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Introvert and pharynx of Mafangscolex, a Cambrian palaeoscolecid
- Jie Yang, Martin R Smith, Xi-guang Zhang, Xiao-yu Yang
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- Journal:
- Geological Magazine / Volume 157 / Issue 12 / December 2020
- Published online by Cambridge University Press:
- 21 April 2020, pp. 2044-2050
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Palaeoscolecid worms are widespread in the Palaeozoic period, and are of key importance to understanding the emergence of moulting animals (superphylum Ecdysozoa). However, palaeoscolecids lack a diagnostic set of morphological characters, and as such are unlikely to form a natural (monophyletic) group. Consequently, detailed anatomical study of individual taxa is necessary in order to evaluate the phylogenetic significance of palaeoscolecids. New specimens of Mafangscolex from the Cambrian Stage 3 Xiaoshiba Lagerstätte in Kunming, China, provide the first detailed account of a proboscis in Palaeoscoelcida sensu stricto, a core group of palaeoscolecids characterized by having a tessellating scleritome of phosphatic plates and platelets. The eversible mouthparts of Mafangscolex comprise an armoured, hexaradially symmetrical introvert, a ring of coronal spines and quincuncially arranged pharyngeal armature, with a range of tooth morphologies. Taken together, this configuration strikingly resembles the proboscis arrangement inferred for the ancestral ecdysozoan. The six-fold symmetry represents an important difference from the pentaradial priapulan proboscis. The retention of key aspects of the ancestral ecdysozoan body plan suggests that palaeoscolecids represent a useful window on the earliest stages of ecdysozoan evolution.
Case studies in novel narial anatomy: 3. Structure and function of the nasal cavity of saiga (Artiodactyla: Bovidae: Saiga tatarica)
- Andrew B. Clifford, Lawrence M. Witmer
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- Journal:
- Journal of Zoology / Volume 264 / Issue 3 / November 2004
- Published online by Cambridge University Press:
- 18 October 2004, pp. 217-230
- Print publication:
- November 2004
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Much of the narial anatomy of the enigmatic antelope Saiga tatarica has been described by previous workers. However, the anatomy of the nasal cavity and the causally associated osteological correlates of proboscis structure require closer attention, because these data are integral for both a more comprehensive understanding of saiga functional morphology and more robust reconstructions of proboscis structure in fossil taxa. Saiga and outgroup specimens were subjected to X-ray computed tomographic (CT) imaging, gross dissection and skeletonization. The nasal cavity of saiga is characterized by an enlarged nasal vestibule and basal conchal fold. Many structures (e.g. turbinates, lateral cartilages, mucosal folds, nasolacrimal duct) are retracted caudally to a small area in the caudodorsal part of the nasal cavity proper. The enlarged vestibule is associated laterally and ventrally with paired sacs. The nasal septum is largely membranous and contains a large patch of cavernous tissue that serves as a dynamic baffle modifying the flow of inspired air. Bones comprising the narial margin have modified attachment sites for buccinator group muscles and the reduced lateral cartilages. The premaxilla is greatly modified by the enlarged musculature associated with nasolabial fusion. Maintenance of the topological relationships of narial structures compared to bovid outgroups has resulted in a nasal cavity with much larger area for seromucous glands of the vestibule as well as narial musculature capable of controlling the aperture of the nasal cavity. Maxillolabial muscles and the lateralis nasi act together both to compress the nasal cavity and to control the dilation of the nostrils such that air flow through the cavity is highly modified relative to bovid outgroups. The lateral vestibular recess is an outpocketing of the nasal vestibule that produces supplementary seromucous secretions and seems to have no homologue among outgroups. The enlarged nasal vestibule, lateral vestibular recess, repositioned basal fold, and septal cavernous mass are regarded as a coordinated adaptation to dusty habitats, such that nasal air flow can be dynamically regulated allowing for collection of inspired particulates in the vestibule and thus cleansing of air destined for the lungs.
Case studies in novel narial anatomy: 2. The enigmatic nose of moose (Artiodactyla: Cervidae: Alces alces)
- Andrew B. Clifford, Lawrence M. Witmer
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- Journal:
- Journal of Zoology / Volume 262 / Issue 4 / April 2004
- Published online by Cambridge University Press:
- 01 June 2004, pp. 339-360
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- April 2004
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The facial region of moose Alces alces is highly divergent relative to other cervids and other ruminants. In particular, the narial region forms an expanded muzzle or proboscis that overhangs the mouth. The nose of moose provides a case study in the evolution of narial novelty within a phylogenetically well-resolved group (Cervidae). The function of the nasal apparatus of moose remains enigmatic, and new hypotheses are proposed based on our anatomical findings. Head specimens of moose and outgroup taxa were subjected to medical imaging (CT scanning), vascular injection, gross anatomical dissection, gross sectioning, and skeletonization. Moose noses are characterized by highly enlarged nostrils accompanied by specialized musculature, expanded nasal cartilages, and an increase in the connective-tissue pad serving as the termination of the alar fold. The nostrils are widely separated, and the rhinarium that encircles both nostrils in outgroups is reduced to a tiny central patch in moose. The dorsal lateral nasal cartilage is modified to form a pulley mechanism associated with the levator muscle of the upper lip. The lateral accessory nasal cartilage is enlarged and serves as an attachment site for musculature controlling the aperture of the nostril, particularly the lateralis nasi, the apical dilatators, and the rectus nasi. Bony support for narial structures is reduced. Moose show greatly enlarged nasal cartilages, and the entire osseocartilaginous apparatus is relatively much larger than in outgroups. The nasal vestibule of moose is very large and houses a system of three recesses: one rostral and one caudal to the nostrils, and one associated with the enlarged fibrofatty alar fold. As a result of the expanded nasal vestibule, osseous support for the nasal conchae (i.e. turbinates) has retracted caudally along with the bony nasal aperture. The nasoturbinate and its mucosal counterparts (dorsal nasal concha and rectal fold) are reduced. The upturned maxilloturbinate, however, is associated with an enlarged ventral nasal concha and alar fold. Moose are the only species of cervid with these particular characteristics, indicating that this anatomical configuration is indeed novel. Although functional hypotheses await testing, our anatomical findings and published behavioural observations suggest that the novel narial apparatus of moose probably has less to do with respiratory physiology than with functions pertaining specifically to the nostrils. The widely separated and laterally facing nostrils may enhance stereolfaction (i.e. extracting directional cues from gradients of odorant molecules in the environment), but other attributes of narial architecture (enlarged cartilages, specialized musculature, recesses, fibrofatty pads) suggest that this function may not have been the evolutionary driving force. Rather, these attributes suggest a mechanical function, namely, an elaborated nostril-closing system.
The proboscis of tapirs (Mammalia: Perissodactyla): a case study in novel narial anatomy
- Lawrence M. Witmer, Scott D. Sampson, Nikos Solounias
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- Journal:
- Journal of Zoology / Volume 249 / Issue 3 / November 1999
- Published online by Cambridge University Press:
- 27 February 2001, pp. 249-267
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- November 1999
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The trunk-like proboscis of tapirs provides a prime case study in the evolution of anatomical novelty. Morphological study of this unique structure was undertaken employing several specimens and a combination of analytical techniques: gross anatomical dissection, radiographic imaging and histological sectioning. Evolution of the proboscis of tapirs entailed wholesale transformation of the narial apparatus and facial architecture relative to perissodactyl outgroups. This transformation involved retraction and reduction of the bony and cartilaginous facial skeleton, such that several structures present in outgroups are completely absent in tapirs, including cartilages surrounding the nasal vestibule (e.g. alar and medial accessory cartilages, rostral portion of the nasal septum) and associated musculature (dilatator naris apicalis, lateralis nasi pars ventralis). At the same time, soft tissues surrounding the upper lip and nose became elaborated to form a mobile, fleshy proboscis. Several key facial muscles (e.g. levator labii superioris, levator nasolabialis, caninus, lateralis nasi) have been co-opted to function in movement of the proboscis. The nasal vestibule is expanded and occupies approximately 75% of the nasal cavity. Vestibular expansion has compressed and simplified caudal components of the nasal cavity (e.g. reduction of dorsal and middle nasal conchae, loss of plica recta and plica basalis). The airway has become dorsally arched causing the ventral conchal complex to become inclined relative to the long axis of the skull. A few anatomical enigmas remain, such as the complicated maxilloturbinate that rostrally contacts the nasal septum and vomeronasal organ. Similarly, the meatal diverticulum, despite being both ancient and anatomically complex, has no obvious functional significance; it is clear that it is not homologous to the nasal diverticulum of horses and other equids. The reduction of the osseocartilaginous portion of the proboscis, coupled with expansion of the muscular and connective tissue components, has resulted in an organ that is best interpreted as a muscular hydrostat.