S stereochemistry at the point of attachment of the 3,4-dimethoxyphenyl substituent, the 1R stereochemistry of the 2-methoxyphenol attachment point was omitted. In the structure of quebrachitol the stereochemistry that distinguishes this particular compound from other O-methylinositol isomers was not indicated. Finally, a double bond was missing in the structure of oxoputerine. The corrected The authors wish to inform readers that there are several minor errors and omissions in the chemical structures shown in Mitrephora species mentioned in our paper, including the title species M. vulpina, has changed, and the names M. vulpina C.E.C. Fisch, M. zippeliana and M. diversifolia are no longer used, while the species M. thorelii Pierre is now M. tomentosa Hook f. & Thomson. Readers are advised to consult a recent monograph on this genus [Finally, we have been alerted by Prof. Christophe Wiart (University of Nottingham) that the taxonomy of several of the is genus  for the


AbstractAsclepiassyriaca L. in Lithuania revealed the occurrence of a new alien plant species, the North American native Asclepiasspeciosa Torr. , in southern parts of Lithuania – the first report of the latter species in Europe. Interestingly, a thorough analysis of herbarium specimens revealed that A.speciosa had first been collected in Lithuania in 1962, but the specimen was misidentified at the time as A.syriaca. The newly discovered population of A.speciosa occupies mesic grasslands, tall-herb fringe communities and arable field habitats. Sexual reproduction of this species was not recorded; it spreads locally by means of vegetative reproduction. We present here an exhaustive analysis of morphological characteristics and differences between A.speciosa and A.syriaca and other species of the genus, as well as a key for identification of alien Asclepias species in Europe. We predict that the effect of A.speciosa on native habitats and communities, and its economic impact, are comparable to those of the highly invasive A.syriaca. Although A.speciosa currently occurs very rarely as an alien species in Europe, its existence in other regions of Europe is highly probable.Studies on populations of  Asclepias L. s. str.  includes about 120 species native to the Western Hemisphere. Most of the species are distributed in North America and the Caribbean, ten species occur in South America  Schltr., have also been reported as alien naturalized species in several countries of southern Europe (Gomphocarpus R.Br. (Gomphocarpusfruticosus (L.) W.T.Aiton and Gomphocarpusphysocarpus E.Mey., respectively) .In Europe, three alien species of the genus riaca L. . Anothern Europe ; howeverctively) . About an Europe . The mos.syriaca , whereasf Europe . BecauseA.syriaca on native plant communities in Lithuania, because available information on this species was only fragmentary. Over several decades, A.syriaca had been reported to occur at several localities in southern Lithuania, and it has been recognized as a naturalized species . Research on morphological characteristics, flowering, and habitats was performed on 26–28 June and 20 September 2018. For comparison of morphological features, we selected a population of A.syriaca located in a similar habitat in the village of Meškučiai, Kaišiadorys district . In each population, we studied characters of 30 flowering shoots.The population of Plants for the study were selected randomly from all over the colony, with a distance of at least 2 m between sampled plants. Stems were cut at ground level with garden shears. Stem height was measured from the soil level to the apex with a precision of 1 cm, using a measuring tape. The number of leaf pairs on the stem was counted, including wilted and fallen leaves. In cases where some of the lower leaves were fallen, their number was determined by leaf scars on the stem. Leaf measurements with a precision of 0.1 cm were taken from the leaf that was situated closest to the middle of the stem. The length and width of the leaf blade, length of the petiole, and diameter of the lowermost inflorescence were measured using a ruler. The length of the leaf blade was measured from the leaf base at the junction with the petiole to its tip, whereas the width of the leaf was measured at its widest point. The number of developed inflorescences on the stem was counted. The diameters of the inflorescences were measured at their middle part and the number of flowers was counted.Asclepias species stored at the Herbarium of the Institute of Botany of the Nature Research Centre (BILAS), Vilnius, Lithuania. Specimens collected during this research were also deposited at the same herbarium.We studied the herbarium specimens of A.speciosa and A.syriaca was tested by applying a 2-sample t-test. All calculations were performed using PAST 3.20 ; the specimen was identified by V. Galinis as A.syriaca . This record was also mentioned by BILAS. Based on the information provided by A.speciosa was erroneously considered as the first record of A.syriaca in Lithuania ; however, it was also erroneously identified as A.syriaca . Just a few days later, A.speciosa was collected by Ž. Sinkevičius . More than a decade later, in 2004, a specimen of A.speciosa was collected by V. Rašomavičius  and also identified as A.syriaca. Therefore, A.speciosa has been present in southern Lithuania in the vicinity of the Žuvintas Biosphere Reserve for at least 56 years and at its recently identified locality for at least 26 years.A new locality for A.speciosa  in the mire of Žuvintas, although the exact size of the population was not given. The label of the herbarium specimen of A.speciosa collected by M. Lapelė in the vicinity of the village of Liepakojai indicated that plants were abundant, but the size of the colony was not specified. The herbarium label of the specimen collected in 2004 by V. Rašomavičius noted that this species formed a colony; however, once again, the size of the stand and density of the colony were not recorded. During the present study in 2018, the area occupied by A.speciosa in the vicinity of the village of Liepakojai was measured. A dense stand of A.speciosa in a mesic meadow and at the edge of a woodland .Both species also differ in the number of flowers in the inflorescence. Although the diameter of the inflorescence is almost the same in both species Table , those oA.speciosa and A.syriaca. The leaf blade of the middle cauline leaves of A.speciosa is broadly ovate or elliptical, widest at its slightly cordate base, whereas the leaf blade of A.syriaca is ovate or elliptical, with a rounded or slightly cuneate base, and is widest near its middle . The hoods of A.speciosa are much longer (10–13 mm) than in both other species and differ in shape, having prolonged acute apices rather than short rounded apices.A.speciosa in Lithuania was recorded in 1962 in the district of Alytus. Considering information provided on herbarium labels, this species has been recorded in two separate locations in what is now the Žuvintas Biosphere Reserve and in its vicinity, on the outskirts of the village of Liepakojai. Therefore, the currently studied colony of A.speciosa has existed for at least 26 years and this species can be considered as naturalized in Lithuania. Although the time of its introduction into Lithuania is unknown, we suppose that A.speciosa might have been introduced at the end of the 19th century or the beginning of the 20th century as an ornamental plant at the nearby Riečiai Manor Park  inflorescences (In its native habitat of Washington State (USA), individual reproductive shoots of escences . In the escences . Such diA.speciosa grows in a broad range of moisture conditions. Usually it is found in moderately wet but well-drained soil, though quite frequently it is found in riparian sites and sub-irrigated or occasionally flooded habitats. Occasionally, populations of this species can also be found in very dry sites  is at approximately 50°N (A.speciosa from southern Manitoba west to British Columbia and south to Minnesota to northwestern Texas and California (A.syriaca (A.speciosa in Lithuania, it can likely invade a broad range of grassland habitats, some types of wetland, many anthropogenic habitats, and become a weed of arable lands, particularly in the Pannonian and Steppic biogeographical regions of Europe. The effect of this species on habitats, their biodiversity and economic impact can be considered comparable to that of A.syriaca. A major obstacle to the spreading of this species is the apparent absence of sexual reproduction in the studied populations in Europe. However, further observations on the fruit setting and possible sexual reproduction of the plant in Europe are required.The northern limit of the native distribution of ely 50°N . Therefolifornia , one canlifornia . Potenti.syriaca . Based oA.speciosa is a very rare alien species and Lithuania is the first documented country of occurrence in Europe. However, the possibility that this species already occurs in other regions of Europe cannot be excluded. It is possible that this species has been overlooked due to resemblance with the quite variable and similar A.syriaca, particularly when plants are examined at the vegetative stage. Therefore, botanists and ecologists should pay particular attention to species of the genus Asclepias in nature, as well as critically review herbarium specimens. It seems unlikely that A.speciosa has only been introduced into Lithuania.Currently,


The Upper Mississippian Bluefield Formation of the Mauch Chunk Group in southeastern West Virginia is known for its preservation of a variety of invertebrate taxa and early tetrapod trackways, but no lower actinopterygian remains have been formally described from these Carboniferous rocks. Here, the first lower actinopterygian fish is described from the Bluefield Formation of West Virginia. This fish is represented by a nearly complete articulated specimen with a three-dimensional snout and an unobstructed view of the gular and branchiostegal region. This new taxon is defined by a unique set of characters, which include features of the snout, circumorbital series, cheek, and operculo-gular region. These features make this fish different and distinct from previously described Carboniferous fishes. Some of the morphological features of note include the presence of a distinct lacrimal, premaxillary, ventral rostral and dorsal rostral bones, a narrow infraorbital ventral to the orbit, and a large crescent shaped infraorbital that contacts a single dermosphenotic. There is an anteriorly inclined hatchet-shaped preoperculum and six small suborbital bones anterior to the expanded region of this bone that filling the space between the preoperculum, dermosphenotic, and infraorbital. Posterior to the preoperculum, there is a single wedge-shaped dermohyal and a series of three rectangular anteopercular bones. The anteopercular bones extend halfway down the anterior border of the rectangular operculum. A median gular, two pairs of lateral gulars, and at least eight branchiostegal rays are present. The heterocercal caudal fin is deeply cleft and inequilobate. The scales have pectinated posterior margins and bear diagonal ridges of ganoine. The description of this new taxon represents the first actinopterygian and the first vertebrate body fossil described from the Bluefield Formation and the second actinopterygian taxon described from the Mauch Chunk Group in West Virginia. The body plan of modern fishes is thought to have originated within a grouping of lower actinopterygian fishes collectively referred to as palaeoniscoids . Though The Upper Mississippian Bluefield Formation is the basal-most formation in the Mauch Chunk Group in the Appalachian Basin . In soutHylopus hamesi have been described from the Bluefield Formation , Lawrence, Kansas, USA.The specimen was examined by stereomicroscopy. To aid in visualization of features, the specimen was viewed under 70% ethanol or shaded with magnesium oxide. Illustrations were prepared using a camera lucida and digital illustrations were constructed using Adobe Photoshop and Illustrator programs. Photographs were taken with a Canon XSi Digital SLR camera with a macro lens. When possible, morphometric measurements and meristic counts were taken following the scheme of http://zoobank.org/. The LSID for this publication is: urn:lsid:zoobank.org:pub:0F2C9D33-9584-4739-B914-5C4C243E0B1F. The online version of this work is archived and available from the following digital repositories: PeerJ, PubMed Central and CLOCKSS.The electronic version of this article in Portable Document Format (PDF) will represent a published work according to the International Commission on Zoological Nomenclature (ICZN), and hence the new names contained in the electronic version are effectively published under that Code from the electronic edition alone. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix Lower actinopterygian skull bones have been identified differently by various researchers See  is accepDiagnosis. As for the type and only speciesType and only Species.Bluefieldius mercerensis n. gen. n. spEtymology. After the Bluefield Formation where the specimen was recovered from.Etymology.mercerensis in reference to Mercer County, West Virginia where the specimen was recovered.Diagnosis. Based on the unique combination of the following characters: Absence of complex bones in the snout and the presence of distinct and separate lacrimal, premaxillary, and dorsal and ventral rostral bones; narrow rectangular infraorbital ventral to the orbit; large crescent shaped infraorbital posterior and posteroventral to the orbit that contacts a single Y-shaped dermosphenotic dorsally; six small rectangular suborbital bones arranged in two distinct rows; wedge-shaped dermohyal posterior to a hatchet-shaped preoperculum; a row of three rectangular, ganoine-bearing anteopercular bones that extend down half the depth of the operculum; rectangular anteriorly inclined operculum with a diagonal ventral margin, suboperculum that is taller posteriorly than anteriorly and shorter in height but wider in length than the operculum; maxilla with a deep posterior plate, a rounded posteroventral process, and a narrow anterior arm that extends to the anteroventral margin of the orbit; a single median gular, two pairs of lateral gulars, a series of at least eight branchiostegal rays; inequilobate and deeply cleft heterocercal caudal fin with an elongated caudal peduncle; anteriorly placed and mid-body scales with pectinated posterior margins and diagonal ridges of ganoine.Holotype and only specimen. KUVP 155843  in age and are good comparisons for this newly described fish. The condition of the bones of the snout, infraorbitals series, suborbitals, and anteopercular bones in B. mercerensis n. gen. n. sp. is compared to that of Bear Gulch actinopterygians and other Carboniferous fishes in the sections below to highlight how this new taxon is distinct from previously described actinopterygians.Bones of the snout. The snout of Carboniferous lower actinopterygians is characterized by a great deal of anatomic diversity in terms of the numbers and identities of its bones of this region. Carboniferous actinopterygians differ from Devonian fishes in that there are numerous Carboniferous species with separate and distinct lacrimal and premaxillary bones, a condition not seen in Devonian fishes (Canobius ramsayi (Kalops (Lineagruan (Beagiscus (Bluefieldius mercerensis n. gen. n. sp. is an example of another Carboniferous actinopterygian with separate lacrimal and premaxillary bones (Bluefieldius n. gen. and the genera Kalops and Paratarrasius from the Bear Gulch Limestone (Bluefieldius n. gen. (Kalops and Bluefieldius mercerensis n. gen. n. sp. differ in the shape of the lacrimal bone (Kalops (B. mercerensis n. gen. n. sp. (Bluefieldius n. gen. n. differs from Kalops in regard to the absence of supraorbital bones (n fishes . Lower a ramsayi  and Bear (Kalops , Lineagrneagruan , and Beaeagiscus . Bluefiery bones ,  4. Theimestone ; Fig. 8.imestone . The elo n. gen. . Kalops . n. sp. . In addial bones .Infraorbital bones.B. mercerensis n. gen. n. sp. has one large crescent shaped infraorbital bone in the posteroventral corner of the orbit (B. n. gen. shares with many Devonian and Carboniferous genera such as Mimipiscis (, Moythomasia (, Howqualepis (Mansfieldiscus (Gonatodus (Woodichthys (Wendyichthys (, Cyranorhis (, Lineagruan, and Beagiascus (Bluefieldius mercerensis n. gen. n. sp. differs from these Devonian and Carboniferous fishes in regards to the composition of the bones of the snout and other features of the cranial bones. It is important to note that the contact between the large crescent shaped infraorbital and the dermosphenotic is another feature that sets Bluefieldius n. gen. apart from the genus Kalops (. Kalops has a series of small infraorbital bones separating the crescent shaped infraorbital in the posteroventral corner of the orbit from the dermosphenotic (he orbit ,  4, 8. mipiscis , Moythomthomasia , Howqualqualepis , Mansfieeldiscus , Gonatodonatodus , Woodichdichthys , and theyichthys , Cyranorranorhis , Lineagragiascus . Bluefies Kalops . Kalops phenotic .Suborbital bones. Suborbitals are anamestic bones that are found posterior to the circumorbital bones, separating these bones from the preoperculum. Suborbital bones are uncommon in Devonian fishes; they are absent in, among others, Cheirolepis (Moythomasia durgaringa (Mimipiscis (Gogosardinia (Moythomasia nitida (Osoriochthys marginis (Mesopoma carricki (Wendyichthys and Cyranorhis (Lineagruan (Paratarrasius (Kalops (Beagiascus (Bluefieldius mercerensis n. gen. n. sp., with six small suborbital bones (irolepis , Moythomrgaringa , Mimipismipiscis , and Gogsardinia . The onla nitida  with onemarginis  which hamarginis . Suborbicarricki  and the ranorhis , are chaneagruan , or a moarrasius , Kalops  (Kalops , and Beaagiascus . Bluefieal bones ,  4, is Anteoperular bones.Bluefieldius mercerensis n. gen. n. sp. is characterized by the third condition—there is a series of anteopercular bones posterior to a wedge-shaped dermohyal and anterior to the operculum (Lambeia pectinatus from the Albert Shale Formation (Paratarrasius (Lineagruan and Beagiascus (Bluefieldius n. gen. differs from Lineagruan and Beagiascus in the extent of the anteopercular bones. In B. mercerensis n. gen. n. sp., the anteopercular bones extend down to about half the height of the operculum; in Lineagruan, Beagiascus, and Lambeia the anteopercular bones extend down the entire height of the operculum, or even to the ventral margin of the suboperculum (Bluefieldius n. gen. and Lineagruan. The anteopercular bones, though ornamented with ganoine, are thin and delicate in Lineagruan judithi whereas the anteopercular bones in Bluefieldius mercerensis n. gen. n. sp. are of the same thickness as the surrounding bones, a condition similar to that seen in Lambeia (Paratarrasius has anteopercular bones that do not extend down to the ventral margin of the operculum (Bluefieldius n. gen. in terms of size, shape, and number (perculum ,  4. Thiormation  and the arrasius  and Lineagiascus . Bluefieperculum . The thi Lambeia . Paratarperculum , but thed number .B. mercerensis n. gen. n. sp. is defined by a unique set of cranial characteristics including morphological characters of the snout, the circumorbital series, suborbital bones, and anteopercular bones. The combination of these specific characteristics separate B. mercerensis n. gen. n. sp. from previously described lower actinopterygian fishes and warrant the description of a new taxon.A new lower actinopterygian genus and species is described from the Upper Carboniferous Bluefield Formation of West Virginia. This new genus is represented by a well preserved articulated specimen which represents the first vertebrate body fossil from the Bluefield Formation and the first described actinopterygian. Lower actinopterygian fishes are characterized by a great deal of anatomic and taxonomic diversity that is not well understood. We have neither a stable classification scheme nor strongly supported hypotheses of relationships for lower actinopterygian fishes. This lack of understanding has been attributed to the need for more well-preserved fishes to be described or redescribed and a better understanding of morphological characters among lower actinopterygians . Taxonom


This article has been retracted: Due to numerous examples of splicing and transposition of data into figures, as well as other errors, the authors believe that retraction of this paper is the only responsible and logical course of action. As a result, all authors have agreed to the retraction of this paper from Oncotarget.1094-1108. https://doi.org/10.18632/oncotarget.378Original article: Oncotarget. 2011; 2:1094–1108.


The evolution of elongated body forms in tetrapods has a strong influence on the musculoskeletal system, including the reduction of pelvic and pectoral girdles, as well as the limbs. However, despite extensive research in this area it still remains unknown how muscles within and around bony girdles are affected by these reductions. Here we investigate this issue using fossorial amphisbaenian reptiles, or worm lizards, as a model system, which show substantial variation in the degree of reductions of girdles and limbs. Using iodine-based contrast-enhanced computed tomography (diceCT), we analyze the composition of the shoulder muscles of the main clades of Amphisbaenia and their outgroups relative to the pectoral skeleton.Bipes is the only taxon that retains forelimbs and an almost complete pectoral girdle. All other amphisbaenian families show more variation concerning the completeness of the pectoral girdle having reduced or absent girdle elements. Rhineura, which undergoes the most severe bone reductions, differs from all other taxa in possessing elongated muscle strands instead of discrete shoulder muscles. In all investigated amphisbaenians, the shoulder muscle agglomerate is shortened and shifted anteriorly relative to the ancestral position as seen in the outgroups.All investigated amphisbaenian taxa retain the full set of 17 shoulder muscles, independent of the degree of limb and girdle reductions, whereas in some cases muscles are fused to complexes or changed in morphology relative to the ancestral condition. Our results show that pectoral muscle anatomy does not necessarily correspond to the loss or reduction of bones, indicating a decoupling of the musculoskeletal system. Muscle attachment sites change from bones to non-skeletal areas, such as surrounding muscles, skin or connective tissue, whereas muscle origins themselves remain in the same region where the pectoral bones were ancestrally located. Our findings indicate a high degree of developmental autonomy within the musculoskeletal system, we predict that the observed evolutionary rearrangements of amphisbaenian shoulder muscles were driven by functional demands rather than by developmental constraints. Nevertheless, worm lizards display a spatial offset of both pectoral bones and muscles relative to the ancestral position, indicating severe developmental modifications of the amphisbaenian body axis.The online version of this article (10.1186/s12862-018-1303-1) contains supplementary material, which is available to authorized users. In tetrapods, the evolution of elongated body forms is tied to substantial modifications of the musculoskeletal system and has shown to be strongly correlated with the reduction of limbs –7. Limb Bipes, possesses forelimbs [Bipes retaining large parts of the bony shoulder girdle and Blanidae still having ossified clavicles and scapulocoracoids, whereas Trogonophidae only possess bony scapulocoracoids and poorly ossified remnants of the sternum; Amphisbaenidae show either no or only very small bony scapulocoracoids and Rhineuridae do not retain any element of the pectoral girdle  lifestyle. Next to their snake-like appearance, amphisbaenians are anatomically characterized by a strongly reinforced skull that is used as a digging tool, the loss of the middle ear, and the reduction of the eyes , 24–29. orelimbs , 30, 31.Meroles cuneirostris, as outgroup taxon for comparison.For our study we used diffusible iodine-based contrast-enhanced computed tomography (diceCT), a recently developed technique for the 3D visualization of metazoan soft tissues –40. As w2O dest; 0,149 g/l in 100 ml H2O = ~ 0,15 w/v I in H2O), and applied this method to individuals belonging to five amphisbaenian species, Rhineura floridana, Bipes biporus, Blanus strauchi, Trogonophis wiegmanni and Cynisca leucura, which are representatives of the five major clades of Amphisbaenia, respectively  [Meroles cuneirostris, see Additional file C. leucura, 33 days for B. biporus, 72 days for T. wiegmanni, 79 days for R. floridana, and 86 days for M. cuneirostris. After completed staining, specimens were scanned using a Phoenix X-ray Nanotom computed tomography machine at MfN using 90 kV, 110 MA and 1000 ms for R. floridana and 40 kV, 250 μA und 1250 ms for all other specimens, with a resolution between 4 and 8 μm. The resulting 3D model was visualized and analysed using the software Volume Graphics 2.0. Muscle terminology follows Jenkins and Goslow (1983) [We used a personally developed protocol for diceCT visualization (iodine-potassium iodine solution from stock solution diluted with HCynisca)  (see AddCynisca) –44, we uw (1983) . The ideWe used the software Mesquite (version 3.10)  to optimBones of the shoulder girdleBipes biporus is the most complete among all amphisbaenians [sternum (st), which is accompanied posteriorly by an extremely long, also calcified and cartilaginous xiphoid process (xp). There is no connection between the sternum and the ribs. Anteriorly it is connected to the well-developed scapulocoracoids (scc) [humerus (hu). Distally, the scapulocoracoids articulate with the cartilaginous, but calcified, rectangular [suprascapulae (ss). The suprascapulae are situated dorsally to the scapulocoracoids and possess an anteroventral process, which extends medial to the clavicles (cl), but does not contact them. The latter are reduced, small, slender and rod-like bones that are connected to the shoulder girdle only via ligamentous connections to the sternum [Bipes biporus is positioned relatively close to the head at the level of the 3rd cervical vertebra [hyoid (hy) is also marked in all specimens.The shoulder girdle of baenians , 48  [Among amphisbaenians, Bipedidae are the only family with forelimbs  and posspiphyses . The bonpiphyses . Bipes b3, 3, 3) . For a cMuscles of the shoulder girdleBipes biporus, as well as those of the other amphisbaenian taxa, was made on the basis of their bony attachment sides, their overall topological arrangement, and generally by outgroup comparison. Because all respective muscle are described in detail for Meroles cuneirostris  and an interior  layer.M. latissimus dorsi (m ld) occurs as a large, flat, dorsolateral muscle, similar to that of Meroles cuneirostris, but a little smaller relative to the other muscles. Posterior it is attached to the M. episternocleidomastoideus and M. trapezius-complex. It lies distally to the M. deltoideus clavicularis and arises from a dorsal layer of connective tissue as well as from the surface of the M. episternocleidomastoideus and M. trapezius-complex and runs ventrally to insert into the deltopectoral crest of the humerus.The M. episternocleidomastoideus and M. trapezius are shown as separate muscles for Bipes canalicatus [, and also are divided into several portions. In our scans of Bipes biporus, M. episternocleidomastoideus and M. trapezius definitely form a large fibrous complex (m etc) with some fibres interacting, and no clear division can be made. Our observation is in accordance with Bolk et al. [Bipes biporus this complex forms a relatively large muscle unit, which lies distally and anteriorly to the M. deltoideus clavicularis. The complex covers most of the M. deltoideus clavicularis and expands laterally and anteriorly in form of a flat layer that extends beyond the level of the M. deltoideus clavicularis. Similar to Meroles cuneirostris, in which both muscles are separate, the complex runs relatively far anterior, i.e. it covers the area from the second vertebra to the posterior part of the skull and covers also parts of the hyoid. The M. episternocleidomastoideus and M. trapezius complex arises from a dorsal layer of connective tissue near the skin and inserts into the cranium.The alicatus , and alsk et al. , state tM. omohyoideus und M. sternohyoideus form a complex (m osc) in Bipes biporus. This complex runs in a layer proximal to the M. episternocleidomastoideus and M. trapezius-complex. Similar to the latter, it arises from the upper third of the lateral body half from an inter-muscular area of connective tissue and runs to the hyoid. Together with the M. episternocleidomastoideus and M. trapezius-complex it is one of the most anteriorly positioned muscle units of the shoulder girdle.As mentioned above, M. deltoideus scapularis (m ds) is a flat muscle that arises from the suprascapula and runs to the deltopectoral crest of the humerus, where it also inserts. It is smaller than the M. deltoideus scapularis of Meroles cuneirostris and much more posteriorly located.The M. deltoideus clavicularis (m dc) is located, which runs distal to the suprascapula and extends beyond its level both anteriorly and posteriorly. It arises from the suprascapula and inserts into the scapulocoracoid as well as into the surface of the adjacent M. supracoracoideus anteriorly and M. deltoideus scapularis posteriorly. In comparison to the M. deltoideus clavicularis of Meroles cuneirostris, the M. deltoideus clavicularis of Bipes biporus exhibits no twist around its own axis. It is a long, flat muscle that is located not only anteroventrally, as in Meroles cuneirostris, but also dorsally to the M. deltoideus scapularis. It forms a muscle layer that is bordered distally by the M. episternocleidomastoideus and M. trapezius-complex as well as by the M. latissimus dorsi, and proximally by the suprascapula.Dorsal to M. deltoideus scapularis and M. serratus anterior the M. supracoracoideus (m supc) is located. It is a long, flat muscle that runs between the attachment of the humerus to the shoulder girdle and the scapulocoracoid, from which it arises. Proximally it is strongly associated with the M. scapulo-humeralis anterior, from which it is also difficult to separate, at least in the contact zone. This observation is in accordance with Bolk et al. [Meroles cuneirostris there is a strong association between the M. supracoracoideus and the M. scapulo-humeralis anterior, but they can be much better separated from each other than in Bipes biporus. Here, the M. supracoracoideus runs with a twist around its own axis in direction of the deltopectoral crest of the humerus, where it also inserts.Anteroventrally to the M. deltoideus scapularis the k et al. , who staM. pectoralis (m p) consists, like in Meroles cuneirostris, of diverse layers, which partly overlap each other. As a superficial muscle it arises from a layer of connective tissue between skin and sternum and runs across the sternum and inserts into the deltopectoral crest of the humerus. Both anterior and posterior to the humerus it extends far dorsally and covers the M. episternocleidomastoideus and M. trapezius- complex anteriorly. Posteriorly it is connected to the M. latissimus dorsi. It runs distally to the M. coraco-brachialis longus.The flat Subjacent muscle layersThe subjacent pectoral muscles are also positioned in two layers in all non-rhineurid amphisbaenians, an exterior  and an interior  layer.M. serratus anterior superficialis (m sas) is located. It is a flat muscle that runs proximally from the suprascapula to the scapulocoracoid and inserts into the deltopectoral crest of the humerus.Proximal to the M. deltoideus scapularis the M. scapulo-humeralis anterior (m sha) runs proximal to the M. supracoracoideus and ventral to the M. deltoideus clavicularis. It arises from both the clavicle and the scapulocoracoid, runs directly along the scapulocoracoid and inserts into the deltopectoral crest of the humerus.The M. scapulo-humeralis posterior (m shp) is located proximolaterally, whereas both are strongly associated with each other. The origin of the M. scapula-humeralis posterior lies on the scapulocoracoid and its insertion is found on the surface of the adjacent muscles, i. e. M. subcoraco-scapularis, M. coraco-brachialis brevis and M. coraco-brachialis longus.Relative to the latter muscle, the M. levator scapulae (m ls) is located, which arises from the lateral process of the atlas in the same way as in Lacertidae [Meroles cuneirostris, in which it is located anterior to the M. serratus anterior superficialis.Proximal to the suprascapula, the certidae . It runsM. coraco-brachialis brevis (m cbb) is located below the articulation of the humerus to the shoulder girdle. It is a flat, bulky muscle that runs ventrally to the M. supracoracoideus and M. scapulo-humeralis anterior, as well as the M. omohyoideus and the M. sternohyoideus-complex. It arises from a layer of inter-muscular connective tissue and inserts into another layer of connective tissue as well as into the surface of the adjacent M. coraco-brachialis longus.The M. coraco-brachialis longus (m cbl) is located. It is larger than the former muscle and arises from the scapulocoracoid, inserting into the deltopectoral crest of the humerus, where it is positioned adjacent to the M. biceps brachii.Proximally, deep below the M. coraco-brachialis brevis, the Bipes biporus is the M. subcoraco-scapularis (m subcs), an aggregation of M. subcoracoideus and M. subscapularis [Meroles cuneirostris. It arises from a ventrally positioned layer of connective tissue inside the body and inserts into the deltopectoral crest of the humerus and is positioned ventrally to the M. serratus anterior superficialis.One of the most proximally situated muscles of apularis , which rM. sternocoracoideus (m sternc) is found, which is relatively large compared to Blanus and Trogonophis. It arises in a layer of connective tissue near the sternum and runs around the deltopectoral crest of the humerus, where it inserts.Laterally to the M. subcoraco-scapularis the Muscles of the upper forelimbM. biceps brachii (m bb), the M. brachialis (m b) and the M. triceps (m t). They all run along the humerus in a similar way and with identical attachment sites as in Meroles cuneirostris  it is apparent that there is variation in both the presence of vestigial clavicles and in the size of the scapulocoracoids: In some specimens the clavicles are absent; in other specimen the scapulocoracoids are smaller than described in the literature. This variation most likely explains the absence of clavicles and the reduced size of the scapulocoracoids in our stained specimen.Several authors , 30, 53 lavicles . In our Muscles of the shoulder girdleOverview of present musclesBlanus strauchi all muscles of the shoulder girdle are present, consisting of nine superficial  are absent.All nine superficial see Fig. a, b and Superficial muscle layersM. latissimus dorsi is a prominent, flat, dorsolaterally running muscle, which is larger than in Meroles cuneirostris, Bipes biporus and Blanus strauchi. It arises dorsally from a layer of connective tissue and runs ventrally, i.e. directly next to the similarly flat complex of M. episternocleidomastoideus and M. trapezius, which results in the absence of any sharp division between the two muscles units. It inserts into another ventral layer of connective tissue near the skin.The M. episternocleidomastoideus and M. trapezius-complex represents a superficial muscle mass that arises from the vertebral column, runs anteroventrally, and inserts near the cranium. Furthermore, the M. episternocleidomastoideus and M. trapezius-complex is the most anteriorly situated muscle mass, similar to Blanus strauchi, but is smaller than in the latter. The M. latissimus dorsi and the M. episternocleidomastoideus and M. trapezius-complex of Trogonophis wiegmanni run far ventrally, similar to Bipes biporus. Another important difference is that the M. latissimus dorsi and the M. episternocleidomastoideus and M. trapezius-complex are located directly next to each other in Trogonophis wiegmanni, whereas in Blanus strauchi there is no association between the M. latissimus dorsi and the M. episternocleidomastoideus and M. trapezius complex.The M. omohyoideus and M. sternohyoideus form a complex, whereas it seems that the M. omohyoideus adapts to the course of the M. sternohyoideus, because the complex arises from the scapulocoracoid and runs anteriorly to insert into the hyoid. In contrast, the course of the M. omohyoideus and M. sternohyoideus-complex is dominated by the M. omohyoideus in the other examined taxa. The M. omohyoideus has generally been described to arise from both the clavicles and the interclavicle in squamates [Trogonophis wiegmanni the clavicles are reduced, whereas the scapulocoracoids and a cartilaginous sternum are still present (see above). The muscle complex arises from the scapulocoracoid, which is why we assumed that the M. sternohyoideus is the more dominant part within this complex. Because the complex consists of two long, parallel running muscles, one can assume that there was a fusion with the M. omohyoideus. A differentiation between the M. omohyoideus and the M. sternohyoideus is therefore not possible.Also quamates , whereasquamates . In TrogM. deltoideus scapularis is dorsally overlain by the M. latissimus dorsi. The muscle shows a more strand-like morphology than in Blanus strauchi. Also, the fibre orientation of the M. deltoideus scapularis of Trogonophis wiegmanni is clearly different from the dorsoventral orientation seen in Blaunus strauchi. In Bipes biporus the M. deltoideus scapularis arises from the suprascapula, whereas in Trogonophis wiegmanni it arises from the surface of the M. levator scapulae and M. longissimus. The M. longissimus is a skeletal muscle of the back and runs in direction to the head. It belongs to the epaxial ancestral muscles, which are situated dorsally to the transverse processes of the vertebrae [The massive and large ertebrae . The oriM. deltoideus clavicularis is found. It arises from the scapulocoracoid and joins the whole course of the M. deltoideus scapularis with parallel-running fibres, until it also ends in the already mentioned area of connective tissue near the cranium. It inserts also in parts into the ventrally running M. supracoracoideus. The M. deltoideus clavicularis also is covered by the M. latissimus dorsi dorsally and by the M. pectoralis ventrally.Ventrally to the M. deltoideus scapularis the narrow, elongated M. supracoracoideus is located ventrally to the M. deltoideus clavicularis that arises from the scapulocoracoid, joins the course of the M. deltoideus scapularis and the M. deltoideus clavicularis, and inserts near the cranium, but a bit further anteriorly than the latter two. The M. deltoideus scapularis, the M. deltoideus clavicularis and the M. supracoracoideus all diverge strongly from their morphology in Meroles cuneirostris and Bipes biporus. While they represent a fan-shaped structure in Meroles cuneirostris and Bipes biporus, they appear as elongated muscle strands in Trogonophis wiegmanni.The M. pectoralis completes the shoulder muscle complex ventrally. It arises from the scapulocoracoid near the sternum and runs in form of several thin, partly overlapping layers across the scapulocoracoid anteriorly. The muscle reaches far dorsally, even more than in Blanus strauchi. In its dorsal region it is in contact with the M. latissimus dorsi, which is slightly overlain in the area where both muscles meet. The M. pectoralis inserts into a layer of connective tissue near the skin.The Subjacent muscle layersM. serratus anterior superficialis is located proximally to the M. deltoideus scapularis, by which it is also almost completely covered. In its morphology it is quite similar to that of Blanus strauchi. It arises posterior to the origin of the M. levator scapulae on the first vertebra and runs caudally, where it inserts near the scapulocoracoid on the surface of the M. scapulo-humeralis anterior and the M. scapulo-humeralis posterior. As in Meroles cuneirostris, fibres run parallel with a posteroventral to anterodorsal propensity.The M. scapulo-humeralis anterior is located, representing a relatively small, elongated muscle, which is covered distally by the M. deltoideus scapularis and the M. deltoideus clavicularis. It arises from the scapulocoracoid and runs anteriorly to the M. serratus anterior superficialis, where it also inserts. The muscle is similar to the M. scapulo-humeralis anterior of Blanus strauchi, whereas in Trogonophis wiegmanni it runs more distally.Lateral to the M. serratus anterior superficialis the M. scapulo-humeralis posterior is situated, which is completely covered by the distally running M. deltoideus clavicularis. It arises near the scapulocoracoid and runs anteroproximally into the center of the body to insert into an area of connective tissue near the M. coraco-brachialis brevis.Ventrally to the M. serratus anterior superficialis the elongated M. levator scapulae belongs to the most proximally situated muscles and arises from an inter-muscular area near the lower level of the cranium. Shortly behind its origin it makes a half-turn around its own axis and runs posteriorly with a dorsal drift, inserting into the surface of the M. longissimus. With respect to its position relative to the cranium, the M. levator scapulae of Meroles cuneirostris and Trogonophis wiegmanni superficially show similarities, although the insertion areas are different. While the M. levator scapulae of Meroles cuneirostris inserts into the scapula and sternum, in Trogonophis wiegmanni this insertion is into the surface of the M. longissimus. The insertion into another muscle is similar to the situation in Blanus strauchi, in which the M. levator scapulae inserts into M. supracoracoideus (see above). The M. levator scapulae is also similar to that of Blanus strauchi with regards to its shape and cranial origin near the hyoid.The M. coraco-brachialis brevis strongly differs in morphology from Meroles cuneirostris and the other examined amphisbaenians. In Meroles cuneirostris it is a relatively large and spacious muscle, whereas in Trogonophis wiegmanni it is rather small and oval in shape. The muscle is situated relatively medially inside the body and is not in contact with any other muscle except for the M. coraco-brachialis longus. It arises and inserts into connective tissue, close to the M. scapulo-humeralis posterior.The M. coraco-brachialis longus arises in the ventral region of the body near the skin. It shows a skewed anterodorsal course and is of trapezoid shape, whereas its apex points anteriorly. It inserts into a layer of connective tissue near the origin of the M. levator scapulae. In Trogonophis wiegmanni this muscle is large in comparison to the other examined species.The M. subcoraco-scapularis is a combination of the M. subcoracoideus and M. subscapularis [Trogonophis wiegmanni there is a visible border between these two units. The larger, bulkier of the two originates from an area of connective tissue near the M. deltoideus clavicularis and runs distally from there to another intramuscular bundle of connective tissue inside the body near the M. serratus anterior superficialis. The other, smaller and more elongated part is situated rather proximally and arises from the surface of the M. omohyoideus and M. sternohyoideus-complex, running in direction towards the M. scapulo-humeralis posterior, where it inserts inside connective tissue situated between the latter and the M. serratus anterior superficialis. A reliable separation of the two units into M. subcoracoideus und M. subscapularis, respectively, is not possible due to the non-skeletal origin and insertion areas in Trogonophis wiegmanni. Therefore, both parts are here treated as M. subcoraco-scapularis.The apularis . In TrogM. sternocoracoideus arises from the sternum and runs along and inserts into the scapulocoracoid, arching slightly anterior. Especially in Trogonophis wiegmanni it represents the linkage between sternum and scapulocoracoids as described by Gans [Trogonophis wiegmanni the M. sternocoracoideus is a relatively small muscle that is strongly associated with the M. omohyoideus und M. sternohyoideus-complex, by which it is also covered distally.The  by Gans . In TrogBones of the shoulder girdleCynisca possesses no shoulder girdle elements, but Zangerl [Cynisca leucura as small, rod-like, bony structures that we interpret as scapulocoracoids, which are located at the level of the third vertebra , four pectoral strands  and one discrete muscle (dm).Within the ancestral shoulder region of see Fig. a are preThe 1st lateral muscle runs as a thick and large muscle from the spinal process of the third vertebra, ventrally to the 1st dorsal strand, and inserts near the anterior end of the 3rd neck muscle inside a narrow area of connective tissue in the postcranial region directly behind the skull.The 2nd lateral muscle is a thin layered muscle that arises from both the skin and an aponeurosis at the level of the third vertebra, running anteriorly to the lower postcranial region behind the skull, where it inserts into the cranium through slender fibres. It partly covers the course of the 1st lateral muscle. It is surrounded by the 1st lateral muscle and the 7th lateral muscle dorsally as well as by the 3rd lateral muscle ventrally.The 3rd lateral muscle runs ventrally to the 2nd lateral muscle, which it also partially covers. It arises ventrally to the origin of the 2nd lateral muscle from both the skin and an aponeurosis right beneath the skin, running anteriorly to the same lower postcranial region as the 2nd lateral muscle, where it inserts into an area of connective tissue near the skin. The 3rd lateral muscle shows mostly the same shape as the 2nd lateral muscle, including the overall elongated and thin-layered condition.The 4th lateral muscle is very thin and flat. It arises from an area of connective tissue near the skin and runs anteriorly to insert inside an inter-muscular region at the level anterior to the transverse process of the first vertebra. The muscle becomes slightly broader anteriorly. It is surrounded by the 3rd lateral strand dorsally and the 1st pectoral strand ventrally.The 5th lateral muscle is very similar to the 4th and strongly associated with it. It has the same origin and follows its course proximally to insert partly into the same inter-muscular region as the 4th lateral muscle as well as inside the surface of the 4th lateral muscle. It is thin and very flat. Distally it is partly overlain by the 1st pectoral strand.The 1st pectoral strand runs ventrally to the 4th and 5th lateral strands and arises at the posterior level of the third vertebra from a tiny inter-muscular region. It is fibrous and thin and runs along the transverse process of the third to first vertebrae along the vertebral column. It becomes flatter and broader anteriorly, until it inserts inside an inter-muscular region at the level of the transverse process of the first vertebra.The 2nd pectoral strand runs ventrally to the 1st pectoral strand, with which it is strongly associated. At some points, the 1st and 2nd pectoral might be even seen as a single muscle. The 2nd pectoral muscle arises from second rib at the level of the third vertebra and runs anteriorly to insert into the skin at the level of the transverse process of the first vertebra. Like the 1st pectoral strand, it is overall very flat.The 3rd pectoral strand is a bulky muscle that runs proximal to the 1st and 2nd pectoral strand, by which it is also partly overlain. It arises from the second rib at the level of the third vertebra and runs anteriorly to insert into an inter-muscular region near the lateral postcranial region right behind the skull. It partly encases the 4th pectoral muscle which is orientated proximally.The 4th pectoral strand runs proximally and ventrally to the 3rd pectoral strand. It arises ventrally from the transverse process of the third vertebra and runs anteriorly to insert near the postcranial region into the transverse process of the 1st vertebra as well as into an inter-muscular region near the lateral postcranial region. It consists of a flat and thin muscle layer that tapers along the transverse processes from the third to the first vertebra. It is the most ventrally positioned muscle.discrete muscle in Rhineura floridana is positioned relatively far anterior within the animal. It arises from a tiny inter-muscular region near the skin at the level of the first vertebra. It is a small, elongated muscle that runs anteriorly and inserts into inter-muscular region near the skin. It has no connection to the other muscle strands and shows a dorsally trending course towards the cranium. It is smaller than all other muscles described above.The only Subjacent muscle layers of the considered shoulder girdle regionRhineura floridana there are eight subjacent muscle strands .Within the ancestral shoulder girdle region of see Fig. b, consisThe 6th lateral strand is a short, but massive muscle. It arises from an inter-muscular region near the lateral side of the anterior end of the neural spine of the second vertebra and runs anteriorly, passing the neural arch of the first vertebra laterally and inserting into the lower cranium.The 7th lateral strand is more elongated than the 6th. It arises from a rounded elevation on the neural arch of the third vertebra and runs anteriorly along the vertebral column, covering the third to first vertebra. It touches the 6th lateral strand near the latter’s point of origin and inserts into the lower cranium distally to the insertion of the 6th lateral strand.The 8th lateral strand is not as large as the 7th lateral strand. It originates from the transverse process of the third vertebra and runs anteriorly along the transverse processes of the second and first vertebrae, inserting into an inter-muscular region anteriorly to the transverse process of the first vertebra. It is distally overlain by the 2nd and 3rd lateral strand.The 9th lateral strand runs ventrally to the 8th lateral strand, to which it is also very similar. It tapers along the transverse processes of the vertebral column anteriorly to the transverse process of the first vertebra, where it inserts into the 8th lateral strand slightly posterior to the latter’s insertion side. The 8th and 9th lateral strands are partly associated. The 9th lateral strand is also distally overlain by the 2nd and 3rd lateral strand.The 10th lateral strand is a very small and short muscle. It arises from the beginning of the second rib near the vertebral column at the level of the third vertebra and runs anteriorly to insert into the beginning of the first rib near the vertebral column. Due to its position, the 10th lateral strand can be seen as an inter-rib muscle, because it links the second and first rib with each other.The 11th lateral strand is an elongated, slender muscle strand and larger than the 10th lateral strand. It arises from the distal tip of the second rib at the level of the third vertebra and runs anteroproximally to the 5th lateral strand, inserting into an inter-muscular region near the insertion of the first rib at the level of the second vertebra.The 12th lateral strand is situated anteriorly to the 11th lateral strand and is of largely similar size. It arises from an inter-muscular region near the 3rd lateral strand and runs anteriorly towards the cranium to insert into the first vertebra laterally.The 13th lateral strand arises from the head of the second rib and runs anteriorly to the head of the first rib, where it also inserts. It is a small muscle similar to the 10th lateral strand, but it runs more ventrally, on the ventral side of the ribs. It might be viewed as an inter-rib muscle, because it creates a linkage between the first and second rib. Distally it is completely covered by the 2nd pectoral strand.Rhineura floridana that features 18 muscles within the pectoral region, all amphisbaenian taxa examined in this study possess the ancestral number of 17 shoulder and pectoral muscles. Some of these muscles occur as complexes, such as the M. omo- and sternohyoideus- complex and the M. episternocleidomastoideus and trapezius complex. In some cases the muscles are also strongly associated with the surrounding neck- and dorsal musculature, with which they seem to form a functional unit. This applies especially to the M. episternocleidomastoideus and trapezius-complex, the M. omo- and sternohyoideus-complex, the M. deltoideus scapularis, and to the M. levator scapulae. In all examined amphisbaenians as well as in Meroles cuneirostris, the M. latissimus dorsi, the M. episternocleidomastoideus and the M. trapezius-complex form an outer muscle arrangement dorsally, whereas the M. pectoralis covers the pectoral region ventrally. In Blanidae, Trogonophidae and Amphisbaenidae, the dorsally covering muscles are elongated ventrally and approach the dorsally elongated M. pectoralis laterally. In Blanus there is no uninterrupted outer muscle cover, whereas in Trogonophis and Cynisca there is an continuous cover in form of a thin external muscle layer, which surrounds the entire pectoral area right beneath the skin. A notable feature observed in all amphisbaenian taxa is that the shoulder girdle agglomerate is shortened and shifted anteriorly relative to the ancestral position as seen in Meroles cuneirostris , sacral and caudal series based on changes in skeletal morphology and integration with the appendicular skeleton. Both the number of regions, their morphologies, and the number of elements composing them are variable across taxa , 59. In Hox gene expressioni, whereas the appendicular skeleton, including the sternum, ventral ribs, girdles, limbs, and most associated muscles, are patterned from lateral plate mesoderm by distinct Hox expression [The relationship of the pectoral girdle to the axial skeleton in amphisbaenians is similarly spatially offset, suggesting dissociation between developmental domains. Dorsal vertebrate and ribs are patterned from paraxial mesoderm by collinear pression . These tpression –66 with pression , 65. Heapression  suggesteBipes a restructuring of the shoulder muscles can be observed, which is likely due to the loss of humerus and forelimbs. In the original condition of a quadrupedal tetrapod, the M. deltoideus scapularis, the M. deltoideus clavicularis, the M. latissimus dorsi, the M. supracoracoideus and the M. pectoralis are closely associated with the deltopectoral crest of the humerus and play an important role in locomotion [Bipes, the attachment sites of the respective muscles have changed relative to the ancestral condition due to the reduction of the humerus. It is likely, therefore, that the function of the single shoulder muscles in Blanus, Trogonophis, Cynisca and Rhineura is no longer the ancestral, humerus-associated function. The muscles that originally served as retractors and protractors of the humerus [Evolutionary dissociation, such as developmental repatterning of one domain relative to another, is a potential mechanism for spatial reorganization of tissues . In the comotion , 68, 69. humerus  are veryRhineura is a special case and may require additional investigations), but the muscles within each layer have different developmental origins and can be considered either axial shoulder or intrinsic limb muscles [Although the functional role of the different shoulder muscles is still unknown for Amphisbaenia, we also predict that the evolutionary rearrangement of the muscles was not primarily driven by developmental constraints but by functional demands. This view is supported by the observation that there is a clearly visible division into two muscle layers, a superficial and a subjacent layer, which appear as clearly separated, ring-like structures in axial view. Both layers contain the same muscles across the different species , implies that also other representatives of Amphisbaenidae, including the shovel-headed forms, will show a similar pattern that is different from that of Rhineura. Although further studies (and more taxon sampling) are required, we thus consider our observations sufficient support for the hypothesis that shovel-headed morphologies evolved convergently and independently within Amphisbaenidae and Rhineuridae, contradicting previous assumptions about a monophyletic origin of shovel-headed amphisbaenians [The results of our study also represent the first, at least indirect, morphological support of the molecular-based hypothesis that shovel-headed amphisbaenians evolved independently, i.e. shovel-headed taxa other than e former . Shovel-e former . If shovbaenians .Using the pectoral region of amphisbaenians as a model system, our study provides novel insights into the relationship between muscles and bones during the reduction of shoulder girdle and limbs, indicating that muscle attachment sites that were originally connected to an ancestrally present bone shift to non-skeletal areas such as surrounding muscles, skin or connective tissue. At the same time, muscle origins themselves remain in the same region where the respective bones were ancestrally located. From this follows that pectoral muscle anatomy does not necessarily correspond to the loss or reduction of bones, indicating a decoupling of the musculoskeletal system. Also, we identified a relative anterior shift of the position of both pectoral bones and muscles relative to the ancestral position as seen in other squamates, suggesting fundamental developmental modifications during the evolution of the amphisbaenian body plan. We predict that the observed evolutionary rearrangements of shoulder muscles in worm lizards were likely driven by functional demands rather than developmental constraints. Furthermore, our study provides indirect morphological support for shovel-headed morphologies to have evolved convergently and independently across Amphisbaenia.For future studies increased taxon sampling would be preferable, especially with respect to shovel-headed forms. In addition, studies on other limb-reduced lizard taxa, such as representatives from Scincidae or Gymnophthalmidae, may provide further, independent insights into to the modifications of the musculoskeletal system during the evolutionary transformation of the squamate body axis.Additional file 1:Table S1. Species table. Table contains information about all specimens used in the study, abbreviations: FMNH: Florida Museum of Natural History, MfN: Museum für Naturkunde Berlin. (DOCX 14 kb)Additional file 2:Text S2. Pectoral muscle morphology in Lacertidae (Meroles cuneirostris). Text contains detailed information about pectoral bones and muscles in Meroles cuneirostris, abbreviations: cc: coracoid, cl: clavicles, dc: deltopectoral crest of humerus, hu: humerus, hy: hyoid, m b: M. brachialis, m bb: M. biceps brachii, m cbb: M. coraco-brachialis brevis, m cbl: M. coraco-brachialis longus, m dc: M. deltoideus clavicularis, m ds: M. deltoideus scapularis, m e: M. episternocleidomastoideus, m etc: M. episternocleidomastoideus and M. trapezius-complex, m ld: M. latissimus dorsi, m ls: M. levator scapulae, m o: M. omohyoideus, m p: M. pectoralis, m sas: M. serratus anterior superficialis, m sha: M. scapulo-humeralis anterior, m shp: M. scapulo-humeralis posterior, m sternc: M. sternocoracoideus, m sternh: M. sternohyoideus, m subcs: M. subcoraco-scapularis, m supc: M. supracoracoideus, m t: M. triceps, m tp: M. trapezius, ss: suprascapulae, st: sternum, v: vertebra. (DOCX 35 kb)Additional file 3:Figure S3. Skull, anterior part of the vertebral column and pectoral region of Meroles cureirostris. A: lateral view, B: ventral view. Abbreviations: cc: coracoid, cl: clavicles, hu: humerus, hy: hyoids, icl: interclavicle, s: scapula, st: sternum, v: vertebra. (TIF 4277 kb)Additional file 4:Figure S4. Pectoral muscles of Meroles cureirostris, A: muscles of the upper forelimb of Meroles cureirostris, B: superficial pectoral muscles of Meroles cureirostris, C: distal subjacent pectoral muscles of Meroles cureirostris, D: superficial and subjacent pectoral muscles of Meroles cureirostris, E: medial subjacent pectoral muscles of Meroles cureirostris. Array shows anterior direction, abbreviations: dc: deltopectoral crest of humerus, hu: humerus, m b: M. brachialis, m bb: M. biceps brachii, m cbb: M. coraco-brachialis brevis, m cbl: M. coraco-brachialis longus, m dc: M. deltoideus clavicularis, m ds: M. deltoideus scapularis, m e: M. episternocleidomastoideus, m etc: M. episternocleidomastoideus and M. trapezius-complex, m ld: M. latissimus dorsi, m ls: M. levator scapulae, m o: M. omohyoideus, m p: M. pectoralis, m sas: M. serratus anterior superficialis, m sha: M. scapulo-humeralis anterior, m shp: M. scapulo-humeralis posterior, m sternc: M. sternocoracoideus, m sternh: M. sternohyoideus, m subcs: M. subcoraco-scapularis, m supc: M. supracoracoideus, m t: M. triceps, m tp: M. trapezius. (TIF 1862 kb)Additional file 5:Table S5. Pectoral muscle origins and insertions. Table contains information about pectoral muscles origins and insertions for all amphisbaenian families. (DOCX 15 kb)


Oxalis corniculata L. is a weed with a world-wide distribution and unknown origin. Though it belongs to a section of the genus from South America, the evidence that this species came from there is weak.O. corniculata using herbarium specimens, historic literature and archaeobotanical research. We also summarized ethnobotanical literature to understand where this species is most used by humans as a medicine.We reviewed the evidence for the origin of O. corniculata occurred in Europe before the 15th century. Nor is there reliable evidence that it occurred in North or South America before the 19th century. However, there is direct archaeobotanical evidence of it occurring in south–east Asia at least 5,000 years ago. There is also evidence from historic literature and archaeobotany that it reached Polynesia before European expeditions explored these islands. Examination of the traditional use of O. corniculata demonstrates that is most widely used as a medicine in south–east Asia, which, while circumstantial, also points to a long association with human culture in this area.Despite numerous claims that it is native to Europe there is no strong evidence that O. corniculata is south–east Asia. This is consistent with a largely circum-Pacific distribution of section Corniculatae of Oxalis. Nevertheless, it is likely that O. corniculata spread to Europe and perhaps Polynesia before the advent of the modern era through trade routes at that time.The most likely origin for  Oxalis corniculata L. is obscure or unknown. The species was first described by Carl Linnaeus from material from the Mediterranean region and some authors suggest that this region is the native range  and the bottle gourd (Lagenaria siceraria (Molina) Standl.) have been explained this way . Of these, O. exilis and O. filiformis are 20th century introductions with relatively narrow distributions (O. corniculata it is important to establish the time of introduction of O. stricta and O. dillenii to Europe.Europe has five extant species of ibutions . Owing tibutions . TherefoO. stricta (O. stricta) was so well established in Europe that O. europaea Jord. He did so based purely on the geographic grounds that the Linnaean O. stricta was North American. He did not give any morphological characters that separate them. Herbarium specimens do exist of O. stricta collected in Europe before 1854, but these are few and only date from earlier in the 19th century  in medieval herbals from the 13th to 15th centuries. In each case only one taxa is illustrated. In Europe we would expect at least one taxon, the native O. acetosella L. This species is quite distinct from O. corniculata in having white flowers, short globular pods, a scaly creeping rhizome and no stem. The illustration in the 15th century Codex Bellunensis and an unnamed codex have no flowers, but depict the scaly rhizome and the lack of stem, which suggests this is O. acetosella   . This isOxalis section Corniculatae remain rare. The RADAR database in the Netherlands contains four records, the earliest of which are from a site that spans the 17th and 18th centuries   . Thunberniculata . The nexniculata , but theO. corniculata and two new species O. procumbens and O. radicosa. O. procumbens and O. radicosa as distinct species. Oxalis procumbens is said to grow in fields and fallow land at altitudes above 1,000 m in tropical Africa and O. radicosa has a larger distribution spanning much of Africa, Asia and Australasia. Examples of specimens are in O. corniculata  Small, O. grandis Small, O. colorea  Fedde, O. stricta, O. suksdorfii Trel., O. dillenii, O. albicans Kunth, O. pilosa Nutt. ex Torr. & A.Gray, O. californica (Abrams) R.Knuth and O. texana  Fedde. Furthermore, in Mexico, Central America and the Caribbean there are other species, O. filiformis, O. rugeliana Urb. and O. thelyoxys Focke  are from 1815 to the 1830s  and Daxi culture   . Then ar7 seeds) . There a7 seeds) . The papO. corniculata the only other possibility is that these are seeds of O. stricta, which is believed to be native to China ; in the Philippines in the 1st millennium BC and in Vietnam between 3,450 and 3,250 BP  (tterfly) . This spO. corniculata we discovered a total of 182 publications containing 892 remedies for ailments at different locations. It is evident from this map that Asia is at the centre of where O. corniculata is used medicinally  and Tahiti (1769) when they were explored during the scientific expeditions of Captain James Cook  about 10 million years ago. Therefore, there is no reason to presuppose that O. corniculata s.s. evolved in South America. Furthermore, additional resolution of the phylogeny of the section Corniculatae is needed. Corniculatae and only two of the North American species.O. corniculata, there is no convincing evidence for its occurrence here earlier than the Renaissance. In contrast, there is clear evidence of O. corniculata in eastern Asia occurring for several thousand years. Archaeological evidence supports this conclusion. Furthermore, early specimens, literature and anthropogenic usage are consistent. This conclusion also makes sense regarding the proximity of its nearest relatives. These are native to Australasia, North America and South America. This means, if one accepts the eastern Asian origin of O. corniculata, that then Oxalis section Corniculatae has a largely circum-Pacific distribution, which we find rather more parsimonious than one member of the section naturally occurring in relative isolation in Europe.Although Europe has often been suggested as the origin of O. corniculata. Although this evidence is weak these results are consistent with a native origin of O. corniculata in Asia and there is little support for an origin elsewhere.There is undoubtedly considerable bias in the publication of ethnobotanical literature and in the intensity of collection of specimens from different parts of the world. In general, native plants are more often used in traditional medicines, but that is probably because there are more available . Plants O. corniculata into the Pacific islands with Polynesian voyagers is also consistent with an east Asian origin. Mitochondrial DNA and Y chromosome studies of Polynesians traces their origins back to Taiwan, the Philippines and Melanesia  Schott (taro) to these island  and the reopening of the silk routes at this time. However, the possible earlier presence of O. corniculata in Carthage indicates an earlier introduction to the Mediterranean Basin. It is interesting that peach stones also occur in the Carthage deposits. Peaches (Prunus persica (L.) Batsch) were introduced to Europe by the 1st century AD and peaches also have an Asian origin (rections . We accen origin . Coincidn origin . TherefoO. radicosa and O. procumbens has been established. In the case of South Africa, Cape Town was originally founded 1652 and was an important trading port for the Dutch East India Company during the 17th and 18th century. As such, it is not hard to envisage that O. corniculata was introduced there. Nevertheless, the testimony of Floras indicates that even if O. corniculata was present in Africa before the modern era, it has become more common and today has a wider distribution than previously.The situation in sub-Saharan Africa is still unclear and may only be resolved once the taxonomic position of O. calachaccensis, which is indistinguishable morphologically and genetically from O. corniculata. Its presence in Bolivia suggests that O. corniculata is either native to South America, or that the specimens described as O. calachaccensis are in fact introduced O. corniculata. Certainly, Knuth described O. calachaccensis from plants collected comparatively recently (O. calachaccensis are modern introductions and that O. calachaccensis should be considered a synonym of O. corniculata.In South America the final difficulty is the status of recently . We are O. corniculata globally. Taxa tend to be most genetically diverse in the area of their origin (O. pes-caprae L. (Relying upon physical evidence to determine the native origins of plant has its limitations and biases. A complementary method would be to examine the genetic diversity of r origin  and suchaprae L. . It is hO. corniculata is lost to time, we have shown that there is sufficient evidence to conclude that it is in fact native to East Asia and unlikely to be native to Europe. Many of the details of the global colonization by species of section Corniculatae remain a mystery, particularly in Africa and Australasia. Furthermore, it is hoped that this paper will raise awareness of the issues, so that new archaeological discoveries of O. corniculata remains are recognised for their importance in understanding the biogeography of this species.While many authors assume that the origin of


Leishmania major and Leishmania tropica are two main species causing cutaneous leishmaniasis (CL) in Iran. Recently, Crithidia spp. has also been reported in the wound of patients with CL. In this study, we determined the species causing CL in the southern of Iran and the role of Crithidia spp. in creating skin ulcers.Leishmania species and then to identify Crithidia spp.In this cross-sectional study from Apr to Sep 2016, 66 patients with CL referred to Diagnostic Lab of Leishmaniasis, Valfajr Health Center, Shiraz, Iran, were selected. After DNA extraction from the Giemsa stained smears, all samples were amplified in two separate steps using specific primers, firstly, to differentiate L. major and L. tropica were responsible for 60 and 6 cases, respectively. Moreover, in two patients, mixed infection with Crithidia was confirmed. In mix infection cases, the morphology of the cutaneous ulcers was not different from the wounds of other patients.Two species Leishmania major is responsible for the most common CL in southern Iran. In addition, in two patients with L. major and L. tropica, mix infection with Crithidia was confirmed. The potential role of Crithidia as the main factor for CL and the probability of this parasite to have synergistic effects on Leishmania, as a hypothesis, requires more comprehensive researches on the ambiguity of this protozoon. Cutaneous leishmaniasis (CL) is one of the six important diseases in tropical and subtropical regions which WHO has recommended and supported researches on its various aspects , 2.Leishmania  are also considered to be infectious agents of the Leishmania culture medium  and informed consent was signed by each of the patients.In this cross-sectional study, in a period of 6 months from Apr to Sep 2016, patients with cutaneous ulcers referred to the Diagnostic Lab of Leishmaniasis, Valfajr Health Center, Shiraz, Iran were tested. Smears of them from exudates of margin of the wounds were prepared and stained with Giemsa. Finally 66 patients with CL were selected. Then a questionnaire containing demographic information and some of the variables such as age, sex, etc. were completed separately for each patient.DNA was extracted from the Giemsa stained smears using the commercial kit . Briefly, the smears on the slides were scratched and collected by lysis buffer into the microtubes and after the addition of proteinase K, incubated initially for 1 h at 60 °C and then 10min at 70 °C. By addition of absolute ethanol and transferring the samples to the column, the rest of the process was carried out according to the kit’s manufacturer’s protocol.L. major, L. infantum and L. tropica, and in the next step, in order to identify Crithidia spp., CRF and CRR primers were used in this purpose. Further details about the sequence of primers used and the PCR programs adjusted for the amplification of both genomic pieces, has been presented in All extracted DNA samples were PCR-amplified in two separate steps. First, a pair of primers LIN4R and LIN17 were used to detect and differentiate of three species -values were considered as statistically significant.The results and data of the questionnaires were analyzed using SPSS software version 18 and chi-square test and P-values. At levels < .05, the PL. major and L. tropica were responsible for 60 and 6 cases, respectively , the morphology of the cutaneous ulcers was not different from the wounds of the other patients.In two patients with mix infection Leishmania spp. (Cutaneous leishmaniasis is one of the vector-borne diseases and is caused by various species of nia spp. , 25. Estnia spp. .The isolation and mass cultivation of parasite in order to determine the species is not only very time-consuming but also is expensive , 27, theL. major, L. infantum and L. tropica create a specific band in the 650, 720 and 760 bases, respectively. The specificity of 100% has been reported for PCR tests using primers LIN4R and LIN17 to detect Leishmania in atypical cases (The success of DNA extraction from these smears has already been studied . Furtheral cases , 30.L. major and in other areas, L. tropica plays a role in causing disease, although, in southern regions such as Fars Province, considered as one of the most important foci in Iran, both species are listed as endemic factors (Crithidia spp. has been reported from cultivation of cutaneous ulcers of patients (In some endemic and hyper-endemic regions of Iran  factors , 31. Recpatients , 16.Crithidia spp. in cutaneous lesions, samples were again amplified with two specific FCR and PCR primers. In two CL cases, there was a mix infection by Leishmania and Crithidia and, as shown in Crithidia spp. are classified in flagellate subphylum and are often considered as insect parasites (Leishmania, they are usually associated with together (Crithidia spp. are considered as infectious agents of the Leishmania culture medium. Crithidia lucilia and C. fasciculata are two important species of this genus (In this study, in order to determine the role of arasites . Since ttogether , 18. CriLeishmania and Crithidia, as well as to study the molecular diversity and the interaction between this parasite and the host (L. major and L. tropica species, Crithidia spp. has also been proven in the wound of patients with CL, and interestingly, even in some of these reports, Crithidia has been reported as the only factor isolated from the wound of mentioned patients. In a study to investigate the polymorphism of two species L. tropica and L. major in the central and desert regions of Iran, Crithidia has been isolated from 6 of 215 and 3 of 125 patients with cutaneous ulcers in Isfahan and Bam, respectively. Molecular analysis of Crithidia-positive samples using BLAST software has shown that their sequence is 97 % similar to C. fasciculata and 90% to C. lucilia (Over the past 20 years, various methods have been developed to determine the different subspecies of the host –34. In rL. major and then L. tropica. Moreover, in two patients with L. major and L. tropica, mix infection with Crithidia was confirmed. And more importantly, the potential role of Crithidia as the main cause of the cutaneous ulcers or the probability to have synergistic effects on Leishmania cutaneous ulcers, as a hypothesis, needs more comprehensive researches on the ambiguity of this protozoon.The main cause of CL in southern Iran is primarily


In the Funding section, one of the grant numbers from the funder Scientific Research from the Ministry of Education, Science, Sports, and Culture of Japan is missing. The correct grant numbers are: Kakenhi 25293124 to K.F., 17K15456 to Y.Y. and 18J20651 to K.M.


Ostertagia leptospicularis draws special attention due to its presence or absence among the same host species in different European countries. Therefore, this paper focuses mainly on the host specificity among nematodes of the subfamily Ostertagiinae. The second aim of this study is to assess the possibility of treating O. leptospicularis as an Ostertagia species complex.Nematodes of the subfamily Ostertagiinae appear to be rather specific to a species or family of hosts, but some are observed in a wide variety of hosts. The nematode post-mortem examinations of domestic and wild ruminants (n = 157), as well as bibliographical references (n = 96), which were pooled and discussed. The research area was limited to European countries, hence the studied ostertagiine species are limited to native ones; likewise, the host species. Special emphasis was placed on the mean abundance values that allowed a typical host or hosts for each nematode species to be specified. Correspondence analysis was performed to confirm the stated host specificity.Data were gathered from Ostertagia leptospicularis, similar to Teladorsagia circumcincta, may represent a potential species complex. This nematode, as the sole member of the subfamily Ostertagiinae, occurs in almost all representatives of the Bovidae subfamily, as well as in the Cervidae.The analysis revealed that nematodes of this subfamily tend to use ruminants from a particular subfamily as their principal host. The results indicate that O. leptospicularis is not strongly connected to any host or is comparably associated with a very wide and diverse group of hosts . The Ostertagia complex may have particular cryptic species or strains typical for any individual host or group of hosts. Such a conclusion requires further investigations on a wider scale.Despite the stated narrow host specificity, the results obtained may suggest that  Parasites are able to colonize one or more host species, and therefore can be classified as specialists, specific for a species or a family of hosts, or generalists, capable of infecting a wide variety of hosts . GeneralTeladorsagia. Results of advanced morphological analyses confirmed by molecular techniques revealed a significant intraspecific diversity in the populations of these nematodes. Until recently, Teladorsagia circumcincta  was treated as a single species that occurs in various hosts (both domestic and wild ruminants). However, some studies on the genus Teladorsagia have demonstrated the existence of two distinct strains within its domestic hosts. The first one was found in sheep and goats, while the other occurred only in goats. In the case of wild ruminants , the nematode regarded previously to be T. circumcincta was discovered to be a distinct species, named Teladorsagia boreoarcticus Hoberg, Monsen, Kutz & Blouin, 1999 to emphasize its origin from a geographically isolated region [Teladorsagia species complex, with particular cryptic species (or strains) [Marshallagia marshalli Ransom, 1907, due to its presence in several host species [Ostertagia; however, such data are not yet available [The ostertagiine taxonomy is complicated, mainly for the species of strains) . Even le species . Similarvailable .Ostertagia leptospicularis Assadov, 1953 especially seems to be a generalist, but such a statement requires broader analysis. From an epizootiological point of view, it is desirable to know which parasite species are more prone to interspecific transmission (i.e. host switching) [Generalist species seem to be able to easily colonize different host populations and pose a serious health risk due to their higher pathogenic effect . Nematoditching) . Such knO. leptospicularis as an Ostertagia species complex.The aim of the present study is to evaluate host specificity among Ostertagiinae, and assess the possibility of treating post-mortem (pm) examination of domestic and wild ruminants from southern Poland, Slovakia and Germany to characterize the parasitic fauna. Only results on the subfamily Ostertagiinae were taken into account. The second step focused on collecting similar data from other European countries and pooling them with those obtained from the pm examinations. The third step was performing the correspondence analysis.Several parasitological approaches were combined for this investigation. The first step was the pm examinations were carried out on 157 animals . The number of animals included in the further analysis is shown in Table Ostertagia leptospicularis / O. kolchida Popova, 1937; O. ostertagi  / O. lyrata Sjoberg, 1926; O. antipini Matschulsku, 1950 / O. lyrateformis ; O. drozdzi Jancev, 1977 / O. ryjikovi ; O. gruehneri Skrjabin, 1920 / O. arctica Mitzkewitsch, 1929; Teladorsagia circumcincta / T. trifurcata ; Spiculopteragia boehmi  / S. mathevosiani Ruchliadev, 1948; S. asymmetrica  / S. quadrispiculata ; S. houdemeri  / S. andreevae ; Mazamastrongylus dagestanica ; M. marshalli / M. occidentalis Ransom, 1907. The majority of the references were gathered using the Web of Science Core Collection citation database.Additional data were collected from 96 bibliographical references , in which at least one species of the subfamily Ostertagiinae derived from domestic or wild ruminants during the pm examinations and the literature were analyzed by pooling the prevalence (P) and all values relating to the intensity of infection . Special emphasis was placed on the mean abundance (MA), i.e. the total number of individuals of a particular parasite species in a sample of a particular host species, regardless of whether or not the host was infected. MA is equivalent to the mean intensity multiplied by P, and therefore is very valuable if biodiversity is considered [The data obtained from the nsidered .In case of ostertagiine species, the identification of males (m) is more reliable; thus, most of the analyzed data applied only to them. However, the total number of females (f) was calculated if the mean intensity of infection and sex ratio were determined. Consequently, it was possible to calculate the actual intensity of females and males in a species for some bibliographical references. As the majority of the papers provided information about the mean intensity of infection, the formula ∑f + ∑m / examined animals or ∑m / examined animals, where ∑f and ∑m are the summed intensity of each sex in a host sample was applied, and the MA was calculated and used in further analyses that allowed the principal host/hosts for each nematode species to be specified. If a major and minor form of one species was included separately, only the major one was counted with the total intensity calculated for both.The correspondence analysis was performed using Statgraphics Centurion XVI software to confirm host specificity. The MA was used as the measure characterizing the association between the rows and columns in the contingency table.pm examinations of 157 animals are shown in Table O. leptospicularis, O. ostertagi, O. antipini, O. drozdzi, T. circumcincta, S. asymmetrica, S. boehmi and M. marshalli. Four of the species (approximately 4050 specimens) were derived from roe deer. The majority of nematode species were found in more than one host species; however, S. antipini and M. marshalli only occurred in roe deer and Tatra chamois, respectively. Spiculopteragia boehmi was the most common parasite species in the Cervidae family, whereas T. circumcincta was the most common in the Bovidae. The prevalence varied from 1.3% (S. asymmetrica infecting roe deer) to 100% . The highest intensity of infection (reaching 829 specimens) with O. ostertagi was in cattle, and the lowest (just 3 specimens) with M. marshalli in Tatra chamois. Interestingly, O. leptospicularis was the only species that was found in both the representatives of the Bovidae (i.e. cattle and sheep) and Cervidae (i.e. roe deer).The results obtained from the pm examinations in 17 countries , whereas Cervidae in 16 countries . The most abundant data were available for roe deer, while those for European bison, Alpine ibex, Pyrenean ibex and reindeer were limited due to their narrow range of occurrence.The comparison of bibliographical data collected showed the differences in the occurrence of ostertagiine species among European ruminant host species Table . AccordiO. drozdzi, S. houdemeri and M. dagestanica are observed only in cervids. Within the countries studied, the most commonly occurring species were O. leptospicularis and T. circumcincta, found in 12 and 13 host species, respectively. Conversely, O. drozdzi and O. gruehneri were found only in two.The parasitic fauna of European cervids consists of eleven species of the Ostertagiinae, and therefore can be considered as more diverse than that of the bovids (eight species); O. leptospicularis was found in almost all representatives of the family Cervidae, with most reports concerning roe deer. Wild and domestic ruminant hosts from Poland and Germany had very similar parasitic faunas. Nevertheless, they varied in some findings, including those referring to O. leptospicularis. The majority of the differences presented concerned the composition of ruminant fauna in both countries . Despite the broad examination of cattle in both countries, O. leptospicularis was noted only in Germany . Lack of this nematode in Polish cattle, or in cattle from other European countries, infers that more attention should be dedicated to this species.In many countries, Both the correspondence map Fig. , as wellM. marshalli and T. circumcincta are most strongly associated with small ruminants (i.e. species of the subfamily Caprinae). This close connection is also observed between O. ostertagi and other bovids (i.e. those of the subfamily Bovinae). However, cattle seem to be a more typical host (the distance between the cross and square is shorter for O. ostertagi and cattle than for O. ostertagi and the European bison). Nematodes of the genus Spiculopteragia, as well as O. drozdzi, show the strongest link to Cervinae representatives, while O. gruehneri, O. antipini and M. dagestanica to those of the subfamily Capreolinae .The correspondence analysis divided the Ostertagiinae and their hosts into clearly distinct groups Fig. . Based oO. antipini and moose, S. houdemeri and sika deer, as well as O. drozdzi, S. asymmetrica and fallow deer . Such data do not provide a clear overview of host specificity. According to these outcomes, members of the subfamilies Bovidae and Cervidae have different helminth fauna, characterized by specific dominant species, which depends on several factors .Some ruminant species have already been treated as the principal host of Ostertagiinae nematodes, but only on the basis of observations of T. circumcincta, O. ostertagi and M. marshalli. The first two are the most important gastrointestinal parasites of domestic ruminants [Teladorsagia circumcincta is the dominant species in all bovids, while M. marshalli seems to be almost entirely restricted to hosts that are well adapted to a cold climate (i.e. representatives of chamois and ibex) [O. leptospicularis and S. boehmi from roe and red deer and in this way enriched the parasitic fauna of the abomasal nematodes with those typical for representatives of the family Cervidae [Among bovids, the most frequently found nematodes are uminants –17. TelaCervidae , 19.Spiculopteragia are generally restricted to cervids, although records of infected bovids in Europe have been documented [S. asymmterica; however, it can be found in sympatric species, such as red deer, roe deer or even European bison if they co-occur with the principal host [O. antipini, M. dagestanica and the largest species of Cervidae, moose, which has currently arisen as its principal host [Cross-transmission is highly visible among cervids, and therefore these hosts are successfully used for studies on formatting the parasitic fauna of ruminants. All representatives of the family Cervidae are the principal hosts of particular ostertagiine species. The origin of such a relationship is dated to over ten million years ago, when the cervids of the subfamily Paleomericinae divided into two lineages, and since then, nematodes and their hosts evolved in parallel. A high degree of co-evolution is also seen among bovids , 20. Thecumented . Fallow pal host . The sampal host . These npal host , 24, 25.Representatives of the family Cervidae  migrate seasonally, which encourage the possibility of parasite transmission not only to other wild ruminants, but also to livestock , 26. ThiOstertagia leptospicularis circulates among cervids, cattle and sheep. This species is not very host-specific; nevertheless, it seems to have mostly adopted roe deer [O. leptospicularis was noted as having become an outstanding and unique species. The equal distance from O. leptospicularis to roe deer and European bison on the correspondence map may suggest that this nematode is not strongly connected to any of the hosts, or that it may be comparably associated with very a wide and diverse groups of hosts.roe deer , 16. BasO. leptospicularis. Ostertagia leptospicularis seems to have a principal host among members of both subfamilies the Bovidae and Cervidae. If the confirmed host specificity can be considered an indication of the species diversity among the Ostertagiinae in ruminants, then O. leptospicularis could be treated as a candidate for an Ostertagia species complex. Further investigations  should firmly establish if, as with T. circumcincta, this nematode represents another species complex with particular cryptic species/strains typical for any individual host/group of hosts, or, as the sole member of the subfamily Ostertagiinae, is a generalist, capable of infecting a wide variety of hosts.The analyses conducted confirmed that nematodes of the Ostertagiinae appear to be specific for a species or family of hosts (i.e. ostertagiines of a particular genus are most strongly connected to hosts from a particular subfamily). For the majority of them, a principal host can be indicated, except for