On the Fine Structure of the Alimentary Tract of Cephalodiscus gracilis (Pterobranchia,Hemichordata)

The U-shaped alimentary tract of Cephalodiscus is of exclusively epithelial structure; on the basis of fine structural criteria the entire tract can be divided into two large subdivisions: an anterior one with mouth, mouth cavity, pharynx and oesophagus, and a posterior one with stomach and intestine. The anterior subdivision is built up of a relatively uniform, innervated, pseudostratified, ciliated epithelium with mucus cells which are concentrated in the initial parts of the mouth cavity. Cilia and mucus presumably constitute a mechanism transporting food particles into the stomach. In the area of the gill slits specific vacuolated cells occur which may lend rigidity to the walls of the slits. The gastric epithelium consists of prismatic cells characterized by, among others, large inclusion bodies, which may represent digestive vacuoles, small dense rod-shaped granules and an elaborate system of microridges, at the base of which abundant endocytotic vesicles occur. The dorsal gastric pouch contains cells rich in rough ER and secretory granules, probably containing digestive enzymes. Thus morphological evidence points both to intra- and extracellular digestion. The intestinal epithelium resembles that of the stomach, however, it is lower, its organelles are fewer and it bears, beside cilia, mainly microridges, which towards its distal end become sparse. Both in the gastric and intestinal epithelium small granulated cells have been found, which presumably represent endocrine cells.     

The scaphopod foot is ventral: more evidence from the anatomy of Rhabdus rectius (Carpenter, 1864) (Dentaliida: Rhabdidae)

Scaphopods comprise about 900 described species of elongate infaunal molluscs, separated into two orders. The phylogenetic position of this class is contentious, having been proposed as a sister-group to bivalves or alternatively cephalopods, all groups that notably represent dramatic modifications of the molluscan body plan and historical confusion over the fundamental body axes. The digging scaphopod foot was previously considered to be anterior. Here we use a three-dimensional tomographic reconstruction of digestive anatomy and partial dorso-ventral musculature, to test the hypothesis that the scaphopod foot is ventral. Similar to cephalopods, the body orientation is confounded by ano-pedal flexion, but rationalising scaphopods is perhaps further undermined by their infaunal lifestyle, which confounds comparison of ecological life position. Some scaphopods are locally abundant, providing good quality material for anatomical study. In our focal species, Rhabdus rectius (Carpenter, 1864), sexes can reliably be differentiated in vivo by differential colour of the gonad (yellow in females; white in males). The gut is composed of three complete loops. Based on the orientation of the digestive tract and the dorso-ventral muscles, we find further evidence to support the interpretation that the concave side of the scaphopod shell is anterior (the site of the mouth) and the foot is ventral.     

MOSQUITO POLLINATION OF HABENARIA OBTUSATA (ORCHIDACEAE)

The floral morphology and pollination of Habenaria obtusata (Orchidaceae) are described. Habenaria obtusata occurs in Alaska, Canada, Norway, Siberia, etc., in great numbers and is pollinated by mosquitoes of the genus Aedes. The pollination process is described as observed in glass cages.     

Functional morphology of the developing alimentary canal in the holothurian Eupentacta fraudatrix (Holothuroidea, Dendrochirota)

Development of the digestive tract of the holothurian Eupentacta fraudatrix was examined using light and transmission electron microscopy. After the blastopore closes, the gut rudiment loses its connection with the blastoderm and becomes an enclosed, tubular chamber, ending blindly at both ends. The differentiation of the digestive and coelomic epithelia is mainly completed by day 12. Since no transient cell types are observed, this differentiation is definitive. By day 20, the mouth and anal openings appear. The cuticular lining in the anterior part of the gut rudiment has an endodermal origin and differentiates before the mouth is formed. The rest of the gut lining is composed of enterocytes typical of holothuroid intestine. At the early stages of development, mitotic figures are encountered among nonspecialized cells of the gut primordium. In more developed digestive epithelium, vesicular enterocytes are capable of mitotic division. Dividing enterocytes retain secretory vacuoles; thus mitosis occurs in actually differentiated cells. After mouth and anus formation, the oesophagus, stomach, intestine and rectum can be distinguished. In the wall of the stomach, powerful musculature is formed.     

Microanatomy of the digestive system of Supachai's caecilian,Ichthyophis supachaii Taylor, 1960 (Amphibia: Gymnophiona)

The gross anatomy and microanatomy of the digestive system of Ichthyophis supachaii were investigated. The microscopic structures of the digestive system are similar to those in other caecilians. Functional and developing teeth are present in adults. The tongue contains the genioglossus muscle. The digestive tract is elongated and consists of the mucosa, submucosa, muscularis and adventitia/serosa. The oesophagus contains longitudinal folds and lacks oesophageal glands. We report for the first time the caecilian gastric rugae and specific localization of oxynticopeptic cells in the anterior gastric region. The intestinal folds are exclusively present in the anterior intestinal region. The liver comprises 30–40 incomplete hepatic lobes, lying in an imbricate manner. Each lobe is enveloped by haematopoietic tissue that produces and delivers blood cells into sinusoids. Hepatic parenchyma is organized into anastomosing, two‐cell‐thick plates, having sinusoids at the basal domain and bile canaliculi at the apical domain of hepatocytes. Pigment cells are scattered inside sinusoids. The pancreas contains pancreatic acini interspersed with islets of Langerhans. The gallbladder proper is thin and continuous with the cystic duct wall. Neutral and carboxylated acid mucosubstances are secreted along the digestive tract, while sulphated mucosubstances are not produced by the stomach and anterior intestinal regions.     

Differential toxic effects of Ulva lactuca (Chlorophyta) on the herbivorous gastropods,Littorina littorea and L. obtusata (Mollusca)

Members of the genus Ulva are widespread and abundant in intertidal and shallow subtidal areas but there are conflicting data regarding susceptibility to herbivory. While some studies have documented that Ulva spp. were favored by a diversity of marine herbivores, other work has revealed herbivore deterrence. We investigated grazing and growth rates of the littorinid species, Littorina littorea and L. obtusata, when offered Fucus vesiculosus, Ascophyllum nodosum, Ulva lactuca, and Chondrus crispus, highlighting distinctive vulnerabilities to toxic effects of U. lactuca. Ulva lactuca was the preferred food of L. littorea, while L. obtusata showed no grazing on this ephemeral algal species. In contrast, F. vesiculosus was highly preferred by L. obtusata. Although L. littorea demonstrated a grazing preference for U. lactuca, growth rate of this gastropod species was nearly 3× greater when fed F. vesiculosus, suggesting a non‐lethal, negative effect of U. lactuca on L. littorea with long‐term exposure. Mortality of L. obtusata ranged from 0% to 100% when held in the presence of various Ulva densities for 1 week, and Ulva exudate depressed herbivory of this gastropod. We conclude that the water‐soluble, toxic exudate produced by U. lactuca in response to herbivory had allelochemical properties, and may contain a cleavage product (acrylic acid) of dimethylsulfoniopropionate or reactive oxygen species (i.e., H₂O₂). Observed differences in susceptibility to Ulva toxicity by the littorinid species may be related to generalist versus specialist feeding and habitat strategies.     

Effects of ambient temperature and of temperature acclimation on nitrogen excretion and differential catabolism of protein and nonprotein resources in the intertidal snails,Littorina saxatilis (Olivi) and L. obtusata (L.)

Nitrogenous excretion in two snails, Littorina saxatilis (high intertidal) and L. obtusata (low intertidal) was studied in relation to temperature acclimation (at 4° and 21°C), including total N excretion rates, the fraction of urea in N excretion, corresponding O:N ratios and the partitioning of deaminated protein between catabolic and anabolic processes at 4°, 11° and 21°C. Aggregate N excretion rates in both species showed no significant compensatory adjustments following acclimation. Total weight specific N excretion rates at 21°C were higher in standard 3 mg L. saxatilis (739 ng N mg⁻¹ h⁻¹) than standard 5 mg L. obtusata (257 ng N mg⁻¹ h⁻¹) for snails acclimated to 21°C. Comparisons of Q₁₀ values of total weight specific N excretion to Q₁₀ values for weight specific oxygen consumption ({xxV}O₂) between 4° to 11 °C and 11° to 21°C indicated that, while total rates of catabolic metabolism ({xxV}O₂) and protein deamination in L. obtusata were essentially parallel, the relationship between N excretion and {xxV}O₂ in L. saxatilis revealed the partitioning of a larger share of deaminated protein carbon into anabolism at 4° and 21°C than at 11°C. Urea N accounted for a larger share of aggregate N excreted in L. saxatilis than in L. obtusata, but in both species urea N is a greater proportion of total N excreted when acclimated at 4°C (urea N: ammonia N ratio range: 1 to 2.15) than in snails acclimated to 21°C (urea N: ammonia N ratio range: 0.46 to 1.39). Molar O:N ratios indicate that the proportion of metabolism supported by protein catabolism is greater in L. saxatilis (O:N range: 2.5–8.4) than in L. obtusata (O:N range: 7.3–13.0). In both species, regardless of acclimation temperature, the O:N ratios are generally lowest (high protein catabolism) at 4°C and highest at 21°C.     

Anatomy of the cecum of the West Indian manatee,Trichechus manatus (Mammalia,Sirenia)

Summary The anatomy of the cecum of the West Indian manatee (Trichechus manatus) is described at the macroscopic and microscopic levels. The cecum possesses a voluminous corpus ceci with two lateral outpocketings; two diverticulae (cecal horns) extend from the corpus at the vicinity of the ileocecal junction. An ampulla ceci is found distal to the corpus ceci, the two compartments communicating via a wide-opened orifice. The most striking microscopic finding is the mucosal covering of a keratinized stratified squamous epithelium overlying the crypts of Lieberkühn.An attempt is made to interpret the morphological observations in terms of the functional digestive process in the manatee, giving alternative suggestions concerning the significance of the cecum and colon in the nutritional status of the manatee.Abbreviations A Ampulla ceci - CC Corpus ceci - CH Cecal horns - I Ileum - C Colon The present investigation was supported by a grant from the Deutsche Forschungsgemeinschaft (La 325/7-1)The author is greatly indebted to the members of the Fish and Wildlife Service, Sirenia Project, Gainesville Station, Gainesville Florida: special thanks are extended to Dr. Thomas O'Shea for organizing this undertaking, to Cathy A. Beck and Robert K. Bonde for their generous hospitality, for allowing the author to participate in the salvaging project, which they carried out with great efficiency, and for help in collecting the material. Without this aid, this study could not have been completed     

The biology of the Pe-ret' Toad, Otophryne robusta- (Microhylidae), with special consideration of its fossorial larva and systematic relationships

Adult and larval Otophryne robusta- were collected in Colombia, South America. Although this genus has been assigned to the family Microhylidae, a variety of both adult and larval features distinguish it from all other microhylids. The adult is a diurnally active leaf mimic with complex agonistic behaviours and vocalizations. It tends to walk rather than hop, and does not burrow in captivity. The larva is unique in having minute, dagger-like, keratinized teeth and a sinistral spiracle at the tip of a long tube. The mature larva lives shallowly buried in sand at the bottom of clear, shallow streams. The anatomy of the tadpole was examined for clues to how it burrows into and survives in sand, as well as to the correct phylogenetic association of the genus. Internally, the tadpole is most similar to microhylid tadpoles and has a wealth of oral features (e.g. unperforated internal nares, branchial food traps arranged in crescentic organs, ventral velum divided on the midline, glottis located rostral to the free edge of the ventral velum, large gill filters etc.) that characterize that family. Most, if not all, of the unique features of the Otophryne larva can be associated with its fossorial existence. The arrangement of the muscles suggest that it can actively raise and lower its snout during burrowing and possibly dorsiflex its head on its vertebral column. Although the larval teeth first suggest carnivory, many features indicate that maerophagy is not possible. Stomach contents include a variety of bacteria and micro-organisms, but no fragments of macroscopic animals. We conclude that the Otophryne tadpole is a microphagous suspension feeder and its teeth are more important for keeping sand grains out of its mouth than for grasping prey. Since the O. robusta larva does not burrow deeply, the tip of its spiracular tube probably extends upward above the sandy bottom and into the current. In this position pressure would be lower at the spiracle than at the mouth and a current could be passively drawn (by the Bernoulli effect) through the oral cavity for feeding and respiration. If this is correct then the Otophryne larva is the first fossorial vertebrate known to passively filter-feed. All features of the tadpole, except the keratinized teeth and sinistral spiracle, indicate an affinity of Otophryne-/i with South American microhylids. The tadpole and adult are, however, so unusual as to warrant subfamilial status+ADs- we propose the subfamily Otophryninac for this monotypic genus.     

Pharyngeal and Gastrodermal Ultrastructure of Prorhynchus stagnalis Schultze, 1851 (Turbellaria,Lecithoepitheliata)

The digestive tract of a freshwater Lecithoepitheliata turbellarian, Prorhynchus stagnalis, has been studied at the ultrastructural level. The buccal tube connecting the mouth opening to the pharynx is lined by an insunk epithelium, and is provided with two kinds of secretory cells. The bulbous pharynx itself, typical of the genus, is a highly muscular organ, also lined internally by an insunk epithelium; it also bears at least two different types of gland cells. Externally the pharynx is enveloped by a thin, flat epithelium. The gastrodermis consists of phagocytes and Minotian gland cells, as typical for most Tricladida. No ciliary covering was observed along the whole digestive system; instead, phagocytic cells have long microvillar projections. Preliminary attempts to follow the digestive process using external markers and cytochemical reaction for acid phosphatase were successful in demonstrating endocytic activity in the phagocytes. These data, besides being a contribution to morphology and systematics, also speak in favour of the general digestive theory of Turbellaria proposed by Jennings and serve as a starting point for experimental studies on intracellular digestion within the Lecithoepitheliata.     

Feeding behavior,mastication, and tooth wear in the western tarsier (Tarsius bancanus)

We assessed feeding and masticatory function in western tarsiers, Tarsius bancanus,from field study, from videotaped recordings of the feeding and chewing behavior of wild-caught animals in temporary captivity, from dissections of the muscles of mastication, and from scanning electron microscopic (SEM) examination of wear features of the teeth. Ingestion of large items of animal prey is made possible by the animal’s extremely wide gape. Anterior translation of the knob-like mandibular condyle in the anteroposteriorly elongated mandibular fossa makes possible a gape angle of 60–70‡. We observed two means of ingestion of grasshopper prey: ingestion by mastication, in which the postcanine teeth sever and reduce bites of the food as it is thrust into the mouth cavity, and repeated gape-shove sequences, during which the tarsier pushed grasshoppers of large diameter into the anterior part of its mouth and attempted to sever a bite with its anterior teeth. Morsels were successfully severed after three to five such sequences, and reduced quickly,with relatively few powerful, crushing chews. The insect cuticle was not evenly comminuted during mastication. We observed a marked side-to-side grinding component in the normal chewing cycle of T. bancanuson videotape and confirmed it by SEM. The main jaw adductors are bulky, long-fibered muscles that can accommodate wide grapes and still generate, at wide degrees of gape,the high occlusal pressures necessary to fracture thick chitinous exoskeletons of the scarabid beetles that form a substantial element of the western tarsier’s diet.     

Myoanatomy of <Emphasis Type="Italic">Barentsia discreta</Emphasis> (Busk, 1886) (Kamptozoa: Coloniales)

The anatomy of the muscular system of Barentsia discreta (Kamptozoa) was studied by confocal laser scanning and transmission electron microscopy. The calyx musculature, muscles associated with the digestive tract, atrial ring muscles, and tentacle muscles are described. The structure of the muscular bulbus located in the upper part of the stalk and the muscle base of the stalk were examined. The middle part of the stalk and the stolon lack musculature. The structure of the star-cell complex lying at the boundary of the stalk and calyx was examined in detail. Emschermann’s (1969) opinion was confirmed that the star-cell complex performs the function of a heart, providing the transport of substances from the calyx to the stalk and stolon. The general plan of the muscle arrangement is similar in all Kamptozoa; it consists of central muscles of the calyx, atrial ring muscles, tentacle muscles, and muscles associated with the digestive tract. Oral, lateral, and aboral muscles extending from the stalk into the calyx, which were described for solitary forms, are lacking in the calyx of colonial B. discreta. The calyx of B. discreta is separated from the stalk by a septum, through which muscles do not penetrate from the stalk.     

Functional Morphology of the Introvert and Digestive System of Myzostoma cirriferum (Myzostomida)

Myzostoma cirriferum feeds by diverting food particles carried by the ambulacral grooves of its comatulid host Antedon bifida. When searching for food, the myzostome uses its protrusible introvert to fulfil two major functions: sensory perception and the capture of food particles. The digestive system is composed of four parts, viz. a pharynx, that is contained within the introvert, a stomach, a series of paired caeca and an intestine that lie in the myzostome's trunk. The pharynx is supplied with a thick muscle which, thanks to peristaltic movements, carries food particles from the mouth to the stomach. Both stomach and caecal cells are able to absorb dissolved nutriments and to store lipids, whereas intestinal cells are only capable of absorption. Due to the beating of their cilia, stomach cells also carry food particles into the caecal lumen, where they are subjected to endocytosis and intracellular digestion by caecal cells. Undigested food fragments eventually gather in a very large, apical vacuole, and the cell apices containing vacuoles are eliminated into the caecal lumen by an apocrinal process. Detached cell apices reach the stomach, where they are embedded in a matrix, together forming a spindle-shaped faecal mass that is expelled through the postero-ventral anus. The observed digestive process—entailing the regular elimination of the apical part of the caecal digestive cells—appears to be unique among the Spiralia.     

Lack of metabolic temperature compensation in the intertidal gastropods,Littorina saxatilis (Olivi) and L. obtusata (L.)

Two intertidal snails, Littorina saxatilis (Olivi, 1972) (upper eulittoral fringe/maritime zone) and Littorina obtusata (Linnaeus, 1758) (lower eulittoral) were collected from a boulder shore on Nobska Point, Cape Cod, Massachusetts, in July and acclimated for 15–20 days at 4 ° or 21 °C. Oxygen consumption rate (Vo₂) was determined for 11–15 subsamples of individuals at 4 °, 11 ° and 21 °C with silver/platinum oxygen electrodes. Multiple factor analysis of variance (MFANOVA) of lo₁₀ transformed values of whole animal Vo₂ with log₁₀ dry tissue weight (DTW) as a covariant revealed that increased test temperature induced a significant increase in Vo₂ in both species (P<0.00001). In contrast, MFANOVA revealed that temperature acclimation did not affect Vo₂ in either L. saxatilis (P= 0.35) or L. obtusata (P= 0.095). Thus, neither species displayed a capacity for the typical metabolic temperature compensation marked by an increase in Vo₂ at any one test temperature in individuals acclimated to a lower temperature that is characteristic of most ectothermic animals. Lack of capacity for metabolic temperature acclimation has also been reported in other littorinid snail species, and may be characteristic of the group as a whole. Lack of capacity for respiratory temperature acclimation in these two species and other littorinids may reflect the extensive semi-diurnal temperature variation that they are exposed to in their eulittoral and eulittoral fringe/maritime zone habitats. In these habitats, any metabolic benefits derived from longer-term temperature compensation of metabolic rates are negated by extreme daily temperature fluctuations. Instead, littorinid species appear to have evolved mechanisms for immediate metabolic regulation which, in L. saxatilis and L. obtusata and other littorinids, appear to centre on a unique ability for near instantaneous suppression of metabolic rate and entrance into short-term metabolic diapause at temperatures above 20–35 °C, making typical seasonal respiratory compensation mechanisms characteristic of most ectotherms of little adaptive value to littorinid species.     

Function and evolution of the adhesion-capture apparatus ofStenus species (Coleoptera,Staphylinidae)

 Further to basic investigations, additional histological and scanning electron microscopical studies of the anatomy and morphology of the adhesion-capture apparatus in species of the genus Stenus have been carried out to achieve a better understanding of its mode of working and evolution. An interspecific morphological comparison of the sticky cushions (paraglossae) of the labium of 47 Stenus species has established that, in most of the investigated species, they represent a general type that has apparently been modified in different ways during the course of evolution. These changes primarily concern the number of adhesive setae on a cushion and the degree of branching of a single adhesive seta. It has been shown experimentally that larger sticky cushions and higher numbers of adhesive setae or adhesive contacts improve adhesion and thus lead to enhanced capture success. In addition to the morphological comparison of the paraglossae, some general aspects of the functional anatomy and morphology of the labium have been examined. It has been established that, during attack, the sticky cushions and the tips of the sense-spines are directed frontally with respect to the prey. This concurrent erection of both paraglossae and the glossae is conditioned by an exocuticular groin that connects the glossae and paraglossae on the left and on the right. The interior of the sticky cushions is made up of a loose reticulum of endocuticular fibres. This cushion-like endocuticle may absorb the thrust caused by the impact of the sticky cushions on the prey. All tissue and chitinous components within the connecting tube of the labium are surrounded by an extra inner covering that separates the tissue within the labium from the exterior wall of the labium. During the protrusion of the labium, this inner tube is most probably required to direct the haemolymph, flowing into the interior of the labium, towards the front. The hypopha- rynx is situated directly in front of the opening of the mouth and narrows the opening into the pharynx. It may be well suited for retaining large food pieces in front of the mouth in order to prevent their premature movement towards the mouth. Accepted: 12 October 1995     

山东寒武系苗岭统三叶虫消化系统的化石证据

三叶虫是寒武纪演化动物群中最引人注目的成员之一,其内部解剖结构一直以来受到广泛关注.与其他非生物矿化软体结构相比,三叶虫的消化系统更容易留下化石记录,为探索其内部结构提供了难得的机会.本文描述了来自山东省潍坊市寒武系馒头组的Proasaphiscus,Lioparia,Deiradonyx和Iranoleesia,以及...     

The Sphyraena obtusata group (Perciformes: Sphyraenidae) with a description of a new species from southern Japan

Sphyraena iburiensis sp. nov. is described, and taxonomic reviews are provided for S. obtusata and S. pinguis. These species, characterized by having 2 gill rakers, are defined as the S. obtusata group. Sphyraena iburiensis, known only from the Pacific coast of southern Japan, is characterized by 8.5–9.5 scales above the lateral line, a single row of scales in the groove along the lower margin of the suborbital region from the posterior tip of the maxilla to below the eye (=suborbital groove) not covered with skin, 2 distinct longitudinal stripes on the lateral surface of the body when fresh (upper stripe usually lost in preserved specimens), the lower stripe reaching the caudal-fin base just below the lateral line. Sphyraena obtusata, distributed in the Indo-West Pacific, is characterized by 5–7.5 scales above the lateral line, a single row of scales in the suborbital groove covered with skin, 2 somewhat indistinct longitudinal stripes on the lateral surface of the body when fresh (upper stripe usually lost in preserved specimens), the lower stripe joining the lateral line midway between the end of the second dorsal-fin base and caudal peduncle and extending to the middle of the caudal-fin base. Sphyraena pinguis, distributed in the Indo-West Pacific, is characterized by 7.5–9.5 scales above the lateral line, a single row of scales in the suborbital groove not covered with skin, and a single longitudinal stripe on the lateral surface of the body joining the lateral line slightly before or just below the end of the second dorsal-fin base and extending to the middle of the caudal-fin base. Seven (S. aureoflammea, S. brachygnathos, S. flavicauda, S. grandisquamis, S. langsar, S. lineata, and S. strenua) and 2 (S. chrysotaenia and S. schlegelii) nominal species are regarded as junior synonyms of S. obtusata and S. pinguis, respectively. In addition, lectotypes are designated for S. flavicauda, S. langsar, S. lineata, and S. obtusata. A key to the three species of the S. obtusata group is provided.     

THE MODE OF POLLINATION IN HABENARIA OBTUSATA (ORCHIDACEAE)

The role of mosquitoes and moths as pollinators of Habenaria obtusata was studied in northern Wisconsin during the summer of 1969. In each population studied, meter-square quadrants were marked and the number of plants, flowers per inflorescence, rate and time of pollinia removed, and capsule set recorded. Carbon dioxide and modified New Jersey Light Traps were set during the flowering period to capture pollinia-bearing insects. The study showed that female mosquitoes of the genus Aedes (particularly A. communis) and two species of Geometrid moths of the genus Xanthorhoe are important pollinators of H. obtusata. Pollination occurred during the last few days of flowering with an average of 18.0% of the pollinia removed and with 14.2% capsule set.     

Molecular evidence for the taxonomic status of three species of the <Emphasis Type="Italic">Sphyraena obtusata</Emphasis> group (Perciformes: Sphyraenidae) from East Asia

Genetic differences among the three species of Sphyraena with two gill rakers from East Asia (S. iburiensis, S. obtusata, and S. pinguis, defined recently as the S. obtusata group), were investigated using 799-bp sequences of the mitochondrial cytochrome b gene. Pairwise sequence differences within each of the three species were 0.0–0.4% (S. iburiensis), 0.0–0.4% (S. obtusata), and 0.0–0.6% (S. pinguis), respectively, pairwise sequence differences among the three species being 18.3–18.6% (S. iburiensis vs. S. obtusata), 14.9–15.4% (S. iburiensis vs. S. pinguis), and 17.6–18.3% (S. obtusata vs. S. pinguis), respectively. The extent of the latter were comparable to those among four other congeneric species, S. africana, S. forsteri, S. helleri, and S. japonica (16.0–24.5%). Phylogenetic analysis using the maximum-likelihood method indicated reciprocal monophyly of the three species, each clade being supported by a bootstrap value of 100%. These findings fully supported the taxonomic status of these species, recently elucidated by detailed morphological comparisons. In addition, the S. obtusata group formed a strongly supported clade against the four other congeneric species.