Primates show distinctions in hearing sensitivity and auditory morphology that generally follow phylogenetic patterns. However, few previous studies have attempted to investigate how differences in primate hearing are directly related to differences in ear morphology. This research helps fill this void by exploring the form-to-function relationships of the auditory system in a phylogenetically broad sample of non-human primates. Numerous structures from the outer, middle, and inner ears were measured in taxa with known hearing capabilities. The structures investigated include the overall size and shape of the pinna, the areas of the tympanic membrane and stapedial footplate, the masses and lever arm lengths of the ossicles, the volumes of the middle ear cavities, and the length of the cochlea. The results demonstrate that a variety of auditory structures show significant correlations with certain aspects of hearing (particularly low-frequency sensitivity). Although the majority of these relationships agree with expectations from auditory theory, some traditional (and possibly outdated) ideas were not supported. For example, the common misconception that higher middle ear transformer ratios (e.g., impedance transformer ratio) result in increased hearing sensitivity was not supported. Although simple correlations between form and function do not necessarily imply causality, the relationships defined in this study not only increase our understanding of auditory patterns in extant taxa but also lay the foundation to begin investigating the hearing in fossil primates.
The auditory region contains numerous structures that have proven useful for phylogenetic classification at various taxonomic levels. However, little work has been done in primates relating differences in morphology to variations in hearing performance. This study documents anatomical and physiological distinctions within primates and begins to address the functional and evolutionary consequences of these and other auditory features. The dimensions of the outer ear (pinna) were measured in cadaveric specimens representing nearly every primate family and used to calculate a shape ratio (height/width). It was found that nonanthropoids have a significantly higher ratio than anthropoids, although the actual height was not found to differ. This indicates that most nonanthropoids have ears that are tall and narrow, whereas monkeys and apes are characterized by ears with more equal height and width dimensions. Eardrum area, stapedial footplate area, and ossicular lever arm lengths were measured in dried specimens to calculate an impedance transformer ratio. A distinction was found between anthropoids and strepsirrhines, with the latter group having a transformer ratio indicative of a higher percentage of acoustic energy transmission through the middle ear. Audiogram data were gathered from the literature to analyze hearing sensitivity and it was found that platyrrhines illustrate more low-frequency sensitivity than like-sized lorisoids. The effects of intraspecific variation on the audiogram results were also examined and were found to produce similar results as the analysis using species mean threshold values. Lastly, correlations between morphological and audiogram variables were examined. Several measures of hearing sensitivity were found to be correlated with pinna shape but correlations with middle ear transmission properties were weaker. In addition to using traditional statistical techniques, phylogenetic corrective methods were applied to address the problem of statistical nonindependence of the data and the results of both analyses are compared. These findings are discussed with respect to how sensory adaptations and phylogenetic history may be related to the current radiation of primates. Key words: primate hearing; outer ears; middle ears; audiogram; phylogenyPrimates express numerous modifications on the fundamental mammalian middle ear morphology and these differences have proven useful for phylogenetic classification at various taxonomic levels (MacPhee and Cartmill, 1986). Among extant strepsirrhines, most Malagasy primates are characterized by a free-floating tympanic ring surrounded by a single relatively large tympanic cavity while lorisoids have a tympanic ring that is fused to the lateral wall of a smaller tympanic cavity. Lorises appear to maintain a substantial effective cavity volume by having additional pneumatic spaces off of the epitympanic recess. Extant haplorhines are similar to lorises in possessing epitympanic sinuses (except tarsiers) (MacPhee and Cartmill, 1986) and a fused tym...
The auditory ossicles in primates have proven to be a reliable source of phylogenetic information. Nevertheless, to date, very little data have been published on the metric dimensions of the ear ossicles in African apes and humans. The present study relies on the largest samples of African ape ear ossicles studied to date to address questions of taxonomic differences and the evolutionary transformation of the ossicles in gorillas, chimpanzees and humans. Both African ape taxa show a malleus that is characterized by a long and slender manubrium and relatively short corpus, whereas humans show the opposite constellation of a short and thick manubrium and relatively long corpus. These changes in the manubrium are plausibly linked with changes in the size of the tympanic membrane. The main difference between the incus in African apes and humans seems to be related to changes in the functional length. Compared with chimpanzees, human incudes are larger in nearly all dimensions, except articular facet height, and show a more open angle between the axes. The gorilla incus resembles humans more closely in its metric dimensions, including functional length, perhaps as a result of the dramatically larger body size compared with chimpanzees. The differences between the stapedes of humans and African apes are primarily size-related, with humans being larger in nearly all dimensions. Nevertheless, some distinctions between the African apes were found in the obturator foramen and head height. Although correlations between metric variables in different ossicles were generally lower than those between variables in the same bone, variables of the malleus/incus complex appear to be more strongly correlated than those of the incus/stapes complex, perhaps reflecting the different embryological and evolutionary origins of the ossicles. The middle ear lever ratio for the African apes is similar to other haplorhines, but humans show the lowest lever ratio within primates. Very low levels of sexual dimorphism were found in the ossicles within each taxon, but some relationship with body size and several dimensions of the ear bones was found. Several of the metric distinctions in the incus and stapes imply a slightly different articulation of the ossicular chain within the tympanic cavity in African apes compared with humans. The limited auditory implications of these metric differences in the ossicles are also discussed. Finally, the results of this study suggest that several plesiomorphic features for apes may be retained in the ear bones of the early hominin taxa Australopithecus and Paranthropus as well as in the Neandertals.
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