identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03DC25380361BF00FF7A391DB6C6394A.text	03DC25380361BF00FF7A391DB6C6394A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hyalogyrina depressa Hasegawa 1997	<div><p>Hyalogyrina depressa Hasegawa, 1997</p> <p>(Figures 7, 8, 9, 10, 11, 12 and 13)</p> <p>External morphology</p> <p>The head bears a conical snout with two anterior bulges. The right side of the head and the neck region are densely ciliated, the neck also shows a glandular epithelium. The cephalic tentacles are dorsoventrally flattened. A small (300 μm long), conical pallial tentacle is visible at the right opening of the mantle cavity. This tentacle is densely ciliated (Fig. 8a) and equipped with prominent longitudinal muscle fibers. Eyes and epipodial tentacles are lacking; the mantle margin is smooth and ciliated.</p> <p>Foot and shell muscles</p> <p>In the contracted animals the foot is transversely folded like a pocket knife (Fig. 7a,b). The anterior foot is deeply bilobed, the posterior end is rounded. The epithelium of the foot’ s sole consists of highly cylindrical, densely ciliated cells interspersed with poorly stained mucous cells. The back of the foot lacks cilia, however. The anterior foot gland forms an irregular mass in the dorsal half of the propodium. Two lobes of this mass fill the anterior foot lappets, another two lobes extend laterally, and one lobe runs backwards ventrally. The central duct (diameter 25 μm) opens at the anterior median edge of the foot sole. The cells are large (20 μm× 36 μm) and have irregularly shaped, basal nuclei (Fig. 7e).</p> <p>The much smaller, posterior foot gland contains weakly stained mucous cells and opens via a short duct in the center of the foot sole. The posterior gland forms a flat mass and is somewhat embedded in the connective tissue of the foot.</p> <p>In the central portion of the foot there is an accumulation of very large (diameter 60–95 μm), clear, oval to deformed calcium cells with large (diameter 15 μm), round nuclei. Whereas the cytoplasm of these cells is restricted to narrow strings, the main volume of the cells is mainly occupied by large vacuoles containing fine concrements (Fig. 7d), which show negative contrast in phase contrast microscopy. Otherwise the foot mass contains muscle and collagen fibers as usual, haemolymph spaces and parenchymous cells.</p> <p>There is a single columellar muscle, which adheres ventrally at the shell (Fig. 9a,b).</p> <p>Mantle cavity (Fig. 7a–c)</p> <p>The mantle cavity occupies about one third of the volume of the last shell whorl. The anterior part of the mantle cavity is occupied by the gill. On the left side, about 200 μm behind the edge, the osphradium forms a thickened ridge. More posterior and to the left the single kidney is situated in front of the monotocardian heart. The posterior, central and right mantle roof is occupied by several rectal loops, followed towards posterior right by the receptaculum seminis and the gonoduct with the gonopore. The posterior wall of the mantle cavity is formed by the bursa copulatrix, the anterior portion of the digestive gland, and to the left by the stomach.</p> <p>The whole mantle roof is underlain by wide blood sinuses, in particular near the anterior mantle edge. On the inner side of the mantle roof there are various glands with propodium, pt = pallial tentacle, r = radula, re = rectum, rs = receptaculum seminis, st = stomach, v = heart ventricle. Supplementary plate 2 offers an interactive 3D model of Hyalogyrina depressa that can be accessed by clicking into Fig. 7 (Adobe Reader version 7 or higher required). Rotate model: drag with left mouse button pressed; shift model: same action+ctrl; zoom: use mouse wheel (or change default action for left mouse button). Select or deselect (or change transparency of) components in the model tree, switch between prefab views, or change surface visualization (e.g. lighting, render mode, crop, etc.)</p> <p>various types of mucous cells: the right pallial gland consists of cells with an empty or bubble-like structure, and occasional dark, apical granules dominate (Fig. 7a: pg; 8a: mg1). On the left side another pallial gland has mucous cells showing metachromatic content (Fig. 7b: pg’; 8b: mg2). In the central pallial roof and occasionally also laterally there are mucous cells which appear empty in the sections (Fig. 8c: mg3).</p> <p>The bipectinate gill shows about 20 alternating lamellae, which have ciliary tracts and regions with very flat (respiratory) epithelia (Fig. 8d). Beneath the efferent blood sinus, the efferent gill axis has a thick muscle which runs into the left mantle, but on the right (afferent) side no distinct retractor is present.</p> <p>Excretory organ, heart and circulatory system (Fig. 9)</p> <p>The rectal sinus runs along the rectum forwards to the right and enters the afferent axis of the gill. After passing the leaflets the efferent gill sinus collects the haemolymph and also receives a mantle sinus, thus becoming the afferent sinus of the kidney. After passing the kidney walls the efferent kidney sinus opens into the auricle, followed by the ventricle (Fig. 9b,c). The head aorta runs forwards along the median mantle floor.</p> <p>The monotocardian heart is situated in the left posterior mantle cavity. The kidney forms a boomerang-shaped organ in the posterior mantle roof and shows a distinct nephropore (Fig. 8e: arrowhead). It is connected with the pericardium by a ciliated renopericardial duct.</p> <p>Genital system (Figs. 10, 11)</p> <p>The true hermaphroditic gland occupies the right half of the visceral body (Fig. 10a). The right portion has a homogenous lumen, from which several lobes expand to the left (Fig. 10b,c). The inner portion of these lobes produces the eggs, which become more mature from left to right and from posterior to anterior (Fig. 11f). Spermiogenesis takes place towards the central parts in the portions of the hermaphroditic gland which lie adjacent to the external epithelium of the visceral sac (Fig. 11f). Mature sperm (only one type is present) have filiform heads and long flagella, and are orientated centrifugally (Fig. 11g).</p> <p>The hermaphroditic duct emerges on the median side of the gonad and runs to the right, where it widens to form a vesicula seminalis containing autosperm with their heads orientated forwards (Fig. 11d). From the proximal part of the gonoduct a prominent blind sac (ca. 350×100× 40 μm) emerges ventrally (Figs. 10a,b; 21c: vs). Its dorsal epithelium consists of thick (up to 22 μm) glandular cells with heavily stained cytoplasms, whereas the ventral epithelium is densely ciliated. The gonoduct as a whole forms an Sshaped loop towards ventral, then anteriorly left, and finally runs in a curve to ventral right towards the genital atrium. It consists of three histologically distinct regions (Fig. 10d: gd1/2/3), as follows. (1) The glandular proximal part shows many dark granules of various sizes, further structures are poorly preserved (Fig. 10d: gd1). From here a prominent sinuses and gill muscle. (d) Circulatory system and kidney (pericardium transparent), left side view. Labels: a = auricle, ags = afferent gill sinus, ao = aorta, cm = columellar muscle, egs = efferent gill sinus, gi = gill, gm = gill muscle (retractor), hm = head muscle (retractor), k = kidney, pc = pericardium, rs = rectal sinus, v = heart ventricle</p> <p>(350 μm×100 μm×40 μm) blind sac emerges (Figs. 10d; 11a: bs1). Its epithelium shows mucous cells with dark cytoplasm and pale nuclei, as well as clear granula or vesicles; the ventral epithelium is densely ciliated. (2) The proximal part is continued by the central part of the gonoduct consisting of long, ciliary cells filled with large, dark vesicles (Figs. 10d; 11a,c: gd2). At the distal end of the central part another glandular blind sac (440 μm× 95 μm×30 μm) is formed, somewhat surrounding the gonoduct (Figs. 10d; 11a: bs2). This gland consists of many lobes with lightish, homogeneous lumina; the lumen opens via a ciliary duct into the gonoduct. (3) The distalmost part of the gonoduct is again formed by long cells with granules, but the latter show metachromic staining (Fig. 11a: gd3). The duct finally enters the genital atrium, from which the following two further structures arise.</p> <p>(A) The bursa copulatrix is embedded in the proximal and median gonoduct (Fig. 10b,c). It is a bean-shaped structure (50 μm×160 μm×50 μm) with a ciliated epithelium. Its large lumen shows organic particles that could not be identified unequivocally, but probably are degraded allosperm (Fig. 11e). A narrow, ciliated duct connects the bursa with the genital atrium.</p> <p>(B) The receptaculum seminis (160 μm×80 μm×70 μm) is situated to the right of the bursa and also lies adjacent to the gonoduct (Fig. 10a–c: rs). It is filled with densely packed allosperm (Fig. 11c). The ciliated duct runs along the gonoduct and again opens into the genital atrium.</p> <p>The genital atrium itself opens via a short duct into the posterior right mantle cavity. A copulatory organ is not present.</p> <p>Alimentary tract (Fig. 12)</p> <p>The small and narrow mouth opening is flanked by lateral oral lappets. Behind the mouth opening two triangular jaw gonoduct (arrow marks position of blind sac 2 along ridge). Labels: b = bursa, bs1/2 = blind sac 1/2, g = gonad, ga = genital atrium, gd = gonoduct, gd1/2/3 = proximal/median/distal portion of gonoduct, rs = receptaculum seminalis, vs = ventral sac plates are situated laterodorsally, consisting of tooth-like elements flanking the oral cavity, which is of inverted Yshape (Fig. 12d). The radula is supported by a purely muscular odontophore; true cartilages are lacking. The radula sheath makes a sharp loop downwards, its blind end consists of numerous odontoblasts (Fig. 12e).</p> <p>The pharynx continues dorsally into the anterior oesophagus, which shows several densely ciliated, longitudinal folds (Fig. 12f). It is flanked by the paired, large (200 μm × 80 μm × 40 μm) salivary glands, which possess many small lobes. The salivary ducts show many loops (due to the contracted condition), their glandular epithelia are heavily stained, whereas the posterior parts show large, pale cells with very large nuclei (Fig. 12g). Effects of torsion are minimal; the posterior part is a simple, narrow (diameter 55 μm) tube that runs backwards and enters the stomach between the openings of the two digestive glands (Fig. 12i).</p> <p>The posterior stomach (diameter 500 μm) is equipped with a large but thin gastric shield with tooth, followed more anteriorly by ciliated areas with high prismatic cells (Fig. 12h). Again a small caecum is present at the ventral side of the stomach (Fig. 12h: ca).</p> <p>The anterior digestive gland occupies the visceral body to the right of the stomach, whereas the posterior gland fills the visceral sac. Both consist of a spreading duct system reaching into irregularly shaped diverticles (Fig. 12i). The epithelium of the digestive glands shows lightish cells with round nuclei and a compact cytoplasm in the apical portion (Fig. 12i).</p> <p>The short and ciliated intestinal tube (diameter 80 μm) emerges from the stomach on the right side, runs forwards and then upwards entering the mantle roof. The rectal portion shows several narrow loops in the mantle roof (Figs. 7c; 12c). The anal opening is situated in the posterior right mantle cavity.</p> <p>Stomach and intestine contain remnants of detritus, including diatom tests.</p> <p>Nervous system (Fig. 13a,b)</p> <p>The cerebral ganglia are situated at the basis of the cephalic tentacles. They are compressed dorsoventrally and connected by a short commissure. The tentacle nerve bifurcates shortly after emergence (Fig. 13a,b: nerve 2). The pleural ganglia are fused with the cerebral ganglia (epiathroid condition); these cerebropleural ganglia are connected to the pedal ganglia by two connectives (n3, n4). The pedal ganglia are connected by a short commissure; one pair of pedal nerves runs forwards (n6), another pair (n7) backwards into the foot. The buccal ganglia with their thin and quite long commissure (n9) are situated at the line of the opening of the salivary glands, and are connected to the cerebral ganglia (n8).</p> <p>stomach (arrowheads: gastric shield). (i) Longitudinal section of posterior oesophagus at line of digestive gland openings, anterior is at right (arrowhead: dense cytoplasm of apical epithelium of digestive gland). Labels: adg = anterior digestive gland, asg = anterior part of salivary gland, ca = stomach caecum, ci = ciliary border, csa = ciliary sorting area of stomach, gs = gastric shield, hao = head aorta, hec = head epidermis, j = jaw, mc = mantle cavity, mt = mouth tube, od = odontoblasts, oe = oesophagus, pdg = posterior digestive gland, ph = pharynx, psg = posterior part of salivary gland, r = radula, re = rectum, sg/sg’ = right/left salivary gland, st = stomach</p> <p>From the right pleural ganglion a connective (n10) crosses above the oesophagus to the left side and reaches the supraoesophageal ganglion, which is situated above the left pleural ganglion. From the latter a short nerve (n11) connects to the osphradial ganglion in the left mantle roof. A left zygoneurosis (Fig. 13b: arrowhead) connects the left pleural with the supraoesophageal ganglion. Posteriorly a mantle nerve (n12) and the posterior portion of the visceral loop (n14’) emerge from the left pleural ganglion.</p> <p>From the left pleural ganglion another connective (n13) crosses below the oesophagus to the right and reaches the suboesophageal ganglion at the line of the right pedal ganglion. Again a mantle nerve (n12) and the posterior part</p> <p>P/P’ = right/left pedal ganglion, Pl/Pl’ = right/left pleural ganglion, Sb = suboesophageal ganglion, Sp = supraoesophageal ganglion, st/st’ = right/left statocyst; 1 = cerebral commissure, 2/2’ = right/ left tentacle nerve, 3/3’ = right/left cerebropedal connective, 4/4’ = right/left pleuropedal connective, 5 = pedal commissure, 6/6’ = right/left anterior pedal nerve, 7/7’ = right/left ventral pedal nerve, 8/8’ = right/left cerebrobuccal connective, 9 = buccal commissure, 10 = supraoesophageal connective, 11 = supraoesophageal-osphradial connective, 12 = visceral loop, 13 = suboesophageal connective, 14/14’ = right/left mantle nerve of the visceral loop (n14) emerge posteriorly. A visceral ganglion could not be detected.</p> <p>Sensory organs (Fig. 13c,d)</p> <p>Eyes and epipodial tentacles are entirely lacking; the cephalic tentacles are smooth (Fig. 13d). Each of the two bean-shaped statocysts (30 μm×20 μm ×10 μm), which are situated at the postero-dorsal sides of the pedal ganglia, contains a small (diameter 12 μm), round statolith (Fig. 13c). The osphradial epithelium is higher than that of the surrounding mantle roof; histological details could not be detected (Fig. 13d).</p> <p>Data from other sources</p> <p>Hasegawa (1997) gave SEM images of the depressed, smooth shell (op.cit.: fig. 29A–C), the hyperstrophic granular to smooth protoconch (fig. 29D, E) forming an anastrophic apex, of the round, multispiral operculum (fig. 29F), the rhipidoglossate radula (fig. 29G, H) with the formula n-1-1-1-n, and of a jaw plate (fig. 29I) consisting of tooth-like elements. He also provided SEM photos of the head-foot (fig. 30A, B) and the gill leaflets (fig. 30C).</p></div> 	https://treatment.plazi.org/id/03DC25380361BF00FF7A391DB6C6394A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Hess, Gerhard Haszprunar Erika Speimann Andreas Hawe Martin	Hess, Gerhard Haszprunar Erika Speimann Andreas Hawe Martin (2011): Interactive 3 D anatomy and affinities of the Hyalogyrinidae, basal Heterobranchia (Gastropoda) with a rhipidoglossate radula. Organisms Diversity & Evolution (New York, N. Y.) 11 (3): 201-236, DOI: 10.1007/s13127-011-0048-0, URL: http://dx.doi.org/10.1007/s13127-011-0048-0
03DC25380379BF0BFF7A3D45B44A3F95.text	03DC25380379BF0BFF7A3D45B44A3F95.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hyalogyrina grasslei Waren & Bouchet 1993	<div><p>Hyalogyrina grasslei Warén &amp; Bouchet, 1993 (Figures 14, 15, 16 and 17)</p> <p>External morphology</p> <p>The epithelium of the head region is covered by a microvillous border with interspersed ciliary tufts. The snout is tapered but slightly bilobate, the mouth opening is quite small. Between the cephalic tentacles there are two short, ciliated knobs (Fig. 14a,b). Another small process—probably a copulatory organ (see below)—is situated at the basis of the right cephalic tentacle close to the openings of receptaculum and gonoduct. A seminal groove could not be detected. In the anterior part of the right mantle cavity there is another small (about 0.2 mm), ciliated tentacle.</p> <p>Pedal structures</p> <p>The foot is folded due to preservation. The single pedal gland opens via a wide pore at the anterior border of the foot, but its posterior portion is difficult to distinguish from the surrounding connective tissue. In the center of the foot there is a densely packed group of large (diameter 80 μm) and transparent calcium cells, each provided with a large (diameter 13 μm) nucleus (Fig. 15g). The remaining foot mass shows large haemocoelic spaces with interspersed collagen and muscle fibers; parenchymous cells are rare.</p> <p>Shell muscle, haemocoel, and mantle cavity</p> <p>The conditions of the shell (columellar) muscle and of the haemocoel resemble those in Xenoskenea pellucida and Hyalogyrina depressa.</p> <p>The entire mantle roof includes large blood sinuses, which are separated from each other by fine fibers of muscle or collagen. The mantle cavity itself is quite large and shows (from left to right) the following structures (Figs. 14a,b; 15a). On the very left, the osphradium forms a round, elevated structure underlain by the osphradial ganglion. Whereas distinct hypobranchial and pallial glands are lacking, the anterior part of the mantle roof is glandular. The single, prominent, bipectinate gill has 19 dorsal and 16 ventral alternating leaflets, which lack a skeleton but are densely ciliated except for flat, interspersed non-ciliated cells, which dominate the distalmost parts of the leaflets (Fig. 15b,c). The gill reaches from the left posterior portion of the mantle roof to the posterior mediodorsal region. Afferent and efferent sinuses are well visible in the sections. At the very left pallial roof the single large, elongated kidney is situated behind the gill, followed by the pericardium with the heart. The right pallial roof is occupied by several loops of the rectum. To the very right there are the openings of the receptaculum and the gonoduct.</p> <p>Excretory organ, heart and circulatory system</p> <p>The efferent gill sinus runs backwards and becomes the afferent sinus of the kidney, the ventral (pallial) wall of the latter shows many blood sinuses. The narrow nephropore lies adjacent to the posterior edge of the gill. Whereas the main portion of the kidney shows the typical epithelium with small, round concrements, which are also present in the lumen, the right posterior part of the kidney is glandular (Fig. 15d,e). Posteriorly a ciliated renopericardial duct connects the kidney with the pericardium. Leaving the kidney posteriorly the sinus enters the auricle and ventricle of the monotocardian heart, which is situated behind the kidney. Again the head aorta runs forwards along the median mantle floor.</p> <p>Genital system (Fig. 16)</p> <p>Hyalogyrina grasslei is a simultaneous hermaphrodite. The tubular hermaphroditic gonad is composed by two main branches, an anterior and a posterior one, which split into a fan-like structure (Fig. 16a). Spermiogenesis takes place in the outer parts, oogenesis in the inner parts of the tubes (Fig. 16c); both types of germ cells show all stages of development. The fused portion of the two main branches becomes the hermaphroditic duct, which is not glandular but densely filled with sperm for a length of about 250 μm, and is meandering later on. Connected with the latter portion of the hermaphroditic gland there is a prominent blind sac (Fig. 16a,c: bs1), which runs backwards for about 200 μm.</p> <p>The hermaphroditic duct opens into the glandular part of the gonoduct; it can be divided in several portions on account of distinct mucous cell types. Along the most proximal portion the glandular cells are cuboidal and contain granules which stain heavily. At the line of the posterior end of the mantle cavity the (still closed) lumen of right/left anterior pedal nerve, 7/7’ = right/left ventral pedal nerve, 9 = buccal commissure, 10 = supraoesophageal connective, 11 = supraoesophageal-osphradial connective, 14’ = left mantle nerve. Supplementary plate 3 offers an interactive 3D model of Hyalogyrina grasslei that can be accessed by clicking into Fig. 14 (Adobe Reader version 7 or higher required). Rotate model: drag with left mouse button pressed; shift model: same action+ctrl; zoom: use mouse wheel (or change default action for left mouse button). Select or deselect (or change transparency of) components in the model tree, switch between prefab views, or change surface visualization (e.g. lighting, render mode, crop, etc.)</p> <p>the gonoduct splits into a ciliated seminal portion (vas efferens) and a thick, glandular oviduct. Both ducts remain associated, however (Fig. 16f). There is a second, narrow, ciliated blind sac (Fig. 16g: bs2) again leading backwards, but this also has a very narrow connection to the distal part of the vas deferens. The thick oviduct shows several prominent mucous cells. The distalmost part of the gonoduct still shows the oviduct and the vas deferens in close association, but there are separate openings for sperm and eggs.</p> <p>Anterior to the female opening there is the opening of a prominent receptaculum seminis, which runs backwards adjacent to the oviduct nearly to the line of the second blind sac. Its epithelium is densely ciliated (Fig. 16b,d) and underlain by a muscular layer, the lumen is filled with sperm (Fig. 16d,h).</p> <p>As outlined above, there is no seminal groove and only a small copulatory organ at the basis of the right cephalic tentacle.</p> <p>Alimentary tract (Fig. 17)</p> <p>The whole buccal apparatus is structured nearly identically to that in Xenoskenea pellucida and Hyalogyrina depressa (see above). Here we recognized a specific paired retractor muscle from the anterior lateral buccal wall to the posterior wall of the head, with an enclosed nerve that originates from the buccal ganglion.</p> <p>The conditions of the oesophagus and stomach also strongly resemble those in Xenoskenea. We recognized five teeth of the gastric shield (Fig. 17f), and the two digestive glands have a common opening into the stomach. The very short intestine starts at the anterior wall of the stomach. The rectum passes beneath the heart, runs forwards to the left, then makes three loops in the pallial roof. The anal opening ciliary sorting area of stomach, dfc = dorsal food channel, dg = digestive gland, ex = external milieu, gd = gonoduct, gs = gastric shield, gt = spines of gastric tooth, hao = head aorta, i = intestine, j = jaw, mc = mantle cavity, oe = oesophagus, ph = pharynx, poe = posterior oesophagus, r = radula, re = rectum, rm = radular musculature, sg/sg’ = right/left salivary gland, st = stomach, x = degraded food</p> <p>(Fig. 17g) is situated in the anterior right mantle cavity close to the male opening and shows a free end of the rectal tube.</p> <p>Nervous system and sensory organs</p> <p>The nervous system is nearly identical to those in Xenoskenea pellucida and Hyalogyrina depressa (see, e.g., Fig. 14c,d).</p> <p>The cephalic tentacles with their bifid tentacular nerves are densely ciliated, except in the outer lateral portions. Eyes are entirely lacking; the small statocysts again contain single statoliths.</p> <p>Available data from other sources</p> <p>Warén and Bouchet (1993) described the shell (op.cit.: figs. 39D–G) as small (max. 2.6 mm), very thin, and colourless, and the smooth protoconch as indistinct with 0.6 to 0.7 whorls (fig. 42E). The operculum is multispiral. The short radula is rhipidoglossate with the formula n 3 1 3 n (op.cit.: SEM figs. 40B, C, E); about 20 marginal teeth are present. Concerning external morphology (SEM figs. 41B– D), the animal has a long, tapered snout; eyes are lacking. Three short, flat tentacle-like structures with ciliated edges are situated between the smooth cephalic tentacles with lateral ciliation. A copulatory organ could not be observed even by SEM; the pallial margin is very thick. The foot is bifurcated anteriorly, whereas the posterior end is truncated; epipodial tentacles are lacking. A prominent, bipectinate gill with about 15 leaflets occupies most of the mantle cavity.</p> <p>The discrepancy concerning the copulatory organ remains obscure: even the SEM photos provided by Warén and Bouchet (1993) do not show any trace of it, whereas our sections revealed it beyond doubt. Since species identity appears certain (we examined paratype material), the only explanation might be that the specimens examined in the two studies differed in sexual maturity.</p></div> 	https://treatment.plazi.org/id/03DC25380379BF0BFF7A3D45B44A3F95	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Hess, Gerhard Haszprunar Erika Speimann Andreas Hawe Martin	Hess, Gerhard Haszprunar Erika Speimann Andreas Hawe Martin (2011): Interactive 3 D anatomy and affinities of the Hyalogyrinidae, basal Heterobranchia (Gastropoda) with a rhipidoglossate radula. Organisms Diversity & Evolution (New York, N. Y.) 11 (3): 201-236, DOI: 10.1007/s13127-011-0048-0, URL: http://dx.doi.org/10.1007/s13127-011-0048-0
03DC25380372BF08FF7A3A60B5C13F95.text	03DC25380372BF08FF7A3A60B5C13F95.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hyalogyrina glabra Marshall 1988	<div><p>Hyalogyrina glabra Marshall, 1988</p> <p>(Figures 18, 19 and 20)</p> <p>External morphology and pedal structures</p> <p>Again the snout is tapered and the mouth opening is quite small. The smooth cephalic tentacles are quite long, but a copulatory organ is not present. A long, densely ciliated pallial tentacle emerges from the right anterior mantle edge.</p> <p>There is a single pedal gland, which opens via a wide pore at the anterior border of the foot. Again a densely packed aggregation of large (diameter 80 μm) and transparent calcium cells is found in the center of the foot. The foot sole is densely ciliated.</p> <p>Shell muscle, haemocoel, and mantle cavity</p> <p>Concerning the columellar muscle and the histology of the haemocoel the conditions in H. glabra appear similar to the preceding species, but histological analysis has been limited due to poor preservation.</p> <p>The mantle roof (Fig. 18d) is equipped with a dense net of blood sinuses throughout its area. At the very left of the mantle cavity a small osphradium with an underlying ganglion is present (Fig. 20). It is followed to the right by a (left) pallial gland (not shown in Fig. 18) with many, small mucous cells, and more posteriorly by the kidney and the heart. The central mantle cavity is occupied by the gill, and posterior of the latter by the rectal loops with the anal opening on the anterior right side. The genital apparatus with gonoduct and receptaculum fills the right corner of the mantle cavity, the most anterior portion is covered by a pallial gland on the right (Fig. 18: pg).</p> <p>The gill is again a bipectinate structure without skeleton or bursicles, but is fixed to the pallial roof only at its left side, otherwise it reaches freely into the cavity.</p> <p>Excretory organ, heart and circulatory system</p> <p>Conditions of excretory organ (single, left, pallial), heart (monotocardian), and circulatory system (with median head aorta) are nearly identical to those described for the other species investigated. We could not detect a nephridial gland, but this might be due to insufficient preservation.</p> <p>Genital system (Fig. 19)</p> <p>The hermaphroditic gonad fills the most posterior part of the body, is a compact organ and shows spermiogenesis in its outer, oogenesis in its inner portions (Fig. 19d). The gonad reaches forwards up to the line of the posterior end of the mantle cavity and is continued by a hermaphroditic duct and the glandular part of the gonoduct, where the lumen is branched into a vas deferens and an oviduct, although both ducts continue forwards in parallel and intimately associated (Fig. 19c). Whereas the vas deferens is a thin, ciliated tube, the oviduct is a thick, glandular structure with a lumen that appears slit-like in cross-sections. Again there are various mucous cell types resulting three distinct portions of the oviduct, and at midlength of the latter there is a small pouch (bs1). Vas deferens and oviduct each have their own openings into the right mantle cavity.</p> <p>Close to the two genital openings there are two further openings. Laterally at left there is the opening of the separated, tube-like receptaculum seminis, which is surrounded by a muscular layer and has a densely ciliated epithelium (Fig. 19b). It includes up to two spermatophores, with densely packed sperm in one specimen (Fig. 19e). Distally the second opening forms a small cavity (‘genital atrium’) around male and female openings. From there a long, thin flagellum reaches backwards.</p> <p>right/left radular musculature, rs = radular sheath, st = stomach. Supplementary plate 4 offers an interactive 3D model of Hyalogyrina glabra that can be accessed by clicking into Fig. 18 (Adobe Reader version 7 or higher required). Rotate model: drag with left mouse button pressed; shift model: same action+ctrl; zoom: use mouse wheel (or change default action for left mouse button). Select or deselect (or change transparency of) components in the model tree, switch between prefab views, or change surface visualization (e.g. lighting, render mode, crop, etc.)</p> <p>Unfortunately, the flagellum in the specimen investigated ends at a break, thus could not be analyzed fully.</p> <p>Alimentary tract (Fig. 18d–f)</p> <p>The whole buccal apparatus is structured nearly identically to those of the preceding species. Again we recognized a specific paired retractor muscle from the anterior lateral buccal wall to the posterior wall of the ovotestis; (e) proximal receptaculum seminis with spermatophore. Labels: ci = cilia, e = eggs, ex = external milieu, fl = flagellum, g = gonad, ovotestis, ga = genital atrium, gd = gonoduct, hd = hermaphroditic duct, mc = mantle cavity, rs = receptaculum seminis, sp = autosperm, vd = vas deferens head, with an enclosed nerve which contacts the buccal ganglion.</p> <p>Also the conditions of the oesophagus and stomach strongly resemble those in the other species. We recognized a single tooth of the gastric shield, and the two digestive glands have a common opening into the stomach. The very short intestine starts at the anterior wall of the stomach. The rectum passes beneath the heart, runs forwards to the left, then makes an anterior and a posterior loop in the pallial roof, and finally reaches the anal opening in the anterior right mantle cavity.</p> <p>Nervous system and sensory organs (Fig. 20)</p> <p>The epiathroid and streptoneurous nervous system is nearly identical to those in the species described above. Due to insufficient preservation we could not clarify the condition of the posterior visceral loop.</p> <p>Sensory structures resemble Hyalogyrina grasslei in that the cephalic tentacles are supplied by a bifid nerve and eyes are entirely lacking. Again the small statocysts contain single statoliths.</p> <p>Available data from other sources</p> <p>Marshall (1988) described the shell as small (max. 1.7 mm), turbinate, transparent, and sculptured (op.cit.: figs. 7F–H, J). The hyperstrophic protoconch has 1.25 whorls (fig. 7I). The radula is rhipidoglossate with the formula n 1 1 1 n, (figs. 16A–D), “the single pair of positionally laterals would seem to be marginals, the original laterals … have presumably been lost” (op.cit.: p. 1000). Marshall also provided a drawing of the head (op. cit.: fig. 9G), which carries a long snout, no eyes, but a filiform ‘copulatory organ’ (according to our results the pallial tentacle) behind the basis of the right cephalic cerebropedal connective, 4’ = left pleuropedal connective, 5 = pedal commissure, 6/6’ = right/left anterior pedal nerve, 7/7’ = right/left ventral pedal nerve, 9 = buccal commissure (not fully reconstructed), 10 = supraoesophageal connective, 11 = supraoesophageal-osphradial connective, 13 = suboesophageal connective tentacle. The foot shows two anterior lappets; epipodial tentacles again are lacking.</p> <p>Hedegaard (1990: pl. 52, fig. 3) figured a shell break demonstrating the shell’ s purely aragonite structure. Finally Warén and Bouchet (1993: fig. 42D) have provided SEM images of the protoconch that clearly show the hyperstrophic condition.</p></div> 	https://treatment.plazi.org/id/03DC25380372BF08FF7A3A60B5C13F95	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Hess, Gerhard Haszprunar Erika Speimann Andreas Hawe Martin	Hess, Gerhard Haszprunar Erika Speimann Andreas Hawe Martin (2011): Interactive 3 D anatomy and affinities of the Hyalogyrinidae, basal Heterobranchia (Gastropoda) with a rhipidoglossate radula. Organisms Diversity & Evolution (New York, N. Y.) 11 (3): 201-236, DOI: 10.1007/s13127-011-0048-0, URL: http://dx.doi.org/10.1007/s13127-011-0048-0
03DC25380371BF0FFC933A60B40F3F10.text	03DC25380371BF0FFC933A60B40F3F10.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hyalogyra expansa Marshall 1988	<div><p>Hyalogyra expansa Marshall, 1988</p> <p>Anatomy and histology</p> <p>Because of the very poor preservation of the specimens available, only a superficial description can be provided which focuses on differences to the species described above.</p> <p>There is only a single pedal gland, which is quite large and lies in the anterior foot.</p> <p>The single (left) ctenidium is quite large, bipectinate, and lacks skeletal support. Its anterior half reaches free into the mantle cavity. Again there is a monotocardian heart and a single (left) kidney lying in the pallial roof. The single (left) hypobranchial gland shows three distinct regions differing in histology. In addition, a huge medio-dorsally placed pallial gland (brood pouch?) with various cell types is present. More posteriorly a separated, glandular (much narrower), tubular vas deferens (prostate gland) opens into the mantle cavity.</p> <p>This species represents another simultaneous hermaphrodite, and again the genital apparatus as a whole is a very complex system. The hermaphroditic gland shows distinct regions of sperm (outwards) and egg (inwards) production. The eggs are very large (more than 200 μm in diameter) and yolk-rich. There is a receptaculum below and posterior to the heart on the left side; a ciliated duct connects the receptaculum to the mantle cavity. A copulatory organ is situated behind the right cephalic tentacle and is supplied by a nerve from the cerebropleural connnective. The posterior basis of the copulatory organ is provided with another short appendix.</p> <p>The entire buccal apparatus resembles those in the Hyalogyrina species, but each buccal retractor clearly shows a nerve inside. The anterior oesophagus has two main dorsal folds and shows torsion, and there are true, separate oesophageal glands (not pouches). The posterior oesophagus is not folded. The stomach is again equipped with gastric shield and ciliary fields. Intestinal loops are found only below the posterior mantle cavity; there is no anterior loop. The very thick pallial gonoduct shifts the pallial rectum far to the left side of the mantle cavity.</p> <p>The nervous system is again epiathroid and streptoneurous and strongly resembles those in the Hyalogyrina species. A so-called juxtaganglionar organ (e.g. Clare 1987; Herbert 1982; Martoja 1965a, b; Switzer-Dunlap 1987) lies at the dorsal side of the short cerebral commissure. The statocysts are attached at the posterior-ventral side of the pedal ganglia and again contain a single statolith each.</p> </div>	https://treatment.plazi.org/id/03DC25380371BF0FFC933A60B40F3F10	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Hess, Gerhard Haszprunar Erika Speimann Andreas Hawe Martin	Hess, Gerhard Haszprunar Erika Speimann Andreas Hawe Martin (2011): Interactive 3 D anatomy and affinities of the Hyalogyrinidae, basal Heterobranchia (Gastropoda) with a rhipidoglossate radula. Organisms Diversity & Evolution (New York, N. Y.) 11 (3): 201-236, DOI: 10.1007/s13127-011-0048-0, URL: http://dx.doi.org/10.1007/s13127-011-0048-0
03DC2538034ABF33FF543FF7B2833ECB.text	03DC2538034ABF33FF543FF7B2833ECB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hyalogyrinidae Waren & Bouchet 1993	<div><p>Affinities of Hyalogyrinidae</p> <p>Generic division of Hyalogyrinidae</p> <p>Marshall (1988) noted the close affinity of Hyalogyrina with Hyalogyra and distinguished the two genera mainly on radula characters: Hyalogyra expansa (formula n 6 1 6 n) has considerably more lateral and marginal teeth than Hyalogyrina (formula n 1 1 1 n). Xenoskenea pellucida has a third kind of radula (n-3-1-3-n according to Warén et al. 1993) and is also clearly different from both other genera in showing eyes, distal appendages of the snout, the single tentacle at the posterior end of the foot, the large mantle pad, and a small but true copulatory organ. Accordingly, Hyalogyra and Hyaloyrina appear more closely related to each other than to Xenoskenea.</p> <p>Position of Hyalogyrinidae among the Ectobranchia and Heterobranchia</p> <p>We regard Linnean ranks solely as expressions of a relative hierarchy, thus we do not argue pro or contra superfamilial (Valvatoidea) versus higher rank (Ectobranchia). Ectobranchia is preferred, because (1) Valvatoidea (often cited as Valvatida or Valvatacea) is also a major group of sea-stars, and (2) Ectobranchia is independent of ranking and refers to a clear synapomorphy of the group, the ectobranch gill condition (the inclusion of the gill-less Orbitestellidae cannot be substantiated by any morphological character and is contradicted by the molecular analyses; Dinapoli and Klussmann-Kolb 2010). Among the Heterobranchia (see below), Ectobranchia (Valvatoidea) are clearly characterized by plesiomorphic and apomorphic characters:</p> <p>Plesiomorphic traits mainly concern the alimentary tract, where paired jaws composed of teeth and a dorsal food channel in the buccal roof and the anterior oesophagus are still present and the stomach is still equipped with a cuticularized gastric shield and tooth as well as with ciliary sorting areas. Concerning the radula there is a clear trend to reduce the number of teeth per row: Hyalogyrinidae still show the rhipidoglossate type; Cornirostridae and Valvatidae have a taenioglossate radula with 7 or 9 teeth per row (see Warén et al. 1993 for discussion); Xylodisculidae (formula 2 1 0 1 2) lack a rhachidian tooth; and Orbitestellidae have only three teeth per row (1-1- 1). This transformation series of the radula type character places Hyalogyrinidae at the basis of Ectobranchia by outgroup comparison, since a re-establishment of the rhipdioglossate type looks very improbable.</p> <p>Van den Biggelaar and Haszprunar (1996) showed that Valvata piscinalis differentiates its mesentoblast 4d cell at the 40-cell stage, whereas opisthobranch and pulmonate taxa do that at the 24-cell stage already. This acceleration has occurred in parallel in the evolution to architaenioglossate (44- to 48-cell stage) and sorbeoconch (40- to 24-cell stage) Caenogastropoda. Together with the other plesiomorphies of the gut (see above) this suggests Ectobranchia as the most basal extant offshoot among the Heterobranchia.</p> <p>Ectobranch synapomorphies are particularly the diagnostic gill (for Orbistestellidae see above) and the pallial tentacles used for exhalant water current. In addition, a couple of sperm characters (see Healy 1993 for details) separate Ectobranchia from Architectonicoidea and all other heterobranch groups. Also the recent study of Dinapoli and Klussmann-Kolb (2010) placed Cornirostra and Valvata (but not Orbitestella) in a single clade close to the origin of Heterobrachia. All current data on the Hyalogyrinidae support the concept of monophyly of the Ectobranchia.</p> <p>The taxon sampling among the ectobranch taxa is still quite poor, and it is likely that several more taxa currently seen as skeneids, vitrinellids or cyclostrematids are in fact ectobranchs. In addition, the Xylodisculidae and several other basal heterobranch taxa (e.g. Tjaernoiidae, Murchisonellidae, Aclididae, Cimidae) remain unstudied. Therefore, an unambiguous phylogenetic analysis of the other families among the Ectobranchia cannot be provided. Nevertheless, Hyalogyrinidae can be reasonably placed at the very basis of Ectobranchia and Heterobrachia, suggesting an origin of Heterobranchia from the rhipidoglossate level of gastropod evolution. However, this does not necessarily mean that the hypothesis of monophyletic Apogastropoda (Caenogastropoda and Heterobranchia) is obsolete: although a synapomorphic taenioglossate condition of the radula is no longer supported, in particular sperm characters (Healy 1993; Ponder and Lindberg 1997) still suggest a common origin of Caenogastropoda and Heterobranchia; the rhipidoglossate sister group for both remains unresolved at the present stage of knowledge. Thus, the story will continue: anatomical, spermatological and molecular studies on more basal heterobranch taxa are needed to provide a robust phylogenetic framework of the origin and early evolution of the Heterobranchia.</p> </div>	https://treatment.plazi.org/id/03DC2538034ABF33FF543FF7B2833ECB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Hess, Gerhard Haszprunar Erika Speimann Andreas Hawe Martin	Hess, Gerhard Haszprunar Erika Speimann Andreas Hawe Martin (2011): Interactive 3 D anatomy and affinities of the Hyalogyrinidae, basal Heterobranchia (Gastropoda) with a rhipidoglossate radula. Organisms Diversity & Evolution (New York, N. Y.) 11 (3): 201-236, DOI: 10.1007/s13127-011-0048-0, URL: http://dx.doi.org/10.1007/s13127-011-0048-0
