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Pectoral fin kinematics and motor patterns are shaped by fin ray mechanosensation during steady swimming in Scarus quoyi
<div class="page"> <div class="layoutArea"> <div class="column"> <p>For many species of fish, rhythmic movement of the pectoral fins, or forelimbs, drives locomotion. In terrestrial vertebrates, normal limb- based rhythmic gaits require ongoing modulation with limb mechanosensors. Given the complexity of the fluid environment and dexterity of fish swimming through it, we hypothesize that mechanosensory modulation is also critical to normal fin-based swimming. Here, we examined the role of sensory feedback from the pectoral fin rays and membrane on the neuromuscular control and kinematics of pectoral fin-based locomotion. Pectoral fin kinematics and electromyograms of the six major fin muscles of the parrotfish, Scarus quoyi, a high-performance pectoral fin swimmer, were recorded during steady swimming before and after bilateral transection of the sensory nerves extending into the rays and surrounding membrane. Alternating activity of antagonistic muscles was observed and drove the fin in a figure-of-eight fin stroke trajectory before and after nerve transection. After bilateral transections, pectoral fin rhythmicity remained the same or increased. Differences in fin kinematics with the loss of sensory feedback also included fin kinematics with a significantly more inclined stroke plane angle, an increased angular velocity and fin beat frequency, and a transition to the body-caudal fin gait at lower speeds. After transection, muscles were active over a larger proportion of the fin stroke, with overlapping activation of antagonistic muscles rarely observed in the trials of intact fish. The increased overlap of antagonistic muscle activity might stiffen the fin system in order to enhance control and stability in the absence of sensory feedback from the fin rays. These results indicate that fin ray sensation is not necessary to generate the underlying rhythm of fin movement, but contributes to the specification of pectoral fin motor pattern and movement during rhythmic swimming.</p> <p> </p> </div> </div> </div>
Pectoral fin kinematics and motor patterns are shaped by fin ray mechanosensation during steady swimming in Scarus quoyi
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FIGURE 3 in Scarus maculipinna, a new species of parrotfish (Perciformes, Scaridae) from the eastern Indian Ocean
FIGURE 3. Underwater photographs of Scarus maculipinna. A. Terminal phase male, approximately 16.5 cm TL, Mentawai Islands, Indonesia. Photograph by J. E. Randall. B. Initial phase, approximately 9 cm TL, Mentawai Islands, Indonesia. Photograph by J. E. Randall. C. Terminal phase male, approximately 18 cm TL, Surin Island, Thailand. Photograph by U. Satapoomin.
FIGURE 2 in Scarus maculipinna, a new species of parrotfish (Perciformes, Scaridae) from the eastern Indian Ocean
FIGURE 2. Specimens of Scarus maculipinna in alcohol. A. Holotype of Scarus maculipinna, PMBC 20408, initial phase coloration, 155.2 mm TL, Similan Island, Thailand. B. Paratype of Scarus maculipinna, FMNH 117285, 186.8 mm TL, terminal phase coloration (same specimen as Fig. 1B). photographs by M. W. Westneat.
FIGURE 1 in Scarus maculipinna, a new species of parrotfish (Perciformes, Scaridae) from the eastern Indian Ocean
FIGURE 1. Specimens of Scarus maculipinna. A. Holotype of Scarus maculipinna, PMBC 20408, 155.2 mm TL, Similan Island, Thailand, photograph by U. Satapoomin. B. Paratype of Scarus maculipinna, FMNH 117285, 186.8 mm TL, photograph by M. W. Westneat. C. Paratype of Scarus maculipinna, BPBM 37666, 153.3 mm TL, Mentawai Islands, Indonesia, photograph by J. E. Randall.
FIGURE 17 in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 17. Double-Distributional Multigraph combining area-based (x-axis; Fig. S2-4) and species-based (y-axis; Fig. S2-8) resemblance patterns of 39 parrotfish species among 10 putative Arabian ecoregions (with separate checklists for the Eastern Gulf of Aden and Southern Oman) and 14 additional areas of the wider Western Indian Ocean across the distributional matrix (Supplement 1), columns and rows of which re-ordered according to the sequence arising from both dendrograms; also presenting 'richness-by-area' and 'presence-by-species' sums.
FIGURE 8. Scarus fuscopurpureus. A in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 8. Scarus fuscopurpureus. A: initial phase, Di Hamri, Socotra Island, 2 m depth; B: terminal male, Ras Qatanahn, Socotra Island, 10 m depth.
FIGURE 9. Scarus ghobban. A in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 9. Scarus ghobban. A: initial phase, Di Timri, Socotra Island, 12 m depth; B: initial phase, Di Hamri, Socotra Island, 10 m depth; C: terminal male, Ras Bidou, Socotra Island, 12 m depth.
FIGURE 3. Chlorurus strongylocephalus. A in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 3. Chlorurus strongylocephalus. A: initial phase female, Di Hamri, Socotra Island, 4 m depth; B: initial phase male, Alameh, Socotra Island, 5 m depth; C: terminal male, Roosh, Socotra Island, 14 m depth.
FIGURE 16 in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 16. Resemblance pattern of 10 putative Arabian ecoregions and Kenya (Scarini, 30 spp.) based on Hellinger distance, represented as plots of the (A) non-metric multidimensional scaling analysis (nMDS), overlaid with the cluster analysis and superposed with symbols representing the statistically (ANOSIM) best supported a priori-defined province-level combination X (Supplement 3); and the (B) hierarchical agglomerative cluster analysis according to separate checklists for the Eastern Gulf of Aden and Southern Oman.
FIGURE 7. Scarus frenatus. A in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 7. Scarus frenatus. A: terminal male, Di Hamri, Socotra Island, 3 m depth; B: terminal male, Qariah, Socotra Island, 5 m depth.
FIGURE 11. Scarus rubroviolaceus. A in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 11. Scarus rubroviolaceus. A: small initial phase, Roosh, Socotra Island, 3 m depth; B: initial phase, Di Hamri, Socotra Island, 10 m depth; C: terminal male, Abd al-Kuri Island, Socotra Archipelago, 8 m depth.
FIGURE 6. Scarus ferrugineus. A in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 6. Scarus ferrugineus. A: terminal male, Di Hamri, Socotra Island, 5 m depth; B: terminal male, Roosh, Socotra Island, 14 m depth.
FIGURE 4. Scarus arabicus. A in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 4. Scarus arabicus. A: initial phase, Abd al-Kuri Island, Socotra Archipelago, 7 m depth; B: SMF uncatalogued, initial phase, 26 cm SL, Hadibo fishmarket, Socotra Island; C: SMF uncatalogued, terminal male, 46 cm SL, Hadibo fishmarket, Socotra Island.
FIGURE 14. Scarus zufar. A in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 14. Scarus zufar. A: initial phase, Eryssel, Socotra Island, 12 m depth; B: terminal male, Ras Bidou, Socotra Island, 14 m depth; C: terminal male, Eryssel, Socotra Island, 12 m depth.
FIGURE 37. Scarus trispinosus Valenciennes 1840 in The fish collection of José Mariano da Conceição Veloso (1742-1811) and the beginning of ichthyological research in Brazil, with a taxonomic description of the extant specimens
FIGURE 37. Scarus trispinosus Valenciennes 1840; a: MCUC ZOO.0000009, 310 mm SL; b: MCUC ZOO.0000055, 284 mm SL. Photos by Luis M. P. Ceríaco.
The origin of the parrotfish species Scarus compressus in the Tropical Eastern Pacific: region-wide hybridization between ancient species pairs
<p class="Normal1"><span><span><span><span><span><span><span><span><span><span><span><b>Background: </b></span></span></span></span></span></span></span></span></span></span></span><span><span><span><span><span><span><span><span><span><span><span>An increasing number of hybrid zones with varying evolutionary outcomes have been documented from different reef fish families. In the Tropical Eastern Pacific (TEP), four species of parrotfishes occur in sympatry on rocky reefs from Baja California to Ecuador: <i>Scarus. compressus</i>,<i>S. ghobban</i>, <i>S. perrico</i>, and <i>S. rubroviolaceus</i>; and have complex phylogeographic histories. The most divergent,<i>S. perrico</i>, belongs to a Tropical American clade that diverged from a Central Indo-Pacific ancestor in the late Miocene (6.6 Ma). We tested the hypothesis that <i>S. compressus</i>was the result of ongoing hybridization among the other three species by sequencing four nuclear markers and a mitochondrial locus in samples spanning 2/3 of the latitudinal extent of the TEP. </span></span></span></span></span></span></span></span></span></span></span></p> <p class="Normal1"><span><span><span><span><span><span><span><span><span><span><span><b>Results: </b></span></span></span></span></span></span></span></span></span></span></span><span><span><span><span><span><span><span><span><span><span><span>A structure model of all samples indicated that K=3 was the best fit to the nuclear data and that individuals identified as <i>S. compressus</i>had admixed assignment values (Q). Power analyses indicated our data could correctly detect and assign pure adults and F1 hybrids with > 0.90 probability, and correct assignment of F2 was also high in some cases. NewHybrids models revealed that 89.8% (n= 59) of the <i>Scarus compressus </i>samples were F1 hybrids of crosses between divergent species pairs: <i>S. perrico </i>× <i>S. ghobban</i>and <i>S. perrico </i>× <i>S. rubroviolaceus</i>. Similarly, <i>S.</i><i>ghobban </i>and <i>S. rubroviolaceus</i>were also hybridizing, with ½ of the admixed individuals assigned to F1 hybrids and the remainder likely deep generation hybrids. We observed strong mito-nuclear discordance in all three hybrid pairs, but found little evidence for accelerated mt vs. nuclear evolution in the paternal species. Bayesian analysis of Migrate models favours gene flow between <i>S. perrico</i>and <i>S. ghobban</i>, but not other species pairs. </span></span></span></span></span></span></span></span></span></span></span></p> <p class="Normal1"><span><span><span><span><span><span><span><span><span><span><span><b>Conclusions: </b></span></span></span></span></span></span></span></span></span></span></span><span><span><span><span><span><span><span><span><span><span><span>Mating between species whose ancestors diverged in the late Miocene is giving rise to region wide, hybrid complex, characterized by a high frequency of parental and F1 genotypes but a low frequency of deep generation hybrids. Trimodal structure, combined with reproductive evidence for fertility of both male and female F1 hybrids, suggest that fitness declines sharply in later generation hybrids. In contrast, the hybrid population of the two younger species had similar frequencies of F1 and > F1 hybrids. These differences are consistent with a model of accelerating post-mating incompatibility with time. Mitochondrial genotypes in hybrids, suggests indiscriminate mating by male <i>S. perrico</i>is driving pre-zygotic breakdown, which may reflect the isolation of this endemic species in the TEP for millions of years and weak selection for conspecific mate recognition. Despite overlapping habitat use, high rates of hybridization, and evidence for historical gene flow, species boundaries are maintained by post-mating processes in this complex. </span></span></span></span></span></span></span></span></span></span></span></p>
FIGURE 13 in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 13. Scarus tricolor, terminal male, Roosh, Socotra Island, 14 m depth.
FIGURE 12 in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 12. Scarus scaber, initial phase, Samha Island, Socotra Archipelago, 10 m depth.
FIGURE 2 in Parrotfishes (Teleostei: Labridae: Scarini) of the Socotra Archipelago: Diversity and distributional biogeography, including a range extension of Scarus zufar Randall & Hoover, 1995
FIGURE 2. Chlorurus sordidus, terminal male, Samha Island, Socotra Archipelago, 10 m depth.
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