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Anomalies in the Vertebral Column and Ilio-Sacral Articulation of Some Anuran Amphibians

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Anomalies in the Vertebral Column and Ilio-Sacral Articulation of Some Anuran Amphibians SALAMANDRA 57(1): 53–64 Anomalies in the anuran vertebral column and ilio-sacral articulation SALAMANDRA 15 February 2021 ISSN 0036–3375 German Journal of Herpetology Anomalies in the vertebral column and ilio-sacral articulation of some anuran amphibians Alexander Haas1, Sophie Schwippert1, Sebastian Büsse2, Thomas Kleinteich3, André Beerlink4, Jörg U. Hammel5, Stephanie Köhnk1, Stanislav Gorb2 & Karolin Engelkes1 1) Centrum für Naturkunde, Universität Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany 2) Functional Morphology and Biomechanics, Institute of Zoology, University of Kiel, Am Botanischen Garten 1–9, 24118 Kiel, Germany 3) TPW Prüfzentrum GmbH, Xantener Str. 6, 41460 Neuss, Germany 4) YXLON International GmbH, Essener Bogen 15, 22419 Hamburg 5) Institute of Materials Research, Helmholtz-Zentrum Geesthacht, Max-Planck-Str. 1, 21502 Geesthacht, Germany Corresponding author: Alexander Haas, e-mail: [email protected] Manuscript received: 17 August 2020 Accepted: 20 November 2020 by Alexander Kupfer Abstract. We present accounts of vertebral anomalies in 17 individuals representing 13 species of anuran amphibians. These cases were detected while perusing a larger survey on the skeleton of frogs, for which µCT scans of a broad range of species were collected and evaluated. Our data and reports from the literature suggest that malformations, asymmetries, and irregularities, if present, appear to be particularly prevalent in the posterior region of the axial skeleton in frogs. Anomalies at the trunk-tail boundary, i.e., at the sacrum and neighbouring segments, were relatively common. Malfor- mations at the trunk-tail boundary often include sacralization of pre- and postsacral elements with asymmetrically or symmetrically developed diapophyses, fusion with the posteriormost presacral vertebra, occurrence of postsacral verte- brae, unusual transverse processes at the proximal end of the urostyle, formation of additional zygapophyses, or fusion of elements that normally articulate. Vertebral fusion in the anterior vertebral column (Presacral Vertebrae I+II) has been reported both in evolutionary context and in cases of individual developmental anomalies. Malformations in the middle section of the vertebral column, such as the case of Epidalea calamita reported herein, are rare. Key words. Amphibia, Anura, axial skeleton, pelvis, sacrum, malformation. Introduction es the articulation between the vertebral column and the pelvis. The sacral vertebra is pivotal for the function of the The anuran body plan is unique among vertebrates mak- sacro-urostylic complex (Emerson 1982). The tail vertebral ing members of the monophylum Anura easily recogniz- column is highly reduced in all extant species; only vestiges able: apart from losing the tail, the truncation of the ver- of the tail vertebrae contribute to the formation of the uro- tebral column, the reduction in the number of vertebrae, style that is otherwise an evolutionary novelty (Ročkova & the elongation of the ilia, expanded sacral diapophyses with Roček 2005, Pugener & Maglia 2009a). ventral articulation to the ilium, and the formation of the The anatomical details and the shape of the ilio-sacral urostyle are the most obvious synapomorphies in early frog joint vary among the frog lineages and are strongly cor- evolution (Shubin & Jenkins 1995, Pugener & Maglia related with a species’ specialization in locomotor mode, 2009a). Frogs are the tetrapod group with the fewest verte- such as jumping versus swimming or burrowing, as well brae. Species of the Ascaphidae and Leio pelmatidae possess as habitat choice (arboreal, fossorial, aquatic, terrestrial) nine presacral vertebrae, all other anurans have a vertebral (Emerson 1979, Jorgensen & Reilly 2013). For example, count of eight or less (Duellman & Trueb 1994). There the very wide sacral diapophyses of the sacral vertebra in is only moderate differentiation into functional groups in pipids allow for a forward sliding of the pelvis (Videler the presacral vertebral column, for example, in the size of & Jorna 1985, Cundall et al. 2017), whereas the almost vertebral transverse processes at the level of the suprascap- cylindrical diapophyses in many ranid frogs provide the ula. Presacral Vertebra I, or Presacral I for short, is the only necessary articular freedom for dorsoventral extension at neck vertebra and stands out by typically lacking transverse the initial jumping phase (Emerson 1979, 1982, Jenkins & processes (Gaupp 1896). A single sacral vertebra establish- Shubin 1998). © 2021 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT), Mannheim, Germany Open access at http://www.salamandra-journal.com 53 Alexander Haas et al. Considering the crucial functional role of the verte- Sensing & Inspection Technologies GmbH Phoenix Nano- bral column, ilio-sacral articulation, and sacro-urostylic tom S or M, or a GE Sensing & Inspection Technologies complex, one would expect that mutational malforma- GmbH Phoenix v|tome|x L 450. Depending on the size of tions of the skeleton were strongly selected against in evo- the specimen, the scanning parameters were set appropri- lution and should be rare in nature (Zamora-Camacho ately (Supplementary document S1). Volumes were recon- & Aragón 2019). Congenital skeletal anomalies, however, structed from X-ray projections using the software deliv- can have many different causes other than genetic muta- ered with the respective scanner. The resulting voxel size tions. Particularly in organisms that complete a substan- ranged from 11.87–106.01 µm, depending on the specimen tial part of their skeletal development during larval life and size and CT system used. metamorphosis, when they can be exposed to teratogens in Amira 6.0.1. software (Thermo Fisher Scientific) was the environment. Recently, Ouellet (2000) and Henle & used to visualize and inspect µCT volumes. For each avail- Dubois (2017) have assessed amphibian anomalies in gen- able µCT image stack an Isosurface of the calcified tissues eral. These hitherto most extensive compilations and eval- (i.e., bones and calcified cartilage) was generated for in- uations of published reports cover anomalies in colour pat- spection. If the specimen showed anomalies in the verte- tern and morphology, and review hypotheses about causes bral column or the ilio-sacral articulation, relevant skele- of such pathologies. Specific skeletal abnormalities, how- tal structures were segmented with Amira’s Segmentation ever, have been covered only marginally apart from exter- Editor (Brush and Magic Wand tools). As this study fo- nal features of general limb or body deformations. Henle cuses on the vertebral column and sacral structures, only & Dubois (2017) provided a glossary, specific cases, and affected skeletal elements were accurately separated dur- general methodological recommendations. ing segmentation; cruder segmentation methods were ap- In a different project on skeletal features of the shoulder plied in other areas of the skeleton. Because cartilage does girdle (Engelkes et al. 2020), we had access to and exam- not visualize reliably in µCT datasets of plain specimens ined micro-computed tomography (µCT) scans of a broad and because it does not contribute additional aspects to taxonomic sample of anuran amphibians. During the ex- the phenomena treated herein, cartilage tissue was gener- amination we recognized several individuals with verte- ally neglected for this study; only calcified cartilage was oc- bral and sacral malformations or anomalies compared to casionally segmented and visualized in skeletal parts that conspecifics or congeners. Various categories of vertebral were unaffected by malformations if its grey values fell anomalies have been described before (Madej 1965). We within the range of bone grey values. Segmented skeletons classify vertebral malformations herein, first, as any asym- were exported with the mulitExport macro (Engelkes et metry that was so substantial that it could be detected visu- al. 2018) as polymesh surfaces (.obj format) and, if neces- ally even without further measurement, such as much in- sary, further processed in MeshLab 1.3.3 (Cignoni et al. flated or present processes on only one side. In specimens 2008); specifically, Quadric Edge Collapse Decimation and that could be considered normal, in contrast, asymmetries Taubin Smooth procedures were applied in MeshLab to re- were not perceivable. A second category of anomalies does duce the number of polygons and to smooth the surfaces not involve asymmetry but the fusion of consecutive ver- while still preserving the original surface as well as pos- tebral structures. Third, supernumerary elements can be sible. Next, the surfaces were imported into various ver- present. Sometimes a combination of these major types of sions of Modo (7–13; The Foundry Visionmongers Ltd.) for malformations can occur in the same specimen. In the fol- subtle further smoothing (if necessary), topology hole fill- lowing we describe the osteological malformations of the ing, assigning colours (malformations red; corresponding vertebral column and ilio-sacral articulation present in structures without indication of malformation green, all specimens examined. Providing a comprehensive compila- other skeletal parts grey), and final rendering (Dome Light tion on the topic is beyond the scope of this work, but by and Directional Light). Colour plates were arranged using reporting the phenomenology of some new cases of axial Graphic 3.1 (Picta Inc.)
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