Independent evolutionary reductions of the phallus in basal birds

Despite a long history of anatomical studies in birds, the genitalia of most avian species remain undescribed. Birds are the only vertebrate taxon with internal fertilization where an intromittent phallus has been lost in most species. Studying the anatomical transitions of the avian phallus in those species where it is still present, allows us to test evolutionary hypotheses of why the phallus was lost in the ancestor of modern birds. As part of an anatomical survey of the evolution of avian phallus morphology, we have examined some avian species whose genitalia have not been described. Previously, there were only two known events of phallus reduction in birds: one transition from intromittent to non‐intromittent in the Galliformes, and a complete loss of phallic structures in the ancestor of Neoaves. Here we report three additional cases of phallus reduction in birds: a transition from intromittent to non‐intromittent phallus in Tinamiformes (Crypturellus, Tinamidae), the presence of a non‐intromittent phallus in Alectura (Megapodidae), and a complete loss of the phallus in Leipoa (Megapodidae). In addition, we report on the unique morphology of the Crypturellus non‐intromittent phallus. These new records of phallus reduction highlight the dynamic nature of phallus evolution in birds. Our findings provide evidence against the hypothesis that the phallus in birds is maintained to insure paternity in taxa with exclusive male parental care, since both groups where we report phallus reduction provide predominately male‐only care.     

Resurrecting embryos of the tuatara,Sphenodon punctatus,to resolve vertebrate phallus evolution

The breadth of anatomical and functional diversity among amniote external genitalia has led to uncertainty about the evolutionary origins of the phallus. In several lineages, including the tuatara, Sphenodon punctatus, adults lack an intromittent phallus, raising the possibility that the amniote ancestor lacked external genitalia and reproduced using cloacal apposition. Accordingly, a phallus may have evolved multiple times in amniotes. However, similarities in development across amniote external genitalia suggest that the phallus may have a single evolutionary origin. To resolve the evolutionary history of amniote genitalia, we performed three-dimensional reconstruction of Victorian era tuatara embryos to look for embryological evidence of external genital initiation. Despite the absence of an intromittent phallus in adult tuataras, our observations show that tuatara embryos develop genital anlagen. This illustrates that there is a conserved developmental stage of external genital development among all amniotes and suggests a single evolutionary origin of amniote external genitalia.     

Histological and carbohydrate histochemical studies of the nasal mucosa of sheep in Saudi Arabia with special reference to its glandular tissue

The histology and carbohydrate histochemistry of the nasal mucosa with attention to glandular tissue had been studied in 7 heads of sheep. Tissues were taken from vestibular region, septum at level of the alar fold, rostral portion of nasal conchae, caudal portion of nasal conchae, middle portion of septum and ethmoidal conchae region. Stratified squamous nonkeratinized epithelium was observed covering the vestibular region. The propria-submucosa of the nasal vestibule was richly permeated with glands having affinity for PAS and non-alcianophilic. The post-vestibular portion of the nasal cavity was lined by transitional epithelium and caudal to it, stratified columnar nonciliated epithelium was noticed. The respiratory epithelium covered the caudal half of the nasal conchae and the major portion of the septum as well as the recesses of the ethmoidal conchae. The glands associated with the respiratory mucosa were thick, coiled and tubular, containing both nonalcianophilic PAS positive and alcianophilic PAS positive cells. The olfactory mucosa covered the ethmoidal conchae and showed predominant serous glands. The results were discussed with that given for other mammals and in regard to the respiratory functions of the nasal mucosa.     

Histochemical and ultrastructural study of the digestive tract of the tortoise Testudo graeca (Testudines)

The digestive tract of the tortoise Testudo graeca (Testudines) was investigated by means of light and electron microscopy. The esophagus of T. graeca was lined by two types of epithelium: non-keratinized stratified squamous in the upper portion and stratified columnar in the lower. The lamina propria of the esophagus contained tubular or tubuloacinar glands. The mucosa of the stomach showed similar characteristics to those of other reptiles. The small intestine exhibited longitudinal folds lined by absorptive and goblet cells. The large intestine was lined by columnar mucous cells. Within the lamina propria of the large intestine, masses of 10–15 epithelial cells connecting with the surface epithelium by means of slender cytoplasmic processes were observed. A battery of six lectins (Con-A, PNA, WGA, DBA, SBA, and LTA) was used to identify the epithelial mucins. WGA and DBA reacted with all types of mucous cells throughout the alimentary canal. PNA was only unreactive in the intestine, LTA in the esophagus, and SBA in the large intestine. These results indicate a similar lectin-binding pattern throughout the gut of T. graeca.     

Histological and ultrastructural study of the digestive tract of rice field eel, Monopterus albus

The histological characteristics of the digestive tract and the ultrastructure of mucosal cells of the stomach and intestine of rice field eel, Monopterus albus, are described to provide a basis for future studies on its digestive physiology. The digestive tract of the rice field eel is a long and coiled tube composed of four layers: mucosa, lamina propria‐submucosa, muscularis and serosa. The pharynx and oesophagus mucosa is lined with a stratified epithelium. The stomach includes the cardiac and pyloric portions and the fundus. Many gastric pits are formed by invaginations of the mucosal layer and tubular gastric glands formed by the columnar cells in the fundus. The intestine is separated from the stomach by a loop valve and divided into a proximal portion and a distal portion. The proximal intestinal epithelium consists of columnar cells with microvilli towards the lumen and goblet cells. The enterocytes are joined at the apical surface by the junctional complex, including the evident desmosomas. Numerous lysosomes and some vesicles are evident in the upper cytoplasm of the cells, and a moderate amount of endoplasmic reticulum and lysosomes are scattered in the supranuclear cytoplasm. The epithelium becomes progressively thicker and the folds containing large numbers of goblet cells are fewer and shorter in the distal portion of the intestine. At the ultrastuctural level, the columnar cells of the tubular gastric glands have numerous clear vacuoles and channels. A moderate amount of pepsinogen granules are present in the stomach. The enterocytes of the intestinal mucosa display a moderate amount of endoplasmic reticulum and lysosomes, and long and regular microvilli.     

A study of morphology and histology of the alimentary tract of Glyptosternum maculatum (Sisoridae,Siluriformes)

Xiong, D., Zhang, L., Yu, H., Xie, C., Kong, Y., Zeng, Y., Huo, B. and Liu, Z. 2011. A study of morphology and histology of the alimentary tract of Glyptosternum maculatum (Sisoridae, Siluriformes). —Acta Zoologica (Stockholm) 92 : 161–169. The structure of alimentary tract has been studied in a cold water fish Glyptosternum maculatum, an endemic teleost species of notable economic importance and with high potential for controlled rearing of the species in Tibet, by light and electron microscope. Glyptosternum maculatum has short oesophagus, large caecal‐type stomach and short intestine, and the digestive tract with four layers: mucosa, submucosa, muscularis and serosa. Taste buds were found in the epithelium of lips, buccopharynx and oesophagus. The stratified epithelium of buccopharynx and oesophagus was located with numerous goblet cells. The U‐shaped stomach has three parts, corresponding to mammalian cardiac, fundus and pyloric portion, lined with a single‐layered columnar epithelium, and tubular gastric glands are present in cardiac and fundic portion, but absent in pyloric portion. No pyloric caeca was detected. The intestine is separated from the stomach by a loop valve. The intestine epithelium is composed of simple columnar cells with a distinct microvillus brush border and many goblet cells. Meanwhile, the intestinal coefficient was 0.898. At the ultrastuctural level, three type cells (mucous, glandular and endocrine cell) were found in the stomach, and glandular cell with a great amount of pepsinogen granules. The enterocytes of the intestinal mucosa display abundant endoplasmic reticulum, mitochondria and well‐developed microvilli. Congxin Xie, College of Fisheries, Huazhong Agricultural University, Wuhan, Hubei 430070, China. E‐mail: xiecongxin@mail.hzau.edu.cn or dreamsail_2005@yahoo.com.cn     

Cyclic histological changes of the oviductal-cloacal junction in the viviparous snake Toluca lineata

The structure and seasonal changes of the oviductal-cloacal junction remain poorly understood in most squamates. This study was undertaken to describe the histology of the oviductal-cloaca junction of a female viviparous snake Toluca lineata, during gestation, previtellogenesis, and vitellogenesis. The oviductal-cloacal junction exhibits a wider lumen and thicker layers of connective tissue, smooth muscle layers, and total wall width compared to the posterior vagina. The lining is characterized by thick, short longitudinal mucosal folds. The luminal epithelia differ morphologically from anterior to posterior portions of the oviductal-cloacal junction. The anterior portion is lined with a simple columnar epithelium composed of nonciliated cells. The middle portion is lined with stratified epithelium that contains an apical columnar cell layer that undergoes morphological changes coincident with the reproductive cycle. The posterior portion is lined with a stratified squamous epithelium. The connective tissue underlying the epithelium contains numerous ovoid cells having abundant acidophilic cytoplasmic granules—eosinophils. Copulation occurs during the previtellogenic stage, as evidenced by the presence of abundant spermatozoa in the lumen of the anterior portion and of a copulatory plug in the middle and posterior portion of the oviductal-cloacal junction. J. Morphol. 237:91–100, 1998. © 1998 Wiley-Liss, Inc.     

Unexpected bird-like features and high intraspecific variation in the braincase of the Triassic relative of dinosaurs

Abstract

Silesaurus opolensis Dzik, 2003 from the Late Triassic (late Carnian) of Poland is a key taxon for understanding the evolution of early dinosaurs. High intraspecific variation observed in the S. opolensis braincase brings caution in taxonomic and diversity studies of early dinosauromorphs. The external and internal osteology of three almost complete braincases of S. opolensis show that this taxon shares several similarities with other early dinosauriforms, which supports a close relationship among these forms. However, the paroccipital processes of S. opolensis are directed ventrally like in birds, reaching the level of the ventral margin of the basioccipital condyle. In dinosauromorphs, these processes usually have an almost horizontal orientation (presumed to be the plesiomorphic condition). Modifications observed in birds and S. opolensis have resulted in the dorsoventral expansion of M. complexus and M. depressor mandibulae, which occupy the dorsolateral part of the posterior side of the skull. In adult birds, these muscles act strongly on the initial upstroke of the head during drinking. Therefore, the inferred condition of these muscles in S. opolensis may imply that Silesauridae evolved toward bird-like feeding behaviour.     

Distribution and structure of glandular tissue in the oropharynx and proximal esophagus of the emu (Dromaius novaehollandiae)

Crole, M.R., Soley, J.T. 2011. Distribution and structure of glandular tissue in the oropharynx and proximal esophagus of the emu (Dromaius novaehollandiae). —Acta Zoologica (Stockholm) 92 : 206–215. The glandular regions of the upper digestive tract in the emu were non‐pigmented (except for the tongue in most specimens) and invested by a non‐keratinised stratified squamous epithelium. The glands found in these regions were exclusively simple in nature and composed of tubular secretory units lined by Periodic Acid Schiff Stain‐positive mucus‐secreting cells. The naming of the various glandular fields was based on previously identified anatomical features and on nomenclature modified from previous studies on birds. The glands were classified into two main types, namely, simple tubular and simple branched tubular mucus‐secreting glands. Simple branched tubular glands were a feature of the regions exposed to the greatest amount of friction during feeding, whereas simple tubular glands were a feature of regions exposed to less friction. The saliva produced by the salivary glands in birds functions to moisturize and lubricate food boli. Mucins in saliva also protect mucosal surfaces from desiccation and mechanical damage, assist in maintaining cellular water balance, provide lubrication and have an antimicrobial action. This study suggests that, in addition to the role of specific gross anatomical features, the type and distribution of glandular tissue in the emu upper digestive tract supports the cranio‐inertial feeding method employed by this species.     

The stomach of Oreochromis niloticus has three regions

The stomach of Oreochromis niloticus was divided into three distinct regions: initial, middle and terminal, corresponding roughly to the cardiac, fundic, and pyloric portions of the mammalian stomach. Grossly, the organ showed initial and terminal portions, the former connected to the distal part of the oesophagus and the latter to the proximal portion of the intestine. There was also a middle region, forming a large blind diverticulum communicating with the first two at their point of junction. The initial or cardiac region was shorter than the middle region but longer than the terminal one, and had a smooth surface devoid of gastric pits. The epithelium in this region was simple columnar devoid of goblet cells, with glandular regions in the lamina propria. The mucosa of the middle or fundic region had gastric pits lined by columnar epithelium, and simple tubular glands filled most of the lamina propria. The terminal or pyloric part of the stomach was very short and its mucosa was slightly folded and devoid of both gastric pits and mucous glandular cells. The lining epithelium of this portion of the stomach was simple columnar and a few goblet cells were seen at its junction with the first part of the intestine. The tunica muscularis of the stomach contained skeletal muscle in the initial and terminal regions, usually intermingled with smooth muscle fibres. Skeletal muscle fibres were also observed in the first portion of the small intestine, near the junction with the stomach.     

A newly discovered sperm transport system in the female of Lygaeidae bugs

In the female reproductive system of the relatively large hemipteran, the western conifer seed bug Leptoglossus occidentalis (Heidemann), a cuticle‐lined tube extends medially along the surface of the vagina from the proximal end of the spermathecal complex anteriorly to the base of the common oviduct. This medial tube houses the proximal end of the spermathecal duct, thereby enabling the transport of material from the spermatheca at the distal end of the spermathecal complex, past the vagina (or bursa copulatrix) and directly to the common oviduct. The proximal portion of the spermathecal complex also contains an insemination duct that is separate from the spermathecal duct. The insemination duct allows the male intromittent organ to extend from the vagina to the spermatheca without navigating through the spermathecal duct. The reproductive systems of two previously studied Hemiptera, the milkweed bug Oncopeltus fasciatus (Dallas) and the box elder bug Leptocoris trivittatus (Say), possess a similar cuticle‐lined medial tube housing the spermathecal duct. This new information provides a clearer understanding of sperm transport in the female reproductive system of Lygaeidae bugs, and helps to clarify the path of the male organ during copulation, as well as the movement of sperm during egg laying.     

Light microscopy and scanning electron microscopy studies on the reduction of the tongue microstructures in the white stork (Ciconia ciconia,Aves)

The structure of the tongue in the white stork (Ciconia ciconia) is observed macroscopically and under light and scanning electron microscopy. Our observations of the tongue reveal a rare terminal reduction of the size of the tongue and microstructures of the lingual mucosa among the investigations of birds published so far. The short, triangular tongue with a pointed tip is approximately 2.5 cm long in the adult and is situated in the caudal part of the oral cavity close to the laryngeal prominence. On the dorsal surface of the tongue, no typical mucosa microstructures like lingual papillae, median groove or lingual prominence are observed. The main structure of the tongue is composed of rostral part of hyoid apparatus, that is, entoglossal cartilage connects with basihyoid. Very thin mucosa is composed of fibrous connective tissue covered with orthokeratinized epithelium. No lingual glands and muscles are observed in the lamina propria of mucosa. Even though the triangular shape of the tongue in the white stork is typical for birds, the inner structure of the reduced organ is composed only of flat cartilagineous entoglossum of hyoid apparatus. During feeding behaviour of the white stork, the food transportation in oral cavity called cranio‐inertial transport is undoubtedly affected by structural reduction of the tongue.     

Données histologiques sur les fosses nasales et leurs annexes chezCrocodylus niloticus Laurenti etCaiman crocodilus (Linnaeus) (Reptilia,Crocodylidae)

Summary The vestibulum is very short and lined by stratified squamous epithelium which contains many alveolar cells. The cavum nasi proprium is exceedingly complex, with three conchal formations and a series of six recesses and sinuses. Olfactory epithelium lines the whole dorsal or dorso-medial half of the cavum, but not the deep sinuses. Non sensory respiratory epithelium lines the ventral or ventro-lateral half of the cavum, all the caviconchal recess, the posterolateral recess, the postturbinal sinus, the postconchal cavity, and the nasopharyngeal duct. In olfactory epithelium the proportion of sensory cells is about 61 % inCrocodylus and 59 % inCaiman; the ratio of sensory cells to supporting cells is about 2.6/1 in the former and 1.8/1 in the latter. Bowman's glands are sero-mucous and normally developed. As in other reptiles, the respiratory epithelium is composed with mucous and ciliated cells; but, in ventrolateral part of the cavum, there are also sero-mucous cells forming small multicellular glands. The hypertrophied lachrymal duct constitutes a very large naso-lachrymal not previously described gland. The lumen is lined by mucous and ciliated cells, the collet of each branched tubular gland by mucous cells and the glandular tubes by sero-mucous cells. Adult crocodilians lack a Jacobson's organ and there are no vomeronasal sensory epithelium in the cavum.In reptiles, aquatic way of life generally involves regression in olfactory epithelium, while Jacobson's organ (or, in Testudines, vomeronasal epithelium) persists and occasionally increases. In crocodilians, things are exactly reverse. After comparing with other Tetrapods, it seems likely that Jacobson's organ has been lost by terrestrial ancestor of crocodilians and birds. Now, only crocodilians posses olfactory epithelium, and naso-lachrymal gland gives them a supplementary protection, necessary in semiaquatic environment.     

Ultrastructural organization of the transition from the distal nephron to the collecting duct in the desert rodent Psammomys obesus

Summary The transition from the nephron to the collecting duct is formed by three tubular segments (convoluted part of the distal tubule, connecting tubule, cortical collecting duct), which in the desert rodent, Psammomys obesus, transform gradually from one segment to the next, due to intermingling of their different cell types.The convoluted part of the distal tubule (DTC) starts abruptly, shortly beyond the macula densa and initially is homogeneously composed of characteristic DTC-cells. Subsequently, the DTC-cells intermingle with intercalated cells. The first appearance of the connecting-tubule cell, which gradually replaces the DTC-cell, is regarded as the beginning of the connecting tubule. The major portion of the connecting tubule is lined by connecting-tubule cells and intercalated cells. The first appearance of the principal cell between them defines the beginning of the cortical collecting duct, which in the medullary ray is lined by principal and intercalated cells only.Each cell type is described in detail and discussed in relation to the assumed function of the tubular segments.Interspecies differences in the cellular composition of the transitional zone from the nephron to the collecting duct are discussed in relation to the different organization of the collecting duct system.     

Histochemical and immunohistochemical adaptations on the gastroesophageal tube of the blue-fronted parrot (Amazona aestiva Linnaeus, 1758)

Amazona aestiva Linnaeus, 1758 is known as the blue-fronted parrot. These animals are considered seed dispersion, contributing to ecological balance. Despite its ecological importance, A. aestiva is listed as an imminent danger of becoming extinct, as it is one of the most illegally traded native species in Brazil, besides losing its habitat due to deforestation and forest fires. The cranial oesophagus mucosa is lined with a non-keratinized stratified squamous epithelium, and the basal layer presents melanocyte accumulation. No mucous glands were observed in the lamina propria/submucosa. The ingluvium two different muscular layer patterns were observed, one internal and circular and one very thick, external and longitudinal. The proventriculum submucosa consists of polygonal proventricular glands and contents into a broad central duct lined with simple columnar epithelium Periodic acid-reactive Schiff positive (PAS+) and Alcian blue positive (AB+). The 5-HT cells are dispersed in the proventricular glands located at base and apex. The ventricle mucous layer contains numerous branched folds lined by a simple columnar epithelium. The epithelium was negative reaction for PAS and positive for AB only on the fold surfaces. The 5-HT cells were scarce and observed only at the base of the ventricular gland, as well as in the adjacent connective tissue.     

Free vascularized appendix transfer for reconstruction of penile urethras with severe fibrosis

Despite the development of newer techniques with a free radial forearm tube flaps for phallus reconstruction, severe urethral strictures are still seen in such cases after irradiation or repeated infection because of the paucity of healthy, well-vascularized tissue. For urethral reconstruction in cases with poorly vascularized tissue as well as for total penile creation, a new technique involving a free vascularized appendix transfer combined with a radial forearm osteocutaneous flap was successfully used in two cases. The appendix provides a normal tube structure composed of a muscular tubular layer lined with mucosal epithelium. It has no hair and has rich vascularization. This results in little stricture at the junction with the original urethra, no occurrence of urethral stones, and possible postoperative enlargement of the diameter with changes in catheters. This method will allow a patient with severe fibrosis around the urethra to undergo one-stage phallus reconstruction with minimal complications.     

Olfactory metamorphosis in the coastal tailed frog Ascaphus truei (Amphibia,Anura, Leiopelmatidae)

The structure of the olfactory organ in larvae and adults of the basal anuran Ascaphus truei was examined using light micrography, electron micrography, and resin casts of the nasal cavity. The larval olfactory organ consists of nonsensory anterior and posterior nasal tubes connected to a large, main olfactory cavity containing olfactory epithelium; the vomeronasal organ is a ventrolateral diverticulum of this cavity. A small patch of olfactory epithelium (the “epithelial band”) also is present in the preoral buccal cavity, anterolateral to the choana. The main olfactory epithelium and epithelial band have both microvillar and ciliated receptor cells, and both microvillar and ciliated supporting cells. The epithelial band also contains secretory ciliated supporting cells. The vomeronasal epithelium contains only microvillar receptor cells. After metamorphosis, the adult olfactory organ is divided into the three typical anuran olfactory chambers: the principal, middle, and inferior cavities. The anterior part of the principal cavity contains a “larval type” epithelium that has both microvillar and ciliated receptor cells and both microvillar and ciliated supporting cells, whereas the posterior part is lined with an “adult‐type” epithelium that has only ciliated receptor cells and microvillar supporting cells. The middle cavity is nonsensory. The vomeronasal epithelium of the inferior cavity resembles that of larvae but is distinguished by a novel type of microvillar cell. The presence of two distinct types of olfactory epithelium in the principal cavity of adult A. truei is unique among previously described anuran olfactory organs. A comparative review suggests that the anterior olfactory epithelium is homologous with the “recessus olfactorius” of other anurans and with the accessory nasal cavity of pipids and functions to detect water‐borne odorants. J. Morphol. 2011. © 2011 Wiley Periodicals, Inc.     

The digestive tract of the amberjack Seriola dumerili, Risso: a light and scanning electron microscope study

The histology of the digestive tract of the amberjack ( Seriola dumerili , Risso) was studied using light and scanning electron microscopy. The anterior oesophagus mucosa displays primary and secondary folds lined with a stratified squamous epithelium with fingerprint-like microridges which is substituted, on the top of the oesogaster folds, by a simple columnar epithelium with short microvilli. Only primary folds are present in the stomach. The anterior portion is rich in simple tubular glands, whereas the oesogaster and the pyloric region are devoid of them. Pyloric caeca and anterior and middle intestine mucosa display the same pattern of folding. The dominant cell type is the enterocyte, which exhibits larger and thinner microvilli in the caeca than in the intestine. The columnar epithelium of the rectum is replaced, in the anal sphincter, by a stratified flattened epithelium. Goblet cells are numerous throughout the whole length of the tract with the exception of the initial part of the oesophagus, the oesogaster, the stomach and the anal sphincter. Mucosubstances have been shown to vary in the different regions of the gut: acid mucines are found in the oesophagus, pyloric stomach, caeca, intestine and rectum, whereas neutral mucosubstances dominate in the anterior portion of the stomach. The muscularis is well developed throughout the length of the tract: two layers of striated muscle at the oesophageal level; two layers of smooth muscle in the stomach wall and three at the intestinal level.     

白胸苦恶鸟消化管的组织学观察

采用石蜡切片技术对白胸苦恶鸟(Amaurornis phoenicurus)的消化道进行了组织学观察.结果表明,食管皱襞发达,黏膜上皮为复层扁平上皮,食管腺发达,颈段多于胸段,黏膜肌层为一层纵肌,厚约0.06 ～0.26 mm.肌层为一层厚约0.19～0.27mm的环肌.腺胃被覆单层柱状上皮,固有层内有单管腺和复管腺两种腺体,单管腺仅深约0.11 ～0.20 mm;复管腺厚约1.19～1.26 mm,占管壁的77.8％ ～80.4％.肌胃的类角质层发达,厚约0.16～0.24 mm.肌胃腺呈管状,与类角质层突起形成皱襞.肌层发达,由内环外纵两层平滑肌构成.肠绒毛无分支和中央乳糜管,十二指肠绒毛长而密集,空肠绒毛呈细长指状,直肠绒毛长且呈叶状.十二指肠与直肠肠绒毛内有大量致密淋巴小结,盲肠绒毛短,肠腺少.     

The gut of the juvenile African lungfish Protopterus annectens: A light and scanning electron microscope study

We describe the microstructure of the alimentary canal of the juvenile lungfish Protopterus annectens. Following the oesophagus, the gut is formed by a long segment that extends down to the pyloric valve. This segment, classically named stomach, is lined by a transitional epithelium but lacks all characteristics of the vertebrate stomach. It has been defined here as the intestinal vestibule. The spiral valve is divided into a first large chamber, which contains mucosal ridges, and a second smooth portion. The entire spiral valve is lined with a pseudostratified columnar epithelium that contains approximately six cell types: enterocytes, goblet cells, ciliated cells, leukocytes, dark pigment cells, and vascular cells. Enterocytes and goblet cells show a high number of cytoplasmic vacuoles. The number and size of the vacuoles, and the number of ciliated cells, decreases from the anterior toward the posterior end, suggesting that most of the digestive processes take place in the anterior part of the spiral valve. The epithelium overlies a lamina propria in the first large chamber and a vascular plexus in the smooth portion. The cloaca has a thick muscular wall covered by a transitional epithelium. An extensive lymphatic system formed by capillaries and lymphatic micropumps is present along the entire wall of the alimentary canal. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.