of the face/palate for inclusion of Xenothrix), MHD 20 is most similar in shape to the atelids, Alouatta, Lagothrix, and Brachyteles, or an otherwise "empty" region of shape space. It groups neither with Cebus nor Callicebus, two taxa that Antillothrix has been associated with in previous studies based on much less complete material. The Antillothrix cranium does not exhibit any of the derived characters classically used to diagnose or define any single clade; rather its morphology shares features with multiple platyrrhine groups. This is consistent with the interpretation that Antillothrix preserves a primitive morphology, which accords with the hypothesis positing an early arrival of platyrrhines in the Caribbean.

Ontogeny of the Postorbital Region in Tarsiers and Other Primates.

Bony structure of the postorbital region is a key trait distinguishing major clades of primates. Strepsirrhines share a postorbital bar, and anthropoids share a complete postorbital septum. At issue is whether the partial postorbital septum of tarsiers unites living tarsiers more closely with anthropoids than with certain large-eyed Eocene fossils. Previously we reported incomplete postorbital closure in tarsiers at birth. In this article, we document comparative analyses of the postorbital region in a broad range of perinatal primates. Virtual reconstructions of microCT data were used to study three-dimensional structure of the perinatal cranium in these taxa. We also describe and illustrate formation of the tarsier partial postorbital septum through the perinatal period using a growth series of Tarsius syrichta. Our results support the hypothesis that partial postorbital septation in the tarsier is secondary to eye hypertrophy. Based on these observations, we propose a structural hypothesis for phylogenetic differences observed in the primate postorbital region. Specifically, we propose that key postorbital traits, including the frontal spur in strepsirrhines and the posterior lamina of the zygomatic in anthropoids, develop as a result of the spatial relationships of brain, eyes, and teeth. Haplorhines are united by expansion of the anterior cranial fossa and loss of the frontal spur. Anthropoids are further united to the exclusion of tarsiers by expansion of the temporal lobes and associated formation of the posterior lamina of the zygomatic. Mechanical forces related to these spatial relationships may be modulated by deep fascia of the orbit to induce formation of the postorbital septum. Anat Rec, 299:1631-1645, 2016. © 2016 Wiley Periodicals, Inc.

Membranous Support for Eyes of Strepsirrhine Primates and Fruit Bats.

Living primates have relatively large eyes and support orbital tissues with a postorbital bar (POB) and/or septum. Some mammals with large eyes lack a POB, and presumably rely on soft tissues. Here, we examined the orbits of four species of strepsirrhine primates (Galagidae, Cheirogaleidae) and three species of fruit bats (Pteropodidae). Microdissection and light microscopy were employed to identify support structures of the orbit. In bats and primates, there are two layers of fascial sheets that border the eye laterally. The outer membrane is the most superficial layer of deep fascia, and has connections to the POB in primates. In fruit bats, which lacked a POB or analogous ligament, the deep fascia is reinforced by transverse ligaments. Bats and primates have a deeper membrane supporting the eye, identified as the periorbita (PA) based on the presence of elastic fibers and smooth muscle. The PA merges with periostea deep within the orbit, but has no periosteal attachment to the POB of primates. These findings demonstrate that relatively big eyes can be supported primarily with fibrous connective tissues as well as the PA, in absence of a POB or ligament. The well-developed smooth muscle component within the PA of fruit bats likely helps to protrude the eye, maintaining a more convergent eye orientation, with greater overlap of the visual fields. The possibility should be considered that early euprimates, and even stem primates that may have lacked a POB, also had more convergent eyes than indicated by osseous measurements of orbital orientation. Anat Rec, 299:1690-1703, 2016. © 2016 Wiley Periodicals, Inc.

Eye Size and Set in Small-Bodied Fossil Primates: A Three-Dimensional Method.

We introduce a new method to geometrically reconstruct eye volume and placement in small-bodied primates based on the three-dimensional contour of the intraorbital surface. We validate it using seven species of living primates, with dry skulls and wet dissections, and test its application on seven species of Paleogene fossils of interest. The method performs well even when the orbit is damaged and incomplete, lacking the postorbital bar and represented only by the orbital floor. Eye volume is an important quantity for anatomic and metabolic reasons, which due to differences in eye set, or position within (or outside) the bony orbit, can be underestimated in living and fossil forms when calculated from aperture diameter. Our Ectopic Index quantifies how much the globe's volume protrudes anteriorly from the aperture. Lemur, Notharctus and Rooneyia resemble anthropoids, with deeply recessed eyes protruding 11%-13%. Galago and Tarsius are the other extreme, at 47%-56%. We argue that a laterally oriented aperture has little to do with line-of-sight in euprimates, as large ectopic eyes can position the cornea to enable a directly forward viewing axis, and soft tissue positions the eyes facing forward in megachiropteran bats, which have unenclosed, open eye sockets. The size and set of virtual eyes reconstructed from 3D cranial models confirm that eyes were large to hypertrophic in Hemiacodon, Necrolemur, Microchoerus, Pseudoloris and Shoshonius, but eye size in Rooneyia may have been underestimated by measuring the aperture, as in Aotus. Anat Rec, 299:1671-1689, 2016. © 2016 Wiley Periodicals, Inc.

1.32 ± 0.11 Ma age for underwater remains constrain antiquity and longevity of the Dominican primate Antillothrix bernensis.

Endemic New World monkeys are an important element of the extinct mammal faunas of the Caribbean's Greater Antilles. Here we report the first geochronometric evidence that the primate Antillothrix bernensis existed in the Dominican Republic during the Pleistocene, based on the uranium-series age of carbonate speleothem that encased a tibia when it was collected in a flooded cave. Three-dimensional geometric morphometrics of laser-scanned living and extinct samples provide evidence to support the hypothesis that this specimen and other Dominican primate tibial remains belong to that same species. U-Th dating of the host cave carbonate returns ages consistently at the 600 ka upper limit of the technique. However, U-Pb, capable of resolving ages of greater antiquity, is more robust in this context, returning a secure age of 1.32 ± 0.11 Ma, which is the oldest chronometric age recorded for a Hispaniolan mammal. While its origins and manner and time of arrival are obscure, the morphometric studies are consistent with phylogenetic analyses that place A. bernensis within the pitheciid clade of the platyrrhines. The species apparently endured for over 1 million years during the climatic perturbations of the Pleistocene, as a frugivorous climbing quadruped, one of two known primate species occupying the hazard prone island of Hispaniola.

Modeling lineage and phenotypic diversification in the New World monkey (Platyrrhini, Primates) radiation.

Adaptive radiations that have taken place in the distant past can now be more thoroughly studied with the availability of large molecular phylogenies and comparative data drawn from extant and fossil species. Platyrrhines are a good example of a major mammalian evolutionary radiation confined to a single continent, involving a relatively large temporal scale and documented by a relatively small but informative fossil record. Here, we present comparative evidence using data on extant and fossil species to explore alternative evolutionary models in an effort to better understand the process of platyrrhine lineage and phenotypic diversification. Specifically, we compare the likelihood of null models of lineage and phenotypic diversification versus various models of adaptive evolution. Moreover, we statistically explore the main ecological dimension behind the platyrrhine diversification. Contrary to the previous proposals, our study did not find evidence of a rapid lineage accumulation in the phylogenetic tree of extant platyrrhine species. However, the fossil-based diversity curve seems to show a slowdown in diversification rates toward present times. This also suggests an early high rate of extinction among lineages within crown Platyrrhini. Finally, our analyses support the hypothesis that the platyrrhine phenotypic diversification appears to be characterized by an early and profound differentiation in body size related to a multidimensional niche model, followed by little subsequent change (i.e., stasis).

The status of platyrrhine phylogeny: a meta-analysis and quantitative appraisal of topological hypotheses.

Phylogenetic or species trees reflect the branching process of lineages and have direct and indirect interest for several branches of evolutionary anthropology. Estimating phylogenetic trees is a necessary first step toward understanding the factors responsible for the ecological and phenotypic diversification of a primate clade. The platyrrhines have become well known as a phylogenetic challenge. Since the 1990s, platyrrhine phylogenetic studies have increasingly analyzed DNA sequences, or other molecular datasets. Several researchers have claimed with confidence that platyrrhine phylogenetic history has been 'resolved' using these molecular data, but the concordance among these studies has never been quantified. Here, we perform a meta-analysis of published platyrrhine trees using topological information and multivariate methods. Specifically, we examine the claim that platyrrhine phylogeny has been determined and explore the relationships between phylogenies and dataset types used for phylogenetic inference (nuclear DNA, mtDNA, Alu sequences, morphology or mixed data). We compare topologies summarizing 31 major neontological studies of the platyrrhines produced since 1975. The analysis reveals that major disparities are rather common among the hypotheses regarding the higher-level relationships of platyrrhines. We also find that the global concordance that appears to emerge at the generic level is less impressive when one looks more finely at particular relationships. Moreover, correspondence among trees appears to be related to the 'type' of dataset analyzed, which suggests that the biological properties of distinct datasets have an inherent influence on the likelihood of producing similar reconstructions of phylogenetic relationships. This serves to remind us that the main questions surrounding the phylogeny reconstruction program begin with experimental design, for both molecular and morphological datasets. Thus, previous claims that platyrrhine genus-level topology have been 'resolved', or that calibrated molecular trees are sufficiently accurate representations of phylogenetic history that they overpower morphological interpretations of fossils, must be considered premature.

Species: beasts of burden.

Ernst Mayr (1904-2005) was the twentieth century's most influential writer to wrestle with the species problem. The following draws heavily on his work, albeit without presumptuously claiming to mirror his thinking or present any original ideas. As a personal meditation, I am thinking mostly of platyrrhines. Following Mayr, I adhere to what is commonly called the Biological Species Concept (BSC) as a way of thinking about a species in the real-world biosphere as a taxon. I also hold to the idea that the Linnaean category called species has the same function as other categories: a linguistic tool for organizing and retrieving information about biodiversity while embodying evolutionary hypotheses. In other words, alpha taxonomy, the area of systematics that involves identifying, naming, and classifying species, is not purely an exercise in either biology or inventory because it involves communication as well. The burdensome work of the species category stems partly from tension created by the several purposes associated with the concept: the objective observation and examination of a fundamental biological phenomenon, the collection and interpretation of data in a selective context of relevance, and the intention to deploy scientific decisions as a form of communication within a dynamic but highly structured language system.

A closer look at the "Protopithecus" fossil assemblages: new genus and species from Bahia, Brazil.

The recently extinct large-bodied New World monkey Protopithecus brasiliensis Lund 1836 was named based on a distal humerus and proximal femur found in the Lagoa Santa cave system in the southeastern Brazilian state of Minas Gerais. These bones are from an animal about twice the size of the largest extant platyrrhines. One hundred and seventy-five years later, a nearly complete skeleton was discovered in the Toca da Boa Vista caves in the neighboring state of Bahia and was allocated to the same taxon as it was the first platyrrhine fossil of comparable size found since the originals. Our detailed study of the equivalent elements, however, reveals important morphological differences that do not correspond to intraspecific variation as we know it in related platyrrhine taxa. The presence of both an expanded brachioradialis flange on the humerus and gluteal tuberosity on the femur of the Bahian skeleton distinguishes it from the Lagoa Santa fossil as well as from all other platyrrhines. Further cranial and postcranial evidence suggests a closer relationship of the former with the alouattine Alouatta, while the limited Lund material fits more comfortably with the ateline clade. Therefore, we propose to limit P. brasiliensis Lund to the distal humerus and proximal femur from Lagoa Santa and erect a new genus and species for the skeleton from Toca da Boa Vista. Cartelles coimbrafilhoi was a large-bodied frugivore with a relatively small brain and diverse locomotor repertoire including both suspension and climbing that expands the range of platyrrhine biodiversity beyond the dimensions of the living neotropical primates.

Fallback foods, preferred foods, adaptive zones, and primate origins.

Appreciation has grown for the impact of tropical forest seasonality and fallback foods on primate diets, behaviors, and morphology. As critically important resources in times of shortage, seasonal fallback foods may have an outsized role in selecting for form and function while the diversity of preferred plant foods has played an equally prominent role in shaping primate evolution. Here, hypotheses of primate origins are examined in the context of food choice models developed by Marshall and Wrangham [2007] and related to the broader concepts of adaptive zones and radiations. The integrated evolution of primate diet and positional behavior is consistent with a growing reliance on angiosperm products--not prey--as preferred and seasonal fallback foods, temporally and phylogenetically coordinated with evolutionary phases of the angiosperm adaptive radiation. Selection for an incisor oriented but non-specialized heterodont dentition, in contrast with most other orders, attests to the universal role of a highly varied vegetation diet as the primates' primary food resource, with diverse physical properties, phenology and high seasonality. A preference by plesiadapiforms for eating small protein- and lipid-rich seeds may have predisposed the primates and advanced angiosperms to diversify their evolving ecological interdependence, which established the primate adaptive zone and became realized more fully with the rise of the modern euprimate and angiosperm phenotypes. The "narrow niche" hypothesis, a recent challenge to the angiosperm co-evolution hypothesis, is evaluated further. Finally, I note support for visual predation as a core adaptive breakthrough for primates or euprimates remains elusive and problematic, especially considering the theoretical framework provided by the Marshall-Wrangham model, updated evidence of primate feeding habits and the counterpoint lessons of the most successful primate predators, the tarsiiforms.

Rethinking Antillothrix: the mandible and its implications.

A mandible of the Hispaniolan primate Antillothrix bernensis, virtually complete and providing the only definitive evidence of the species' lower dentition, has been discovered in a submerged Dominican Republic cave. The new specimen enables a more certain assessment of the species' phylogenetic position than previously possible. It belongs to the same individual as the nearly complete young adult cranium and postrcranial elements found earlier at the same site. Of the extinct Caribbean platyrrhines, the jaw compares well with partial mandibles representing Xenothrix mcgregori, from Jamaica. Among living platyrrhines, it closely resembles Callicebus and Aotus, as documented in a biometric analysis employing three-dimensional geometric morphometrics of Callicebus, Aotus, Pithecia, Chiropotes, Cacajao, Cebus, and Saimiri. The jaw falls within the morphological variability of Callicebus and Aotus in this three-dimensional analysis, is otherwise most similar to Pithecia, and is distinct from cebines. Lower molars resemble the Haitian primate, Insulacebus, a genus known by a full dentition and gnathic fragments with a pattern of derived features also present in Xenothrix. Considering the available craniodental and postcranial evidence, we conclude that Antillothrix is not properly classified as cebid but rather is best grouped with Pitheciidae, an idea long central to discussions of the phylogenetic affinities of the Greater Antillean primates. Since Antillothrix and Insulacebus are more primitive anatomically than the highly modified Xenothrix, it is tempting to surmise that the origins of the latter involved a vicariance or dispersal event via Hispaniola isolating it on Jamaica.

At birth, tarsiers lack a postorbital bar or septum.

Among primates, partial or complete posterior closure of the orbit has been widely accepted as a shared derived characteristic justifying an exclusive tarsier-anthropoid clade, while some regard the tarsier lateral orbit as an elaborated postorbital bar (POB). To test these competing hypotheses while minimizing the confounding effect of tarsier orbital hypertrophy, we compared tarsiers and other primates at early (fetal and newborn) ages using dissection, micro-CT scans and soft tissue histology. Our findings demonstrate unanticipated variation in the anatomy and development of the zygomaticofrontal (ZFA) articulation, which forms the orbit's lateral framework. Tarsiers uniquely exhibit a combination of two features: absence of a pre- and peri-natal frontal spur to join with the zygomatic to form the ZFA; and, the spur's substitution by an elaborate ligament, which envelops the eye laterally as an expansive postorbital membrane (POM) that merges with the anterolateral fontanelle of the lateral cranial vault. In lacking a frontal spur, tarsiers are distinct from strepsirhines, while the ligamentous structure of the POM distinguishes its ZFA from that of anthropoids, which is a typical facial suture at and prior to birth. The POM of tarsiers may be thought of as an accessory fontanelle, a structural compromise that provides flexible stability and spatial separation of bones while anticipating rapid postnatal growth of an enormously enlarged eye. We regard the tarsier POM as part of a neomorphic eyeball hypertrophy complex, and reject the hypothesis of derived homology of the postorbital septa of adult tarsiers and anthropoids on histological, developmental and functional grounds.

New world monkey nightmares: science, art, use, and abuse (?) in platyrrhine taxonomic nomenclature.

The taxonomy and nomenclature of New World monkeys is becoming precariously unstable and impractical, plagued by revisions aimed at conforming to approaches that reject the Biological Species Concept for narrowly construed reasons and resulting in a hyperinflated taxonomy at species (often) and genus (sometimes) levels. This undermines a major goal of classification at the most basic taxonomic levels to ease communication and facilitate research. Since it is difficult to justify extensive changes in terminology without a deeply justified theoretical purpose or without showing what scientific benefits these alterations can bring, working primatologists need not accept this doctrinaire trend. Knowing as little as we do about what a species actually is, does not justify contorting the value of a species nomenclature so that it reflects nothing more than coat color, a node, or endpoint of a dendrogram.

The making of platyrrhine semifolivores: models for the evolution of folivory in primates.

Among living New World monkeys, Howlers and Muriquis are by far the most folivorous. We examine how well the morphology and behavior of Alouatta and Brachyteles conform to leaf-eating adaptational models derived from other studies. Both genera match these expectations unevenly, which suggests a broader conception of primate folivory is in order. Hence the notion of "semifolivory." While their dentitions prove highly sensitive to selection for leaf-eating, core features relating to body size, brain size, ranging behavior and presumed energy budgets are less predictable corollaries. Leaf-eating in atelines and colobines may have evolved from a preadaptive reliance on seed-eating, which would have necessitated comparable gastric adaptations. Fossils suggest semifolivory in the low-energy Howler lineage may have begun with an increase in body size, a relatively small brain and, possibly, a concomitantly enlarged gut, followed by dental adaptations. It may have advanced via body-size reduction, part of a pioneering adaptation in marginal ecologies on the periphery of rich Amazonian habitats or as a strategy to minimize competition among an abundance of frugivores within the lowland forest-perhaps not as a fallback scheme. In the high-energy Muriqui, semifolivory may have evolved in more intensely seasonal, low-yield forests where frugivores were constrained and rare, a model more consistent with the fallback paradigm. The seed-to-leaves evolutionary pathway hypothesized for anthropoid leaf-eaters may be a widespread phenomenon in primates. We propose it is ultimately rooted in a pre-euprimate reliance on the seeds and seed coats of primitive angiosperms before the latter evolved attractive sugary fruits to coax primates into becoming dispersers of seeds, instead consumers.

Evolutionary morphology, platyrrhine evolution, and systematics.

This special volume of the Anatomical Record focuses on the evolutionary morphology of New World monkeys. The studies range from three-dimensional surface geometry of teeth to enamel ultrastructure; from cranioskeletal adaptations for eating leaves and seeds to the histology of taste bud proxies; from the architecture of its bones to the mechanoreceptors of the tail's skin; from the physical properties of wild foods to the feeding biomechanics of jaws and skull; from the shapes of claws and fingertips, and of elbows, to the diversity and morphology of positional behavior; from the vomeronasal organ and its biological roles to links between brains, guts, sociality, and feeding; from the gum-eating adaptations of the smallest platyrrhines to the methods used to infer how big the largest fossil platyrrhines were. They demonstrate the power of combining functional morphology, behavior, and phylogenetic thinking as an approach toward reconstructing the evolutionary history of platyrrhine primates. While contributing new findings pertaining to all the major clades and ecological guilds, these articles reinforce the view that platyrrhines are a coherent ecophylogenetic array that differentiated along niche dimensions definable principally by body size, positional behavior, and feeding strategies. In underlining the value of character analysis and derived morphological and behavioral patterns as tools for deciphering phylogenetic and adaptational history, doubts are raised about a competing small-bore morphological method, parsimony-based cladistic studies. Intentionally designed not to enlist the rich reservoir of platyrrhine evolutionary morphology, an empirical assessment of the costs incurred by this research stratagem reveals inconsistent, nonrepeatable, and often conflicting results.

Relative brain size, gut size, and evolution in New World monkeys.

The dynamics of brain evolution in New World monkeys are poorly understood. New data on brain weight and body weight from 162 necropsied adult individuals, and a second series on body weight and gut size from 59 individuals, are compared with previously published reports based on smaller samples as well as large databases derived from museum records. We confirm elevated brain sizes for Cebus and Saimiri and also report that Cacajao and Chiropotes have relatively large brains. From more limited data we show that gut size and brain mass have a strongly inverse relationship at the low end of the relative brain size scale but a more diffuse interaction at the upper end, where platyrrhines with relatively high encephalization quotients may have either relatively undifferentiated guts or similar within-gut proportions to low-EQ species. Three of the four main platyrrhine clades exhibit a wide range of relative brain sizes, suggesting each may have differentiated while brains were relatively small and a multiplicity of forces acting to maintain or drive encephalization. Alouatta is a likely candidate for de-encephalization, although its "starting point" is difficult to establish. Factors that may have compelled parallel evolution of relatively large brains in cebids, atelids and pitheciids may involve large social group sizes as well as complex foraging strategies, with both aspects exaggerated in the hyper-encephalized Cebus. With diet playing an important role selecting for digestive strategies among the seed-eating pitheciins, comparable in ways to folivores, Chiropotes evolved a relatively larger brain in conjunction with a moderately large and differentiated gut.