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Lungs and Mesopneumonia of Scincomorph Lizards (Reptilia: Squamata) Wilfried Kleina,Ã, Christian Reutera, Wolfgang Bo¨ Hmeb, Steven F

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Lungs and Mesopneumonia of Scincomorph Lizards (Reptilia: Squamata) Wilfried Kleina,Ã, Christian Reutera, Wolfgang Bo¨ Hmeb, Steven F ARTICLE IN PRESS Organisms, Diversity & Evolution 5 (2005) 47–57 www.elsevier.de/ode Lungs and mesopneumonia of scincomorph lizards (Reptilia: Squamata) Wilfried Kleina,Ã, Christian Reutera, Wolfgang Bo¨ hmeb, Steven F. Perrya aAbt. Morphologie und Systematik, Institut fu¨r Zoologie, Universita¨t Bonn, Poppelsdorfer Schloss, 53115 Bonn, Germany bSektion Herpetologie, Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany Received 25 April 2004; accepted 21 June 2004 Abstract Mesopneumonia of 28 and lungs of 44 species of scincomorph lizards are described, representing mesopneumonia in six of the seven scincomorph families and lungs of all seven families of this taxon. Except for gymnophthalmids and scincids, a family typical organization of mesopneumonia occurs. In cordylids, gerrhosaurids, xantusiids and lacertids the complete right ventral mesopneumonium (VMp) inserts cranially on the posterior vena cava and caudally on the dorsal surface of the right liver lobe. The left VMp is attached to the ventrolateral body wall in cordylids and lacertids; in gerrhosaurids the left VMp is short and inserts on the pericardium and the posterior vena cava; in xantusiids the left VMp is reduced and the lungs are fused cranially with the pericardium. In scincids the VMp of both sides vary in length and insert on the ventral mesentery, or may be lacking completely. The visceral topology of the gymnophthalmids Calyptommatus and Notobachia differs from the general scincomorph pattern, with the liver and stomach elongated and situated on the right and left side of the body cavity, respectively. The left and the right VMp extend over the entire length of the lungs and insert on the ventral mesentery. All lungs examined in the study are single-chambered and show no major structural variability of the inner surfaces, except in lacertids, some gerrhosaurids, Cordylus, and two genera of gymnophthalmids (Echinosaura and Neusticurus). In these groups, rows of dorsomedial niches are present. r 2005 Elsevier GmbH. All rights reserved. Keywords: Scincomorpha; Lizards; Morphology; Lungs; Mesopneumonia; Viscera See also Electronic Supplement at: http://www.senckenberg.de/odes/05-06.htm Introduction and Gymnophthalmidae (Fig. 1; Estes et al. 1988). The number of species exceeds 1900 for all scincomorph Scincomorpha comprises seven families forming two lizards, with approximately 1227 scincids (Greer 2001), clades. One group (Scincoidea) comprises Scincidae, 54 cordylids (Uetz 2004), 32 gerrhosaurids (Uetz 2004), Cordylidae and Gerrhosauridae, whereas the other 26 xantusiids (Uetz 2004), 264 lacertids (Uetz 2000), (Lacertoidea) contains Xantusiidae, Lacertidae, Teiidae 117 teiids (Uetz 2000), and 182 gymnophthalmids (Uetz 2000). The lungs of 13 and the mesopneumonia of 24 Ã Corresponding author. Departamento de Zoologia, Instituto de scincomorph species are described in detail, representing Biologia, Universidade Federal da Bahia, Rua Bara˜ o de Geremoabo, all or six of the seven scincomorph families, res- 147, Campus de Ondina, CEP: 40170-290, Salvador, Bahia, Brazil. Tel.: +55 71 2636560; fax: +55 71 2456909. pectively (see Part 1 of the accompanying Organisms E-mail address: [email protected] (W. Klein). Diversity and Evolution Electronic Supplement at 1439-6092/$ - see front matter r 2005 Elsevier GmbH. All rights reserved. doi:10.1016/j.ode.2004.06.004 ARTICLE IN PRESS 48 W. Klein et al. / Organisms, Diversity & Evolution 5 (2005) 47–57 water, 6–8 h; (2) periodic acid (0.5%), 60 min; (3) Schiff’s reagent, 10 min; (4) tap water, 5 min; (5) acid ethanol (70%, 1% HCl conc.), 16 h; (6) toluidine blue (0.25%, pHo2), at least 4 h; (7) ethanol (70%), several repetitions to wash out superfluous toluidine blue; (8) ethanol (50%), 5 min; (9) clearing and storage of lungs in glycerol. The following data for the respiratory system (glottis, trachea and lungs) were collected [units of measurement in square brackets]: length of respiratory system measured from cranial tip of glottis to most caudal Fig. 1. Phylogenetic relationships of scincomorph lizards end of lungs (LRS [mm]), length of lungs (LLu [mm]), following Estes et al. (1988). length of left lung (Lle [mm]), length of right lung (Lri [mm]), length of trachea (LTr [mm]), length of extra- pulmonary bronchus (LBr [mm]), length of prehilar region (LPh [mm]), number of niches (NNi), number http://www.senckenberg.de/odes/05-06.htm). In most of tracheal cartilages (NTC), parenchymal type cases qualitative data for only one or two species per (ed ¼ ediculae, fav ¼ faveoli) and parenchymal distri- family is available (e.g. Milani 1894; Broman 1904; Becker bution (het ¼ heterogeneous, hom ¼ homogeneous). 1993). However, within the Teiioidea the mesopneumonia LRS, LLu and LTr were standardized to SVL, LTr and (Klein et al. 2000), and within the Varanoidea the lung LBr were standardized to LRS, LPh was standardized to structure (Becker et al. 1989), comprise a useful character LLu, and the quotients of lung length over trachea length only at the family level. Therefore, description of a small (LLu/LTr), and length of left lung over length of right number of species per family represents a meaningful lung (Lle/Lri) were calculated. contribution. The aim of the present paper is to describe the mesopneumonia and provide quantitative morpholo- gical data for the lungs of representatives of all families of Results scincomorph lizards, and thereby to develop some hypotheses about the functional and systematic value of the mesopneumonia–lung complex. Scincomorpha General visceral topology Materials and methods The heart lies in its pericardial cavity at the level of the shoulder girdle, connected caudally via the vena cava to the liver. The latter is composed of two lobes, Specimens from the Alexander Koenig Zoological divided in the mediosagittal plane. The right lobe is Research Institute and Museum (ZFMK) in Bonn, Germany, and from the collection of P.L.B. Rocha larger than the left one and extends dorso-caudally, contacting the meso of the right gonad with its caudal (Universidade Federal da Bahia, Brazil) were used. end. The gall bladder is embedded in the ventro-caudal Snout-vent length (SVL), total length (TL), and body part of the right liver lobe and the stomach lies on the mass (M ) were recorded as reference data (see Electr. B left side of the abdominal cavity. The lungs lie dorso- Suppl. 05-06, Pt. 2). caudally to the heart and dorsally to the cranial parts of The visceral topology and location of mesos in each liver and stomach. No intrapulmonary bronchi are group are presented, followed by a description of the present and the extra-pulmonary bronchi, when present, lungs. Definitions of the mesos and representation of the results follow Klein et al. (2000). Briefly, a standard are short. Intestine, gonads and fat bodies, whose presence and size vary according to the nutritional schematic diagram, representing the parietal and visc- status of a specimen, fill the caudal part of the body eral surfaces within a body cavity, is used to document cavity. the origin and insertion of the investigated mesos (DM ¼ dorsal mesentery, VM ¼ ventral mesentery; DMp ¼ dorsal mesopneumonium, VMp ¼ ventral me- Scincidae sopneumonium). The part of a meso attached to the organ is defined as its origin; the distal part as insertion. Visceral topology and mesos To visualize the lungs, a combined toluidine blue-PAS The viscera of skinks show no deviation from the (Periodic-Acid–Schiff reaction) en-bloc staining method general scincomorph pattern. DMp originate on the was used. This method, which selectively stains tracheal dorsal mid-line of the lungs and insert on the border cartilage and lung parenchyma, is as follows: (1) tap alimentary canal-DM. The VMp vary in their development ARTICLE IN PRESS W. Klein et al. / Organisms, Diversity & Evolution 5 (2005) 47–57 49 Fig. 2. (a) Attachments of mesopneumonia in Chalcides, Tiliqua and Tribolonotus; left: transverse sections through body cavity (ventral side up), right: parietal surfaces in body cavity; DM ¼ dorsal mesentery, DMp ¼ dorsal mesopneumonium (dashed line), li ¼ liver, lu ¼ lung, vcp ¼ vena cava posterior, VM ¼ ventral mesentery, VMp ¼ ventral mesopneumonium (solid line). (b) Attachments of mesopneumonia in Scincus scincus. (c) Respiratory system in S. scincus, dorsal view, left lung open to show internal structure, right lung showing major blood vessels on outer surface; scale bar ¼ 1 cm. on both sides. A cranial fusion of lungs with pericar- bronchi 2.3% LRS (min. 1.6% in Mabuya, max. 3.2% in dium occurs in Chalcides, Tiliqua and Tribolonotus Scincus), the prehilus 6.2% LLu (min. 2.5% in Novoeu- (Fig. 2a). Novoeumeces and Mabuya show a complete meces, max. 12.3% in Chalcides), and the lungs 22.8% right VMp originating in the ventral mid-line of the lung SVL (min. 13.2% in Chalcides, max. 32.3% in Tribolo- and inserting at the border esophagus-VM. The left notus). The lungs are of equal length, with Lle/Lri 1.03 VMp of Novoeumeces extends from the cranial 1/4 of the (min. 0.92 in Mabuya, max. 1.12 in Scincus). The NTC is lung and inserts at the base of the esophageal VM. In 51.3 (min. 33.3 in Tribolonotus, max. 62.7 in Mabuya), Mabuya, on the other hand, the left VMp is lacking and and LLu/LTr is 1.22 (min. 0.9 in Chalcides, max. 1.6 in the left lung is fused cranially with the pericardium. The Tribolonotus). The lungs are long and slender, show a right VMp of Scincus originates in the cranial 3/4 of the slightly heterogeneous parenchymal distribution with lung and inserts on the VM. The left VMp, by contrast, faveoli in the cranial part of the lung and ediculae in the originates only on the cranial 1/4 of the lung, also most caudal part. No niches are present (Fig. 2c). inserting on the VM (Fig. 2b). Respiratory system Cordylidae For results per species see Electr.
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