Musculature of the penial complex: A new criterion in unravelling the phylogeny of Hygrophila (Gastropoda: Pulmonata)

The taxonomy of freshwater pulmonates (Hygrophila) has been in a fluid state warranting the search for new morphological criteria that may show congruence with molecular phylogenetic data. We examined the muscle arrangement in the penial complex (penis and penis sheath) of most major groups of freshwater pulmonates to explore to which extent the copulatory musculature can serve as a source of phylogenetic information for Hygrophila. The penises of Acroloxus lacustris (Acroloxidae), Radix auricularia (Lymnaeidae), and Physella acuta (Physidae) posses inner and outer layers of circular muscles and an intermediate layer of longitudinal muscles. The inner and outer muscle layers in the penis of Biomphalaria glabrata consist of circular muscles, but this species has two intermediate longitudinal layers separated by a lacunar space, which is crossed by radial and transverse fibers. The muscular wall of the penis of Planorbella duryi is composed of transverse and longitudinal fibers, with circular muscles as the outer layer. In Planorbidae, the penial musculature consists of inner and outer layers of longitudinal muscles and an intermediate layer of radial muscles. The penis sheath shows more variation in muscle patterns: its muscular wall has two layers in A. lacustris, P. acuta, and P. duryi, three layers in R. auricularia and Planorbinae and four layers in B. glabrata. To trace the evolution of the penial musculature, we mapped the muscle characters on a molecular phylogeny constructed from the concatenated 18S and mtCOI data set. The most convincing synapomorphies were found for Planorbinae (inner and outer penis layers of longitudinal muscles, three-layered wall of the penis sheath). A larger clade coinciding with Planorbidae is defined by the presence of radial muscles and two longitudinal layers in the penis. The comparative analysis of the penial musculature appears to be a promising tool in unraveling the phylogeny of Hygrophila.     

Histology and Ultrastructure of the Body Wall in the Phoronid Phoronopsis harmeri

The histology and ultrastructure of the body wall in Phoronopsis harmeriwere studied using light microscopy and TEM. The ectoderm epithelium of tentacles, anterior body region, and ampulla consists of monociliary cells. Gram-negative bacteria were found between microvilli, in the protocuticle of the anterior region, and in the ampulla. The epithelium of the posterior body region lacks both monociliary cells and bacteria. The bundles of nerve fibers run between the layer of epithelial cells and basal membrane. The musculature of the body wall comprises circular and longitudinal muscles. The circular muscle fibers are applied to the basal membrane and constitute a solid layer extending almost throughout the length of the body. This pattern is broken in the posterior body region, where there is no solid layer of circular musculature, and the latter is arranged in isolated muscle bands. In the ampullar (terminal) body region, the inversion of circular and longitudinal muscle layers takes place, so that the latter appears to be pressed against the basal membrane. The apical surfaces of longitudinal muscle cells bear cytoplasmic processes; some of the cells have a flagellum. The basal portion of the longitudinal muscle cells forms a cytoplasmic process containing bundles of tonofilaments. The processes of all cells making up the muscle bands are interwoven and anchored to the basal membrane.     

Paralineopsis taki gen. et sp. nov., a littoral heteronemertean from Japan,provided with special proboscideal,circulatory and sensory organs of significance to nemertean systematics

Some morphological features with major systematic significance are recorded in the heteronemertean Paralineopsis taki gen. et sp. nov. from Onomichi, Japan as follows: horizontal band of specialized epithelium extends from near apex to the opening of the cerebral organ canal on either side of the head; precerebral region filled with gelatinous (hyaline) connective tissue in which longitudinal muscles are absent; body wall muscles do not accompany rhynchodaeal invagination; rhynchodaeum initially only epidermal; inner longitudinal muscles of ventral wall of cephalic blood lacuna become intimately associated with rhynchodaeum forming a dorsal saddle over it; cerebral organs do not penetrate inner longitudinal muscles, and do not contact blood vascular system; proboscideal diaphragm post-cerebral; outer longitudinal muscles absent throughout body; longitudinal muscles of proboscis derived from inner longitudinal musculature. The systematic relationship of P. taki and Paralineus elisabethae (Schütz, 1911) from Villefranche, France is discussed.     

A new genus and species of heteronemertean from the Changjiang (Yangtze) River Estuary

A new genus and species of heteronemertean, Yinia pratensis gen. nov. and sp. nov., collected from low salinity waters (salinity 0.2–0.4 ‰) at Changjiang River Estuary, is described and illustrated. The species possesses a proboscis with an outer circular and an inner longitudinal muscle layer, and is placed in family Lineidae sensu Gibson. The following combination of morphological features distinguishes the new species from any other genera in this family: proboscis with two muscle crosses; dermis without connective tissue layer between gland cells and body wall outer longitudinal muscle layer; rhynchocoel wall circular muscles not interweaving with adjacent body wall longitudinal muscles; foregut with circular somatic muscles and subepithelial gland cell layer; neurochord cells present in central nervous system; caudal cirrus missing; blood system developed into alimentary plexus extending almost the full length of the body. Another significant character is that the lobular excretory cells are extremely well developed which may represent adaptation to water of low salinity. This revised version was published online in July 2006 with corrections to the Cover Date.     

Three-dimensional reconstruction of the musculature of Cossura pygodactylata Jones, 1956 (Annelida: Cossuridae)

The musculature of adult specimens of Cossura pygodactylata was studied by means of F-actin labelling and confocal laser scanning microscopy (CLSM). Their body wall is comprised of five longitudinal muscle bands: two dorsal, two ventral and one ventromedial. Complete circular fibres are found only in the abdominal region, and they are developed only on the border of the segments. Thoracic and posterior body regions contain only transverse fibres ending near the ventral longitudinal bands. Almost-complete rings of transverse muscles, with gaps on the dorsal and ventral sides, surround the terminal part of the pygidium. Four longitudinal bands go to the middle of the prostomium and 5–14 paired dorso-ventral muscle fibres arise in its distal part. Each buccal tentacle contains one thick and two thin longitudinal muscle filaments; thick muscle fibres from all tentacles merge, forming left and right tentacle protractors rooted in the dorsal longitudinal bands of the body wall. The circumbuccal complex includes well-developed upper and lower lips. These lips contain an outer layer of transverse fibres, and the lower lip also contains inner oblique muscles going to the dorsal longitudinal bands. The branchial filament contains two longitudinal muscle fibres that do not connect with the body musculature. The parapodial complex includes strong intersegmental and segmental oblique muscles in the thoracic region only; chaetal retractors, protractors and muscles of the body wall are present in all body regions. Muscle fibres are developed in the dorsal and ventral mesenteries. One semi-circular fibre is developed on the border of each segment and is most likely embedded in the dissepiment. The intestine has thin circular fibres along its full length. The dorsal blood vessel has strong muscle fibres that cover its anterior part, which is called the heart. It consists of short longitudinal elements forming regular rings and inner partitions. The musculature of C. pygodactylata includes some elements that are homologous with similar muscular components in other polychaetes (i.e., the body wall and most parapodial muscles) and several unique features, mostly at the anterior end.     

The neuroactive substances and associated muscle system in Rhipidocotyle campanula (Digenea,Bucephalidae) from the intestine of the pike Esox lucius

We report about the muscular system and the serotonergic and FMRFamidergic components of the nervous system of the Bucephalidae trematode, Rhipidocotyle campanula, an intestinal parasite of the pike. We use immunocytochemical methods and confocal scanning laser microscopy (CLSM). The musculature is identified by histochemical staining with fluorescently labeled phalloidin. The body wall musculature of R. campanula contains three layers of muscle fibres – the outer thin circular, intermediate longitudinal and inner diagonal muscle fibres running in two opposite directions. The digestive system of R. campanula possess of a well-developed musculature: radial, longitudinal and circular muscle elements are detected in the pharynx, circular and longitudinal muscle filaments seen in the oesophagus, and longitudinal and the circular muscle fibres were found in the intestinal wall. Specific staining indicating the presence of actin muscle filaments occurs in the cirrus sac localized in the posterior body region. The frontal region of anterior attachment organ, the rhynchus, in R. campanula is represented by radial muscle fibres. The posterior part of the rhynchus comprise of radial muscles forming the organ's wall, and several strong longitudinal muscle bundles. Serotonergic and FMRFamidergic structures are detected in the central and peripheral compartments of the nervous system of R. campanula, that is, in the paired brain ganglia, the brain commissure, the longitudinal nerve cords, and connective nerve commissures. The innervations of the rhynchus, pharynx, oesophagus and distal regions of the reproductive system by the serotonergic and FMRFamidergic nervous elements are revealed. We compare our findings obtained on R. campanula with related data for other trematodes.     

Anatomy of the planarian Dugesia japonica I. The muscular system revealed by antisera against myosin heavy chains

The planarian Dugesia japonica has two genes encoding myosin heavy chain, DjMHC-A and B (Kobayashi et al., 1998). We produced antibodies specifically recognizing each myosin heavy chain protein using their carboxyl terminal regions expressed in E. coli as antigens. Immunohistochemical analyses of sections and whole-mount specimens revealed the detailed structure and distribution of each type of muscle fiber in the planarian. In general, the MHC-A muscle fibers were distributed beneath the epithelial layers, namely, they were observable in the pharynx, the mouth, the intestine, the eyes and the body wall. In the pharynx, only MHC-A muscle fibers were present. In contrast, the MHC-B muscle fibers were distributed in the mesenchyme as dorso-ventral and transverse muscles, and in the body wall. The body-wall muscles were composed of an outer layer of circular MHC-A muscles and inner longitudinal and intermediate diagonal MHC-B muscle layers. Thus, two types of muscle fibers were distinguished by their distribution in the planarian.     

免疫荧光法显示贝居腹盾吸虫肌肉神经组织的实验研究

从背角无齿蚌(Anodonta woodiana)围心腔内检获自然感染的贝居腹盾吸虫(Aspidogaster conchicola),根据虫体内睾丸、卵巢、子宫及卵黄腺4种生殖器官的发育程度,将所获虫体分为幼虫组和成虫组,所有虫体用4%甲醛固定后,分别采用鬼笔环肽染色法和乙酰化微管蛋白免疫组织化学法显示贝居腹盾吸虫成虫和幼虫的肌肉纤维和神经纤维,使用激光共聚焦显微镜观察。结果显示,鬼笔环肽荧光染色的贝居腹盾吸虫成虫较幼虫肌肉组织更为发达,口周围、盾盘附着器、子宫末段及阴茎囊肌肉纤维分布较为密集。乙酰化微管蛋白荧光染色的贝居腹盾吸虫成虫较幼虫神经网络更为复杂,在部分生殖器官如子宫末段、阴茎囊和睾丸部位存在神经纤维。双重染色结果显示,该吸虫不同器官上的肌肉纤维分布与神经纤维的分布相互吻合。该结果提示,贝居腹盾吸虫可能通过神经系统对其肌肉活动进行调控。     

Muscular system in polychaetes (Annelida)

The structure of the polychaete muscular system is reviewed. The muscular system comprises the muscles of the body wall, the musculature of the parapodial complex and the muscle system of the dissepiments and mesenteries. Various types of organisation of the longitudinal and circular components of the muscular body wall are distinguished. In Opheliidae, Polygordiidae, Protodrilidae, Spionidae, Oweniidae, Aphroditidae, Acoetidae (=Polyodontidae), Polynoidae, Sigalonidae, Phyllodocidae, Nephtyidae, Pisionidae, and Nerillidae circular muscles are lacking. It is hypothesised that the absence of circular muscles represents the plesiomorphic state in Annelida. This view contradicts the widely accepted idea of an earthworm-like musculature of the body wall comprising an outer layer of circular and an inner layer of longitudinal fibres. A classification of the various types of parapodial muscle construction has been developed. Massive and less manoeuvrable parapodia composed of many components like those of Aphrodita are regarded to represent the plesiomorphic state in recent polychaetes. An analysis of the diversity of the muscular structure supports the hypothesis that the primary mode of life in polychaetes was epibenthic and the parapodial chaetae had a protective function.     

Functional morphology of the muscles in Philodina sp. (Rotifera: Bdelloidea)

Whole-mounts of Philodina sp., a bdelloid rotifer, were stained with fluorescent-labeled phalloidin to visualize the musculature. Several different muscle types were identified including incomplete circular bands, coronal retractors and foot retractors. Based on the position of the larger muscle bands in the body wall, their function during creeping locomotion and tun formation was inferred. Bdelloid creeping begins with the contraction of incomplete circular muscle bands against the hydrostatic pseudocoel, resulting in an anterior elongation of the body. One or more sets of ventral longitudinal muscles then contract bringing the rostrum into contact with the substrate, where it presumably attaches via adhesive glands. Different sets of ventral longitudinal muscles, foot and trunk retractors, function to pull the body forward. These same longitudinal muscle sets are also used in `tun' formation, in which the head and foot are withdrawn into the body. Three sets of longitudinal muscles supply the head region (anterior head segments) and function in withdrawal of the corona and rostrum. Two additional pairs of longitudinal muscles function to retract the anterior trunk segments immediately behind the head, and approximately five sets of longitudinal retractors are involved in the withdrawal of the foot and posterior toes. To achieve a greater understanding of rotifer behavior, it is important to elucidate the structural complexity of body wall muscles in rotifers. The utility of fluorescently-labeled phalloidin for the visualization of these muscles is discussed and placed in the context of rotifer functional morphology.     

Functional morphology of musculature in the acoelomate worm, Convoluta pulchra (Plathelminthes)

Convoluta pulchra is a small worm living in the surface sediment of mud flats where it feeds on diatoms. It is roughly teardrop in shape with a ventral groove in which the mouth sits, and it can move in a variety of ways, readily distorting its body in bending, twisting, and turning motions. Fluorescently labeled probes for filamentous actin revealed the musculature in whole mounts of the worm. In the body wall, the musculature consisted of a grid of circular, longitudinal crossover (that is, with a longitudinal orientation in the anterior half of the body but arcing medially to cross over to the contralateral side of the body behind the level of the mouth), and a few diagonal fibers. Inside the body was a strong, irregular brush of muscles originating at the rostral tip of the body and anchoring laterally and posteriorly along the body wall, and strong dorsoventral muscles flanked the ventral groove. Two fans of muscles in the ventral and dorsal body wall reached posteriorly and laterally; that on the dorsal side originated at junctures of the dorsoventral muscles with the body wall and that on the ventral body wall originated from the mouth. By their positions, certain groups of muscles could be correlated with given movements: the crossover muscles with some turning motions and feeding, and the inner muscles with probing and retraction motions of the rostrum and with a tuck-and-turn motion the worm used to turn itself around. Electron microscopy showed numerous maculae adherentes junctions linking all muscle types and special junctions linking the musculature with the epidermis. The latter myoepidermal junctions were of dimensions larger than those of maculae adherentes and contained an interlaminar material which we believe represents islands of basal matrix comparable to basement membrane. Accepted: 12 July 1999     

On the absence of circular muscle elements in the body wall of Dysponetus pygmaeus (Chrysopetalidae, 'Polychaeta', Annelida)

The annelid body wall generally comprises an outer layer of circular muscle fibres and an inner layer of longitudinal muscle fibres as well as parapodial and chaetal muscles. An investigation of Dysponetuspygmaeus (Chrysopetalidae) with confocal laser scanning microscopy showed that circular muscles are entirely absent. Further studies indicate that this feature is characteristic for all Chrysopetalidae. A scrutiny of the literature showed a similar situation in many other polychaetes. This lack of circular muscle fibres may either be due to convergence or represent a plesiomorphic character. Since circular muscles are very likely important for burrowing forms but not necessary for animals which proceed by movements of their parapodial appendages or cilia, this problem is also related to the question of whether the ancestral polychaete was epi‐ or endobenthic.     

Relating divergence in polychaete musculature to different burrowing behaviors: A study using opheliidae (Annelida)

Divergent morphologies among related species are often correlated with distinct behaviors and habitat uses. Considerable morphological and behavioral differences are found between two major clades within the polychaete family Opheliidae. For instance, Thoracophelia mucronata burrows by peristalsis, whereas Armandia brevis exhibits undulatory burrowing. We investigate the anatomical differences that allow for these distinct burrowing behaviors, then interpret these differences in an evolutionary context using broader phylogenetic (DNA‐based) and morphological analyses of Opheliidae and taxa, such as Scalibregmatidae and Polygordiidae. Histological three‐dimensional‐reconstruction of A. brevis reveals bilateral longitudinal muscle bands as the prominent musculature of the body. Circular muscles are absent; instead oblique muscles act with unilateral contraction of longitudinal muscles to bend the body during undulation. The angle of helical fibers in the cuticle is consistent with the fibers supporting turgidity of the body rather than resisting radial expansion from longitudinal muscle contraction. Circular muscles are present in the anterior of T. mucronata, and they branch away from the body wall to form oblique muscles. Helical fibers in the cuticle are more axially oriented than those in undulatory burrowers, facilitating radial expansion during peristalsis. A transition in musculature accompanies the change in external morphology from the thorax to the abdomen, which has oblique muscles similar to A. brevis. Muscles in the muscular septum, which extends posteriorly to form the injector organ, act in synchrony with the body wall musculature during peristalsis: they contract to push fluid anteriorly and expand the head region following a direct peristaltic wave of the body wall muscles. The septum of A. brevis is much thinner and is presumably used for eversion of a nonmuscular pharynx. Mapping of morphological characters onto the molecular‐based phylogeny shows close links between musculature and behavior, but less correlation with habitat. J. Morphol. 275:548–571, 2014. © 2014 Wiley Periodicals, Inc.     

贵州松桃寒武系清虚洞组Mongolitubulus squamifer 壳体显微结构初步研究

本文主要对贵州松桃寒武系清虚洞组灰岩中酸泡获得的管状化石Mongolitubulus squamifer壳体显微及亚显微结构进行研究。研究结果显示, M. squamifer不仅内层广泛发育纵向纤维结构,具鳞片的外层外壁也发育有微弱的纵向纤维结构,可能解释了管体外壁广泛发育的纵向开裂现象。在一枚保存有圆卵形鳞片以下部位的标本中发现,鳞片排列形式是由基部密集的小瘤点状颗粒然后过渡为不规则的圆卵形鳞片,到管体中上部逐渐形成规则排列的三角形鳞片。通过测量部分已报道的M.squamifer管体宽度与鳞片宽度发现正常发育的棘刺,鳞片大小与管体宽度有一定相关性。贵州松桃的M.squamifer壳层原始结构为2层,包括致密的具鳞片的外层和具明显纵向纤维结构的内层,管体内外层之间发育空隙导致管体容易破损。一些标本近基部断口处的层间空隙容易被次生矿物充填,导致内层增厚致密纤维结构消失或产生中间填充层。该研究揭示了M.squamifer的鳞片排列特征和壳体微观结构,为解释其亲缘关系提供新的形态学证据。     

Ultrastructure of the subepidermal musculature of Xenoturbella bocki, the adelphotaxon of the Bilateria

 Xenoturbella bocki is the only species of the high-ranked taxon Xenoturbellida. The species lives on marine mud bottoms at a depth of 20–120 m and moves extremely slowly by ciliary gliding. Nevertheless it possesses a well-developed body wall musculature with outer circular muscles, a prominent layer of inner longitudinal muscles and radial muscles that extend from the outer circular myocytes to the musculature surrounding the gastrodermis. The longitudinal myocytes are not compact cells, but form fascicles of fibrils running parallel to each other. Fine cytoplasmic cords connect the fibres of a cell to each other and with its nuclear region. The muscles are embedded within a sometimes expansive extracellular matrix (ECM) that lacks any fibrillar components. All muscle cells display conspicuous and numerous cytoplasmic extensions that are intermingled with each other. Tight coupling between adjacent cell membranes is not found, but zonula adhaerens-like junctions exist. Fibrils belonging to different myocytes, but also fibrils of the same cell, are coupled by such cytoplasmic extensions. Circular, radial and at least the peripheral longitudinal myocytes display cell-matrix connections with the internal lamina, a component of the subepidermal ECM. This internal lamina projects down into the centres of the fascicles with longitudinal muscle fibrils and forms extensive attachment zones with the muscle cells, reminiscent of focal contacts. For the ingestion of food, X. bocki opens the simple mouth pore and protrudes the aciliated gastrodermis. The body wall musculature is responsible for this protrusion and also for the withdrawal of the gastrodermis. In the past, possible phylogenetic kinships with the Acoelomorpha (Plathelminthes) or the Enteropneusta and Holothuroidea were discussed, but, on the basis of all information available, X. bocki is hypothesized to be the sister taxon of the Bilateria. Accepted: 2 April 1997     

The pregenital abdominal musculature in phasmids and its implications for the basal phylogeny of Phasmatodea (Insecta: Polyneoptera)

Recently several conflicting hypotheses concerning the basal phylogenetic relationships within the Phasmatodea (stick and leaf insects) have emerged. In previous studies, musculature of the abdomen proved to be quite informative for identifying basal taxa among Phasmatodea and led to conclusions regarding the basal splitting events within the group. However, this character complex was not studied thoroughly for a representative number of species, and usually muscle innervation was omitted. In the present study the musculature and nerve topography of mid-abdominal segments in both sexes of seven phasmid species are described and compared in detail for the first time including all putative basal taxa, e.g. members of Timema, Agathemera, Phylliinae, Aschiphasmatinae and Heteropteryginae. The ground pattern of the muscle and nerve arrangement of mid-abdominal segments, i.e. of those not modified due to association with the thorax or genitalia, is reconstructed. In Timema, the inner ventral longitudinal muscles are present, whereas they are lost in all remaining Phasmatodea (Euphasmatodea). The ventral longitudinal muscles in the abdomen of Agathemera, which span the whole length of each segment, do not represent the plesiomorphic condition as previously assumed, but might be a result of secondary elongation of the external ventral longitudinal muscles. Sexual dimorphism, common within the Phasmatodea, also applies to the muscle arrangement in the abdomen of some species. Only in the females of Haaniella dehaanii (Heteropteryginae) and Phyllium celebicum (Phylliinae) the ventral external longitudinal muscles are elongated and span the length of the whole segment, possibly as a result of convergent evolution.     

The musculature of horsehair worm larvae (<Emphasis Type="Italic">Gordius aquaticus</Emphasis>,<Emphasis Type="Italic"> Paragordius varius</Emphasis>, Nematomorpha): F-actin staining and reconstruction by cLSM and TEM

The musculature of larvae of Gordius aquaticus was investigated by laser-scanning microscopy and compared to transmission electron microscopic data for the larva of Paragordius varius. In the anterior portion of the body, the preseptum, four different muscle groups can be distinguished: (1) 12 anterior parietal muscles in the body wall, (2) six oblique muscles that function as retractors of the introvert, (3) six proboscideal muscles, which function as retractors for the proboscis, and (4) six muscles associated with spines of the outermost of the three rings of spines. The posterior portion of the body, the postseptum, possesses four pairs of longitudinal muscle strands in G. aquaticus, the postseptal parietal muscles, that are located dorsolaterally and ventrolaterally. These are not clearly visible in P. varius, where instead three pairs of parietal muscles are present. Additional small muscles are associated with the terminal spines and with the duct running from the pseudointestine to the body wall. All fibers show a cross-striated pattern although this striation is less obvious at the ends of the fibers.     

Body wall organization in enchytraeids

The muscle organization of the body wall in some species of oligochaetes belonging to the Enchytraeus genus is described. No differences have been detected in their circular muscles, whereas longitudinal muscles show significant differences, allowing an easy identification of the various worm species. In particular, differences are noticeable in the external longitudinal layer. These observations suggest that structural and ultrastructural muscle fiber organizations can be used as a taxonomic tool.     

Body wall muscles in the haplotaxidsHaplotaxis gordioides andPelodrilus leruthi (Annelida,Oligochaeta)

Summary The muscle organization of the body wall in two species of Haplotaxida is described.Haplotaxis gordioides andPelodrilus leruthi show significant differences in the longitudinal muscle layer. The observations suggest that inH. gordioides, the muscle fibres could foreshadow the flat circomyarian organization of microdrile muscles, while theP. leruthi organization recalls the pinnate arrangement of megadrile body wall.     

Onychophoran‐like myoanatomy of the Cambrian gilled lobopodian Pambdelurion whittingtoni

Arthropods are characterized by a rigid, articulating, exoskeleton operated by a lever‐like system of segmentally arranged, antagonistic muscles. This skeletomuscular system evolved from an unsegmented body wall musculature acting on a hydrostatic skeleton, similar to that of the arthropods’ close relatives, the soft‐bodied onychophorans. Unfortunately, fossil evidence documenting this transition is scarce. Exceptionally‐preserved panarthropods from the Cambrian Lagerstätte of Sirius Passet, Greenland, including the soft‐bodied stem‐arthropod Pambdelurion whittingtoni and the hard‐bodied arthropods Kiisortoqia soperi and Campanamuta mantonae, are unique in preserving extensive musculature. Here we show that Pambdelurion's myoanatomy conforms closely to that of extant onychophorans, with unsegmented dorsal, ventral and longitudinal muscle groups in the trunk, and extrinsic and intrinsic muscles controlling the legs. Pambdelurion also possesses oblique musculature, which has previously been interpreted as an arthropodan characteristic. However, this oblique musculature appears to be confined to the cephalic region and first few body segments, and does not represent a shift towards arthropodan myoanatomy. The Sirius Passet arthropods, Kiisortoqia and Campanamuta, also possess large longitudinal muscles in the trunk, although, unlike Pambdelurion, they are segmentally divided at the tergal boundaries. Thus, the transition towards an arthropodan myoanatomy from a lobopodian ancestor probably involved the division of the peripheral longitudinal muscle into segmented units.