DOCSLIB.ORG

Terrestrial Force Production by the Limbs of a Semi-Aquatic Salamander Provides Insight Into The

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Terrestrial Force Production by the Limbs of a Semi-Aquatic Salamander Provides Insight Into The bioRxiv preprint doi: https://doi.org/10.1101/2021.05.01.442256; this version posted May 2, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 1 Terrestrial force production by the limbs of a semi-aquatic salamander provides insight into the 2 evolution of terrestrial locomotor mechanics 3 4 Sandy M. Kawano1*, and Richard W. Blob2 5 6 1Department of Biological Sciences, The George Washington University, Washington, D.C. 7 20052, U.S.A. 8 2Department of Biological Sciences, Clemson University, Clemson, SC 29634, U.S.A. 9 10 *Corresponding author. E-mail: [email protected] 11 12 Running title: Kinetics of fins and limbs on land 13 Keywords: biomechanics, ground reaction force, terrestrial locomotion, salamander, fish, water- 14 to-land transition 15 16 # of tables: 7 # of figures: 3 17 Total word count (Intro, Results, Discussion, Acknowledgements, Funding, Conclusion, Figure 18 legends): 1107 + 4012 + 884 = 6003 19 20 Summary statement (15-30 words): Semi-aquatic salamanders had limb mechanics that were 21 intermediate in magnitude yet steadier than the appendages of terrestrial salamanders and 22 semi-aquatic fish, providing a framework to model semi-aquatic early tetrapods. Page 1 of 34 bioRxiv preprint doi: https://doi.org/10.1101/2021.05.01.442256; this version posted May 2, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 23 ABSTRACT 24 Amphibious fishes and salamanders are valuable functional analogs for vertebrates that 25 spanned the water-to-land transition. However, investigations of walking mechanics have 26 focused on terrestrial salamanders and, thus, may better reflect the capabilities of stem 27 tetrapods that were already terrestrial. The earliest tetrapods were aquatic, so salamanders that 28 are not primarily terrestrial may yield more appropriate data for modelling the incipient stages of 29 terrestrial locomotion. In the present study, locomotor biomechanics were quantified from semi- 30 aquatic Pleurodeles waltl, a salamander that spends most of its adult life in water, and then 31 compared to a primarily terrestrial salamander (Ambystoma tigrinum) and semi-aquatic fish 32 (Periophthalmus barbarus) to evaluate whether walking mechanics show greater similarity 33 between species with ecological versus phylogenetic similarities. Ground reaction forces 34 (GRFs) from individual limbs or fins indicated that the pectoral appendages of each taxon had 35 distinct patterns of force production, but hind limb forces were comparable between the 36 salamanders. The rate of force development (‘yank') was sometimes slower in P. waltl but 37 generally comparable between the three species. Finally, medial inclination of the GRF in P. 38 waltl was intermediate between semi-aquatic fish and terrestrial salamanders, potentially 39 elevating bone stresses among more aquatic taxa as they move on land. These data provide a 40 framework for modelling stem tetrapods using an earlier stage of quadrupedal locomotion that 41 was powered primarily by the hind limbs (i.e., “rear-wheel drive”), and reveal mechanisms for 42 appendages to generate propulsion in three locomotor strategies that are presumed to have 43 occurred across the water-to-land transition in vertebrate evolution. Page 2 of 34 bioRxiv preprint doi: https://doi.org/10.1101/2021.05.01.442256; this version posted May 2, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 44 INTRODUCTION 45 The evolutionary invasion of land was a seminal event in vertebrate history that has received 46 intense study, with integrative approaches filling major gaps in our understanding of how this 47 transition transpired (Ashley-Ross et al. 2013). Skeletal remains and trackways in the fossil 48 record can provide some of the most direct evidence about such historical events, yet also 49 present challenges given the limits to which fossils preserve behavior and functional 50 performance (see (Gatesy et al., 1999; Maidment et al., 2014); and references therein). 51 Consequently, extant amphibious fishes, amphibians, and reptiles have been used as functional 52 models to infer the biology of extinct tetrapodomorphs (tetrapods and tetrapod-like 53 sarcopterygian fishes) (Nyakatura et al., 2019, 2014; Pierce et al., 2013), with extant taxa 54 representing alternative strategies for invading land and potentially serving as models for 55 different functional stages as vertebrates became terrestrial. Investigations of extant taxa 56 exhibiting morphological and/or behavioral traits that are consistent with those of extinct 57 tetrapodomorphs offer particularly intriguing opportunities to evaluate how tetrapods were able 58 to leave the water’s edge (Pierce et al., 2013). 59 In considerations of locomotor performance during the invasion of land, salamanders 60 are often used as functional analogues for early tetrapods since they move between water and 61 land (Karakasiliotis et al., 2012), and exhibit a relatively generalized tetrapod Bauplan that has 62 not changed substantially for at least 150 million years (Gao and Shubin, 2001) and possibly as 63 long as 230 million years (Schoch et al., 2020). Previous studies used living salamanders to 64 gain insights into the biomechanics and muscle physiology of walking underwater (Ashley-Ross 65 et al., 2009; Azizi and Horton, 2004; Deban and Schilling, 2009; Frolich and Biewener, 1992) Page 3 of 34 bioRxiv preprint doi: https://doi.org/10.1101/2021.05.01.442256; this version posted May 2, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 66 and on land (Ashley-Ross et al., 2009; Brand, 1996; Deban and Schilling, 2009; Delvolvé et al., 67 1997; Frolich and Biewener, 1992; Kawano and Blob, 2013; Pierce et al., 2020; Sheffield and 68 Blob, 2011), transitioning between water and land (Ashley-Ross and Bechtel, 2004), and 69 assessing how bone histology relates to ecological habits (Canoville and Laurin, 2009; Laurin et 70 al., 2004). Given the greater effect of gravitational loads on the musculoskeletal system in 71 terrestrial environments (Young et al., 2017), one of the most fundamental requirements for 72 moving on land is the ability to support body weight for maintaining posture and physical forces 73 for locomotion. Evaluations of the weight-bearing capabilities in the limbs of early tetrapods 74 have been approached through measurements of ground reaction forces (GRFs) experienced 75 by the tiger salamander, Ambystoma tigrinum, which demonstrated that the hind limbs 76 supported a similar proportion of body weight as the forelimbs but had a greater contribution to 77 acceleration (Kawano and Blob, 2013). To what extent might these results relate to the 78 locomotor function of early tetrapods? Fossil evidence suggests that some of the first tetrapods, 79 such as Acanthostega, were still aquatic (Coates, 1996), and other early tetrapods, such as 80 Ichthyostega, may have had only limited terrestrial capabilities (Pierce et al., 2012). In contrast, 81 A. tigrinum are found in various terrestrial habitats, ranging from conifer forests to deserts, and 82 adults rarely spend extended durations in water for reasons other than reproduction (Petranka, 83 1998). As such, they may not provide an ideal model for the initial invaders of land, in which 84 terrestrial capacity may not have been fully acquired. How might limb function differ for a 85 species that spends most of its time in water? 86 Because salamanders have a diverse range of habitat preferences and life histories 87 (Wake, 2009), they provide an opportunity to model different evolutionary stages in the adoption Page 4 of 34 bioRxiv preprint doi: https://doi.org/10.1101/2021.05.01.442256; this version posted May 2, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 88 of terrestrial habits. Examinations of amphibious species with greater aquatic tendencies could 89 yield substantial insights into the limb function of earlier stem tetrapods with digit-bearing limbs. 90 In this study, we compared GRFs of individual limbs produced by semi-aquatic salamanders 91 (Spanish ribbed newts, Pleurodeles waltl Michahelles 1830; see Electronic Supplementary 92 Materials for details), to published data on terrestrial tiger salamanders, Ambystoma tigrinum 93 Green 1825, and semi-aquatic African mudskippers, Periophthalmus barbarus (Linnaeus 1766) 94 (Kawano and Blob, 2013). We also calculated new measures of yank, the rate of force 95 development (sensu Lin et al., 2019), to evaluate how the rate of force production might differ 96 between these species. Quantifying the dynamics of force-time measures contributes valuable 97 information to explain kinematic and muscle activation patterns (Lin et al., 2019). Our aim in 98 these comparisons was to examine extant taxa that model important stages during the transition 99 to land to quantify the functional differences between fins and limbs for terrestrial locomotion. 100 During aquatic locomotion, dominant force production by the rear appendage (“rear- 101 wheel drive”) may have appeared as early
Load more

Recommended publications
  • Your Inner Fish
    CHAPTER THREE HANDY GENES While my colleagues and I were digging up the first Tiktaalik in the Arctic in July 2004, Randy Dahn, a researcher in my laboratory, was sweating it out on the South Side of Chicago doing genetic experiments on the embryos of sharks and skates, cousins of stingrays. You’ve probably seen small black egg cases, known as mermaid’s purses, on the beach. Inside the purse once lay an egg with yolk, which developed into an embryonic skate or ray. Over the years, Randy has spent hundreds of hours experimenting with the embryos inside these egg cases, often working well past midnight. During the fateful summer of 2004, Randy was taking these cases and injecting a molecular version of vitamin A into the eggs. After that he would let the eggs develop for several months until they hatched. His experiments may seem to be a bizarre way to spend the better part of a year, let alone for a young scientist to launch a promising scientific career. Why sharks? Why a form of vitamin A? 61 To make sense of these experiments, we need to step back and look at what we hope they might explain. What we are really getting at in this chapter is the recipe, written in our DNA, that builds our bodies from a single egg. When sperm fertilizes an egg, that fertilized egg does not contain a tiny hand, for instance. The hand is built from the information contained in that single cell. This takes us to a very profound problem.
    [Show full text]
  • Crawford Reservoir
    Crawford Reservoir FISH SURVEY AND MANAGEMENT INFORMATION Eric Gardunio, Fish Biologist Montrose Service Center General Information: Crawford Reservoir is a popular fishery that provides angling opportunity for yellow perch, channel catfish, northern pike, rainbow trout, black crappie, and largemouth bass. This reser- voir, located in Crawford State Park, covers 414 surface acres at full capacity and is open year round to an- gling. Visit the State Parks website for information on regulations, camping, and recreation: http://parks.state.co.us/Parks/Crawford Location: 2 miles south of the town of Crawford on Hwy 92. Primary Management: Warmwater Mixed Species Lake Category 602 Amenities Previous Stocking Sportfishing Notes 2019 Black Crappie Boat Ramps (2) Rainbow Trout (10”): 9,100 Good spots include the East Campgrounds (2) Largemouth Bass (2”): 30,088 shore primarily around the Showers Clear Fork boat ramp cove or Largemouth Bass (6”): 150 anywhere with brush Visitors Center Largemouth Bass (20”): 70 Good baits include small tube Restrooms Channel Catfish (7”): 1,500 jigs and worms Parking Areas 2018 Channel Catfish Picnic Shelters Rainbow Trout (10”): 12,184 Good spots include the north Largemouth Bass (2”): 30,000 side of peninsula cove and near the dam Channel Catfish (7”): 4,250 Good baits include night 2017 crawlers and cut-bait WARNING !!! Rainbow Trout (10”): 12,184 Largemouth Bass Prevent the Spread of Largemouth Bass (2”): 20,000 Good spots include the rocky Zebra Mussels and other Largemouth Bass (16”): 70 areas near the dam and flood- Aquatic Nuisance Species ed brush and vegetation in the Channel Catfish (9”): 2,000 spring and summer.
    [Show full text]
  • Fish Locomotion: Recent Advances and New Directions
    MA07CH22-Lauder ARI 6 November 2014 13:40 Fish Locomotion: Recent Advances and New Directions George V. Lauder Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138; email: [email protected] Annu. Rev. Mar. Sci. 2015. 7:521–45 Keywords First published online as a Review in Advance on swimming, kinematics, hydrodynamics, robotics September 19, 2014 The Annual Review of Marine Science is online at Abstract marine.annualreviews.org Access provided by Harvard University on 01/07/15. For personal use only. Research on fish locomotion has expanded greatly in recent years as new This article’s doi: approaches have been brought to bear on a classical field of study. Detailed Annu. Rev. Marine. Sci. 2015.7:521-545. Downloaded from www.annualreviews.org 10.1146/annurev-marine-010814-015614 analyses of patterns of body and fin motion and the effects of these move- Copyright c 2015 by Annual Reviews. ments on water flow patterns have helped scientists understand the causes All rights reserved and effects of hydrodynamic patterns produced by swimming fish. Recent developments include the study of the center-of-mass motion of swimming fish and the use of volumetric imaging systems that allow three-dimensional instantaneous snapshots of wake flow patterns. The large numbers of swim- ming fish in the oceans and the vorticity present in fin and body wakes sup- port the hypothesis that fish contribute significantly to the mixing of ocean waters. New developments in fish robotics have enhanced understanding of the physical principles underlying aquatic propulsion and allowed intriguing biological features, such as the structure of shark skin, to be studied in detail.
    [Show full text]
  • Forelimb Position Affects Facultative Bipedal Locomotion in Lizards Chase T
    © 2018. Published by The Company of Biologists Ltd | Journal of Experimental Biology (2018) 221, jeb185975. doi:10.1242/jeb.185975 RESEARCH ARTICLE Forelimb position affects facultative bipedal locomotion in lizards Chase T. Kinsey* and Lance D. McBrayer‡ ABSTRACT 2004; Clark and Higham, 2011; Tucker and McBrayer, 2012; Parker Recent work indicates that bipedal posture in lizards is advantageous and McBrayer, 2016). During predation events or social ’ during obstacle negotiation. However, little is known about how interactions, a terrestrial vertebrate s behavior, speed and stability bipedalism occurs beyond a lizard’s acceleratory threshold. traversing obstacles may impinge upon their survivorship and/or Furthermore, no study to date has examined the effects of forelimb fitness (Stiller and McBrayer, 2013; Schulte et al., 2004; Arnold, position on the body center of mass (BCoM) in the context of 1983; but see Garland and Losos, 1994). bipedalism. This study quantified the frequency of bipedalism when Stereotyped limb movement in quadrupedal locomotion, or gait, sprinting with versus without an obstacle at 0.8 m from the start of a has predictable footfalls across various speeds (Snyder, 1952, 1954, sprint. Forelimb positions were quantified during bipedal running at 1962; Irschick and Jayne, 1999; Farley and Ko, 1997). Some the start of a sprint and when crossing an obstacle. Two species with terrestrial lizards alter their gait and/or posture while sprinting contrasting body forms (and thus different BCoM) were studied (Schuett et al., 2009; for review, see Russell and Bels, 2001). (Sceloporus woodi and Aspidoscelis sexlineata) to assess potential Facultative bipedalism occurs in some quadrupeds when only the variation due to body plan and obstacle-crossing behavior.
    [Show full text]
  • Pterosaurs Flight in the Age of Dinosaurs Now Open 2 News at the Museum 3
    Member Magazine Spring 2014 Vol. 39 No. 2 Pterosaurs Flight in the Age of Dinosaurs now open 2 News at the Museum 3 From the After an unseasonably cold, snowy winter, will work to identify items from your collection, More than 540,000 Marine Fossils the Museum is pleased to offer a number of while also displaying intriguing specimens from President springtime opportunities to awaken the inner the Museum’s own world-renowned collections. Added to Paleontology Collection naturalist in us all. This is the time of year when Of course, fieldwork and collecting have Ellen V. Futter Museum scientists prepare for the summer been hallmarks of the Museum’s work since Collections at a Glance field season as they continue to pursue new the institution’s founding. What has changed, discoveries in their fields. It’s also when Museum however, is technology. With a nod to the many Over nearly 150 years of acquisitions and Members and visitors can learn about their ways that technology is amplifying how scientific fieldwork, the Museum has amassed preeminent own discoveries during the annual Identification investigations are done, this year, ID Day visitors collections that form an irreplaceable record Day in Theodore Roosevelt Memorial Hall. can learn how scientists use digital fabrication of life on Earth. Today, 21st-century tools— Held this year on May 10, Identification Day to aid their research and have a chance to sophisticated imaging techniques, genomic invites visitors to bring their own backyard finds have their own objects scanned and printed on analyses, programs to analyze ever-growing and curios for identification by Museum scientists.
    [Show full text]
  • Amphibious Fishes: Terrestrial Locomotion, Performance, Orientation, and Behaviors from an Applied Perspective by Noah R
    AMPHIBIOUS FISHES: TERRESTRIAL LOCOMOTION, PERFORMANCE, ORIENTATION, AND BEHAVIORS FROM AN APPLIED PERSPECTIVE BY NOAH R. BRESSMAN A Dissertation Submitted to the Graduate Faculty of WAKE FOREST UNIVESITY GRADUATE SCHOOL OF ARTS AND SCIENCES in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY Biology May 2020 Winston-Salem, North Carolina Approved By: Miriam A. Ashley-Ross, Ph.D., Advisor Alice C. Gibb, Ph.D., Chair T. Michael Anderson, Ph.D. Bill Conner, Ph.D. Glen Mars, Ph.D. ACKNOWLEDGEMENTS I would like to thank my adviser Dr. Miriam Ashley-Ross for mentoring me and providing all of her support throughout my doctoral program. I would also like to thank the rest of my committee – Drs. T. Michael Anderson, Glen Marrs, Alice Gibb, and Bill Conner – for teaching me new skills and supporting me along the way. My dissertation research would not have been possible without the help of my collaborators, Drs. Jeff Hill, Joe Love, and Ben Perlman. Additionally, I am very appreciative of the many undergraduate and high school students who helped me collect and analyze data – Mark Simms, Tyler King, Caroline Horne, John Crumpler, John S. Gallen, Emily Lovern, Samir Lalani, Rob Sheppard, Cal Morrison, Imoh Udoh, Harrison McCamy, Laura Miron, and Amaya Pitts. I would like to thank my fellow graduate student labmates – Francesca Giammona, Dan O’Donnell, MC Regan, and Christine Vega – for their support and helping me flesh out ideas. I am appreciative of Dr. Ryan Earley, Dr. Bruce Turner, Allison Durland Donahou, Mary Groves, Tim Groves, Maryland Department of Natural Resources, UF Tropical Aquaculture Lab for providing fish, animal care, and lab space throughout my doctoral research.
    [Show full text]
  • Development and the Evolvability of Human Limbs
    Development and the evolvability of human limbs Nathan M. Younga,1, Günter P. Wagnerb, and Benedikt Hallgrímssonc aDepartment of Orthopaedic Surgery, University of California, San Francisco, CA 94110; bDepartment of Ecology and Evolutionary Biology, Yale University, New Haven, CT 06405; and cDepartment of Cell Biology and Anatomy, University of Calgary, Calgary, AB, Canada T2N4N1 Edited* by David Pilbeam, Harvard University, Cambridge, MA, and approved December 29, 2009 (received for review October 14, 2009) The long legs and short arms of humans are distinctive for a fore- and hindlimb modules was reduced, leading to a higher primate, the result of selection acting in opposite directions on degree of variational independence of limb size and shape. Our each limb at different points in our evolutionary history. This model predicts that selection for independent function of the mosaic pattern challenges our understanding of the relationship limbs should lead to alterations to limb covariational structure. of development and evolvability because limbs are serially homol- Specifically, humans and apes would be predicted to exhibit ogous and genetic correlations should act as a significant con- decreased phenotypic correlations between fore- and hindlimb straint on their independent evolution. Here we test a compared to quadrupeds. If decreases in integration between developmental model of limb covariation in anthropoid primates fore- and hindlimb are a product of selection for more inde- and demonstrate that both humans and apes exhibit significantly pendent function of limbs, then the timing of any change in reduced integration between limbs when compared to quadrupe- integration has implications for the reconstruction of ancestral dal monkeys.
    [Show full text]
  • Largemouth Bass, Smallmouth Bass, and Spotted Bass Management and Fishing in Pennsylvania
    Largemouth Bass, Smallmouth Bass, and Spotted Bass Management and Fishing in Pennsylvania Prepared by R. Lorantas, D. Kristine and C. Hobbs PFBC Warmwater Unit 2005 (updated 2013; R. Lorantas) Goal: Maintain or enhance largemouth, smallmouth, and spotted bass sport fishing through harvest management of naturally sustained bass populations and through habitat preservation and enhancement. Judiciously stock largemouth and smallmouth bass in compatible new and reclaimed habitats. Largemouth bass and smallmouth bass occur throughout Pennsylvania and were originally indigenous to the Ohio River and Lake Erie Drainage. Spotted bass, Pennsylvania’s most rare black bass, occurs only in the Ohio River drainage. The Ohio drainage includes the Ohio River, Allegheny River and Monongahela River drainages. Largemouth bass typically predominate in reservoirs and lakes and occur at lower densities in slow moving rivers and streams within these drainages. Smallmouth bass are typically abundant in rivers, warmwater streams and medium to large size lakes and reservoirs in these drainages. Spotted bass are most abundant within a 20 mile radius of the confluence of the Ohio, Allegheny and Monongahela Rivers. In the Lake Erie drainage largemouth bass are largely confined to Presque Isle Bay, however smallmouth bass are abundant in Lake Erie as well as Presque Isle Bay. Smallmouth bass and largemouth bass stocking by the Pennsylvania Fish and Boat Commission and other agencies over a century ago into the Delaware, Susquehanna, and Potomac River Drainages lead to colonization of waters within these drainages, and both species are now self­sustaining in these drainages. Most natural warm­ water lakes and man­made reservoirs in Pennsylvania contain self– sustaining largemouth and smallmouth bass populations.
    [Show full text]
  • Evaluating the Ecology of Spinosaurus: Shoreline Generalist Or Aquatic Pursuit Specialist?
    Palaeontologia Electronica palaeo-electronica.org Evaluating the ecology of Spinosaurus: Shoreline generalist or aquatic pursuit specialist? David W.E. Hone and Thomas R. Holtz, Jr. ABSTRACT The giant theropod Spinosaurus was an unusual animal and highly derived in many ways, and interpretations of its ecology remain controversial. Recent papers have added considerable knowledge of the anatomy of the genus with the discovery of a new and much more complete specimen, but this has also brought new and dramatic interpretations of its ecology as a highly specialised semi-aquatic animal that actively pursued aquatic prey. Here we assess the arguments about the functional morphology of this animal and the available data on its ecology and possible habits in the light of these new finds. We conclude that based on the available data, the degree of adapta- tions for aquatic life are questionable, other interpretations for the tail fin and other fea- tures are supported (e.g., socio-sexual signalling), and the pursuit predation hypothesis for Spinosaurus as a “highly specialized aquatic predator” is not supported. In contrast, a ‘wading’ model for an animal that predominantly fished from shorelines or within shallow waters is not contradicted by any line of evidence and is well supported. Spinosaurus almost certainly fed primarily from the water and may have swum, but there is no evidence that it was a specialised aquatic pursuit predator. David W.E. Hone. Queen Mary University of London, Mile End Road, London, E1 4NS, UK. [email protected] Thomas R. Holtz, Jr. Department of Geology, University of Maryland, College Park, Maryland 20742 USA and Department of Paleobiology, National Museum of Natural History, Washington, DC 20560 USA.
    [Show full text]
  • 2021 Fish Suppliers
    2021 Fish Suppliers A.B. Jones Fish Hatchery Largemouth bass, hybrid bluegill, bluegill, black crappie, triploid grass carp, Nancy Jones gambusia – mosquito fish, channel catfish, bullfrog tadpoles, shiners 1057 Hwy 26 Williamsburg, KY 40769 (606) 549-2669 ATAC, LLC Pond Management Specialist Fathead minnows, golden shiner, goldfish, largemouth bass, smallmouth bass, Rick Rogers hybrid bluegill, bluegill, redear sunfish, walleye, channel catfish, rainbow trout, PO Box 1223 black crappie, triploid grass carp, common carp, hybrid striped bass, koi, Lebanon, OH 45036 shubunkin goldfish, bullfrog tadpoles, and paddlefish (513) 932-6529 Anglers Bait-n-Tackle LLC Fathead minnows, rosey red minnows, bluegill, hybrid bluegill, goldfish and Kaleb Rodebaugh golden shiners 747 North Arnold Ave Prestonsburg, KY 606-886-1335 Andry’s Fish Farm Bluegill, hybrid bluegill, largemouth bass, koi, channel catfish, white catfish, Lyle Andry redear sunfish, black crappie, tilapia – human consumption only, triploid grass 10923 E. Conservation Club Road carp, fathead minnows and golden shiners Birdseye, IN 47513 (812) 389-2448 Arkansas Pondstockers, Inc Channel catfish, bluegill, hybrid bluegill, redear sunfish, largemouth bass, Michael Denton black crappie, fathead minnows, and triploid grass carp PO Box 357 Harrisbug, AR 75432 (870) 578-9773 Aquatic Control, Inc. Largemouth bass, bluegill, channel catfish, triploid grass carp, fathead Clinton Charlton minnows, redear sunfish, golden shiner, rainbow trout, and hybrid striped bass 505 Assembly Drive, STE 108
    [Show full text]
  • Vestibular Evidence for the Evolution of Aquatic Behaviour in Early
    View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Publications of the IAS Fellows letters to nature .............................................................. cetacean evolution, leading to full independence from life on land. Vestibular evidence for the Early cetacean evolution, marked by the emergence of obligate evolution of aquatic behaviour aquatic behaviour, represents one of the major morphological shifts in the radiation of mammals. Modifications to the postcranial in early cetaceans skeleton during this process are increasingly well-documented3–9. Pakicetids, early Eocene basal cetaceans, were terrestrial quadrupeds 9 F. Spoor*, S. Bajpai†, S. T. Hussain‡, K. Kumar§ & J. G. M. Thewissenk with a long neck and cursorial limb morphology . By the late middle Eocene, obligate aquatic dorudontids approached modern ceta- * Department of Anatomy & Developmental Biology, University College London, ceans in body form, having a tail fluke, a strongly shortened neck, Rockefeller Building, University Street, London WC1E 6JJ, UK and near-absent hindlimbs10. Taxa which represent bridging nodes † Department of Earth Sciences, Indian Institute of Technology, Roorkee 247 667, on the cladogram show intermediate morphologies, which have India been inferred to correspond with otter-like swimming combined ‡ Department of Anatomy, College of Medicine, Howard University, with varying degrees of terrestrial capability4–8,11. Our knowledge of Washington DC 20059, USA § Wadia Institute of Himalayan Geology, Dehradun 248 001, India the behavioural changes that crucially must have driven the post- k Department of Anatomy, Northeastern Ohio Universities College of Medicine, cranial adaptations is based on functional analysis of the affected Rootstown, Ohio 44272, USA morphology itself. This approach is marred by the difficulty of ............................................................................................................................................................................
    [Show full text]
  • Fins, Limbs, and Tails: Outgrowths and Axial Patterning in Vertebrate Evolution Michael I
    Review articles Fins, limbs, and tails: outgrowths and axial patterning in vertebrate evolution Michael I. Coates1* and Martin J. Cohn2 Summary Current phylogenies show that paired fins and limbs are unique to jawed verte- brates and their immediate ancestry. Such fins evolved first as a single pair extending from an anterior location, and later stabilized as two pairs at pectoral and pelvic levels. Fin number, identity, and position are therefore key issues in vertebrate developmental evolution. Localization of the AP levels at which develop- mental signals initiate outgrowth from the body wall may be determined by Hox gene expression patterns along the lateral plate mesoderm. This regionalization appears to be regulated independently of that in the paraxial mesoderm and axial skeleton. When combined with current hypotheses of Hox gene phylogenetic and functional diversity, these data suggest a new model of fin/limb developmental evolution. This coordinates body wall regions of outgrowth with primitive bound- aries established in the gut, as well as the fundamental nonequivalence of pectoral and pelvic structures. BioEssays 20:371–381, 1998. ௠ 1998 John Wiley & Sons, Inc. Introduction over and again to exemplify fundamental concepts in biological Vertebrate appendages include an amazing diversity of form, theory. The striking uniformity of teleost pectoral fin skeletons from the huge wing-like fins of manta rays or the stumpy limbs of illustrated Geoffroy Saint-Hilair’s discussion of ‘‘special analo- frogfishes, to ichthyosaur paddles, the extraordinary fingers of gies,’’1 while tetrapod limbs exemplified Owen’s2 related concept aye-ayes, and the fin-like wings of penguins. The functional of ‘‘homology’’; Darwin3 then employed precisely the same ex- diversity of these appendages is similarly vast and, in addition to ample as evidence of evolutionary descent from common ances- various modes of locomotion, fins and limbs are also used for try.
    [Show full text]