Introduction to Physidae (Gastropoda: Hygrophila); biogeography, classification, morphology

Physidae, a world-wide family of freshwater snails with about 80 species, are reclassified by progressive characters of the penial complex (the terminal male reproductive system): form and composition of penial sheath and preputium, proportions and structure of penis, presence or absence of penial stylet, site of pore of penial canal, and number and insertions of penial retractor muscles. Observation of these characters, many not recognized previously, has been possible only by the technique used in anesthetizing, fixing, and preserving. These progressive characters are the principal basis of 23 genera, four grades and four clades within the family. The two established subfamilies are divided into seven new tribes including 11 new genera, with diagnoses and lists of species referred to each. Proposed as new are: in Aplexinae, Austrinautini, with Austrinauta g.n. and Caribnautu harryi g.n., nom.nov.; Aplexini; Amecanautini with Amecanauta jaliscoensis g.n., sp.n., Mexinauta g.n., and Mayabina g.n., with M. petenensis, polita, sanctijohannis, tempisquensis spp.nn., Tropinauta sinusdulcensis g.n., sp.n.; and Stenophysini, with Stenophysa spathidophallus sp.n.; in Physinae, Haitiini, with Haitia moreleti sp.n.; Physini, with Laurentiphysa chippevarum g.n., sp.n., Physa mirollii nom.nou.; and Physellini, with Chiapaphysa g.n., and C. grijalvae, C. pacifica spp.nn., Utahphysa g.n., Archiphysa g.n., with A. ashmuni, A. sonomae spp.nn., Physella hemphilli sp.n., and Ultraphysella sinaloae g.n., sp.n. The simplest reproductive system is found in Austrinauta of the Aplexinae; its penial complex approaches that in the related family Lymnaeidae. Within Physinae a close approximation is found in Haitia. By these two genera the two subfamilies are drawn close together. Four grades of progressive complexity are recognized: (I) penial sheath entirely muscular; (II) penial sheath with both glandular and muscular tissue; (III) penis with penial stylet or other specialization of the tip of the penis; and (IV) pore of penial canal lateral rather than terminal as in the lower grades. In both subfamilies there are clades with glandular tissue in the penial sheath, a penial sheath subdivided into two parts, and tip of penis specialized in various ways. These clades are formalized as new tribes. Of 23 genera of Physidae, 17 occur in Pacific drainages of North and Central America, eight of these restricted to the region. Concentration of primitive genera along the Pacific coast from Mexico to Costa Rica conforms to previous observations that primitive pulmonate families are concentrated within, or along the continental margins of, the Pacific Ocean. An ancestral origin of Physidae along an ancient eastern Pacific coast is probable. From this region the several lineages have spread north, south and east in the Americas, and through Siberia to Europe. Although Physinae have fewer genera than Aplexinae (11 v. 12), they have more species (47 v. 34). Greater land area in the temperate zone has provided more opportunity for speciation of Physinae, in contrast to the generally tropical and warm-temperate range of Aplexinae. Furthermore, 10 species of Physinae are localized in individual lakes, whereas Aplexinae are not lake-dwellers. Both well-developed egg strings and capsular strings are found in the spawn of Sibirenauta elongatus. These structures have been known in Lymnaeidae, but not hitherto in Physidae; they are a link with some marine groups, such as Siphonariidae. Spiral color bands and white streaks in the shell of Mexinauta recall those in Lancidae (Lymnaeacea), whereas the radula of Physidae is like that of Chilinidae. Physidae thus show affinities to various basal stocks of aquatic pulmonates; no clear-cut sister-group can be recognized. Most species have a restricted range; out of 55 with sufficiently detailed information for analysis, 25 are limited to a single 1 degrees x 1 degrees quadrangle. Only a few species are widespread, on one or even two continents. Accordingly, more species of Physidae are threatened by habitat destruction than in other families of Hygrophila with generally wider distributions. Other features are a key to genera; catalog of more than 430 names applied to living Physidae, with original reference, type locality, and location of type specimens; summary of museums with types; and glossary.     

云南管螺属一新种记述(肺螺亚纲,柄眼目,烟管螺科)

记述采自云南省石林县陆生贝类1新种,即石林管螺Phaedusa shilinensis sp.nov.,文中对新种形态特征、栖息环境进行了描述,并对其相似种进行了分析和讨论。     

RADULAR PRODUCTION RATES IN TWO SPECIES OF LACUNA TURTON (GASTROPODA: LITTORINIDAE)

The molluscan radula is a dynamic organ, both in terms of itsuse and production. New rows of teeth are constantly producedat the posterior end of the radula, while older, worn teethare shed anteriorly, producing a dynamic equilibrium. We useda cold-shock to mark the radular ribbon and measure tooth rowproduction rates in two gastropod species, Lacuna vincta (Montagu)and L. vanegata Carpenter. We found that the average tooth rowproduction rate at 10–11°C did not differ betweenthese two species, and was 2.94 (SE = 0.002) rows per day forLacuna vincta and 2.97 (SE = 0 002) for L. vanegata Inter-individualvariability in production rate was very low, and was correlatedwith shell length, smaller individuals had slightly higher productionrates. The total length of the radular nbbon varied greatlyamong individuals, ranging from 47 to 94 (2.57 to 5.68 mm) rowsin L vincta and 53 to 99 rows (2.80 to 7.14 mm) in L vanegata,and was only somewhat correlated with the length of the shelLThis great variability will result in large differences amongindividuals in the time it takes to replace the radula totally,from 14.96 to 35.44 days in L vincta and from 17 43 to 39 69days in L. vanegata. (Received 1 September 1995; accepted 20 November 1995)     

Redescriptions of Phallodrilus parthenopaeus Pierantoni and P. obscurus Cook (Oligochaeta, Tubificidae)

The type species of Phallodrilus Pierantoni, 1902, P. parthenopaeus Pierantoni, 1902; is redescribed on the basis of new material from W Scotland. The species is characterized by its specific spermathecal and penial setae, its very slender atria, and its long spermathecal ducts. Sexually fully mature specimens of P. obscurus Cook, 1969 are described for the first time, on the basis of material from New Jersey (E USA). The new material shows that this species possesses 5 to 9 penial setae in each ventral bundle of segment XI, and that its atria are coated with a relatively thick musculature.     

Action and immunoreactivity of neuropeptides in the buccal neuromuscular system of a prosobranch mollusc,Rapana thomasiana

Summary In a prosobranch mollusc, Rapana thomasiana, the catch-relaxing peptide H-Ala-Met-Pro-Met-Leu-Arg-Leu-NH₂ (CARP) was found to depress the contraction of the radula protractor and retractor elicited by electrical stimulations. The action of CARP was in contrast to that of other neuropeptides, H-Phe-Met-Arg-Phe-NH₂ (FMRFamide) and H-Phe-Leu-Arg-Phe-NH₂ (FLRFamide), which enhanced the contraction of the radula protractor and retractor, respectively. By immunohistochemical examinations, FMRFamide-like immunoreactive neurons were found on the rostral side of the right buccal ganglion and the caudal side of the left ganglion, where some CARP-like immunoreactive neurons were also distributed, indicating a possible coexistence of FMRFamide and CARP. FMRFamide- and CARP-like immunoreactivities were also detected in the neuropile of buccal ganglia, radula nerves arising from the ganglia, and nerve fibers in the radula muscles. The present results suggest that FMRFamide- and CARP-like peptides are involved in the regulation of the contraction of the radula muscles.     

A new species of Sesara Albers, 1860 from northeastern Thailand (Stylommatophora: Helicarionidae)

Sesara triodon n.sp. is described from the Phu Pha Lom limestone area in Loei, northeastern Thailand. It possesses a small and depressed–conoid, transparent shell with a shiny external surface. The aperture possesses three apertural teeth, in contrast to other Thai species, Sesara parva Solem, 1966 (two teeth) and Sesara megalodon Blanford, 1902 (four teeth). The genital system of the new species has some distinctive characters, including a vagina longer than the free oviduct, and the epiphallic retractor caecum is rounded and bulbous. The radula has 104 rather straight rows of teeth, with each row having74–81 teeth. This is the easternmost record of the genus Sesara in Thailand.

http://zoobank.org/urn:lsid:zoobank.org:pub:52EF0AB6-1E17-4B96-BC6F-4257FE5FBF00     

Age determination and estimation of larval period in field caught abalone (Haliotis discus hannai Ino 1953) larvae and newly metamorphosed post-larvae by counts of radular teeth rows

We developed an age determination method for larval and newly metamorphosed post-larval abalone Haliotis discus hannai in a laboratory experiment and determined the age of field caught individuals. Laboratory experiments showed that competent veliger larvae (4 days after fertilization) had a radula and regularly added rows of radular teeth with age in the absence of metamorphosis. Under environmentally relevant temperatures (17-22 °C), the number of rows of radular teeth increased linearly with age, but slopes of the regression lines were different among temperatures. Rows of radular teeth were added more slowly at lower temperatures. The effect of temperature on the development rate of the radula was quantified by the regression and the temperature coefficient, Q₁₀. The radular development of newly metamorphosed post-larvae, which had not acquired a peristomal shell (adult shell), was comparable with that of veliger larvae, although older post-larvae had a larger number of rows of radula than those of the same age of veliger larvae. From these results, an age determination method of veliger larvae and newly metamorphosed post-larvae was established, using the number of rows of radular teeth. The age of veliger larvae and newly metamorphosed post-larvae was determined by the age determination method for samples collected in August to October of 2003 and 2004 for which the thermal history of the coastal water of Miyagi Prefecture Japan was available. Only 9.1% of veliger larvae (n = 8) captured in the field had formed a radula and these were estimated to be 4-6 days old. The remaining 90.9% of larvae (n = 80) that had not formed a radula were classified as younger than 4 days old. All newly metamorphosed post-larvae (n = 24) that had metamorphosed on substrata were estimated to be 4-6 days old. Results of the field study indicate that these abalone metamorphosed within a few days after the acquisition of competence (4 days after fertilization) at this site, which has suitable crustose algal habitat.     

Aeolid nudibranchs (Gastropoda: Opisthobranchia) of the family Glaucidae from New Zealand waters

Two new species of aeolid, a new genus being created for one of them, are described along with another two species, one Phidiana militaris (Alder & Hancock) (= Learchis militaris ) a new record for New Zealand, the other, Glaucus atlanticus , found only once before. Jason mirabilis gen. et sp. nov. is distinguished by a ventral tongue-like process with a ridged chitinous covering which replaces the radula (still present but vestigial) and by the greatly subdivided rhinophoral lamellae. Babaina caprinsulensis sp. nov. differs from its Californian relative in having papillate rhinophores. The New Zealand specimens of Phidiana militaris differ slightly in colouration from those described for elsewhere in the Indo-Pacific.
The Glaucidae is enlarged to include the Babainidae, Facelinidae and Favorinidae, the main familial characteristics being the simple oral glands, sharp radular tooth and pugnacious behaviour. Possible evolutionary trends within the family based on the arrangement of the cerata and penial structure are traced out. The Babaininae, pleuroproctic with a notal ridge and irregular disposition of the cerata, is considered the most primitive subfamily and the Hervellinae, with cerata in single rows, the most advanced. The many facelinid genera previously recognized are reduced to two, Antionetta Schmekel and Phidiana Gray.     

Original molluscan radula: comparisons among Aplacophora,Polyplacophora, Gastropoda,and the Cambrian fossil Wiwaxia corrugata

As the original molluscan radula is not known from direct observation, we consider what the form of the original radula may have been from evidence provided by neomenioid Aplacophora (Solenogastres), Gastropoda, Polyplacophora, and the Cambrian fossil Wiwaxia corrugata (Matthews). Conclusions are based on direct observation of radula morphology and its accessory structures (salivary gland ducts, radular sac, anteroventral radular pocket) in 25 species and 16 genera of Aplacophora; radula morphogenesis in Aplacophora; earliest tooth formation in Gastropoda (14 species among Prosobranchia, Opisthobranchia, and Pulmonata); earliest tooth formation in four species of Polyplacophora; and the morphology of the feeding apparatus in W. corrugata. The existence of a true radula membrane and of membranoblasts and odontoblasts in neomenioids indicates that morphogenesis of the aplacophoran radula is homologous to that in other radulate Mollusca. We conclude from p redness of salivary gland ducts, a divided radular sac, and a pair of anteroventral pockets that the plesiomorphic state in neomenioids is bipartite, formed of denticulate bars that are distichous (two teeth per row) on a partially divided or fused radula membrane with the largest denticles lateral, as occurs in the genus Helicoradomenia. The tooth morphology in Helicoradomenia is similar to the feeding apparatus in W. corrugata. We show that distichy also occurs during early development in several species of gastropods and polyplacophorans. Through the rejection of the null hypothesis that the earliest radula was unipartite and had no radula membrane, we conclude that the original molluscan radula was similar to the radula found in Helicoradomena species.     

A new species of Gudeodiscus Páll-Gergely, 2013 from China,with extraordinary conchological and anatomical features (Gastropoda,Pulmonata, Plectopylidae)

A new species of the Plectopylidae, Gudeodiscus longiplica is described from northern Guangxi Province, southern China. The shell, anatomical and radular characters are figured and described. This new species is characterized by long plicae on its parietal shell wall, which have not been observed in any other Gudeodiscus species. In contrast, the long parietal plicae are characteristic for the genera Plectopylis and Chersaecia, which mainly inhabit Thailand and Myanmar. These two genera are, however, only distantly related to the new species, as other characters (anatomy, protoconch sculpture, parietal plicae) suggest. The male portion of the genital structure of the new species is characterized by two separate penial caeca with different lengths, but similar in outer and inner structure. The relevance of this anatomical character is discussed. Gudeodiscus longiplica sp. n. occurs sympatrically with Gudeodiscus soosi Páll-Gergely, 2013. The anatomy and radula characters of the latter species are also described and figured.     

不同滴灌方式下棉花生物量和产量的水氮调控效应

通过3个水平的灌水量和施氮量（低、中、高）的田间试验,研究了田间不同滴灌方式下棉花生物量和产量的水氮调控效应.结果表明：在1带4行、2带4行、2带6行滴灌模式下灌水量由低（分别为90、140、140 mm）到中（分别为150、200、200 mm）时,地上部干物质量分别提高9.2%、37.9%和23.5%,籽棉产量分别提高19.1%、14.1%和16.0%;灌水量由中到高（分别为210、260、260 mm）时,地上部干物质量分别提高15.8%、19.1%和16.7%,籽棉产量分别提高7.7%、11.2%和9.5%.施氮量由低（67.6 kg·hm^-2）到中（95.2 kg·hm^-2）时,地上部干物质量2带4行模式提高14.3%,籽棉产量1带4行模式提高22.2%,其他模式无显著变化;施氮量由中到高（122.8 kg·hm^-2）时,籽棉产量3种模式分别提高7.4%、13.9%和9.9%,地上部干物质量无显著变化.与1带4行和2带6行模式相比,2带4行模式地上部干物质量和籽棉产量的水氮调控效应更明显,相同水氮处理下2带4行地上部干物质量和籽棉产量均高于2带6行和1带4行.表明2带4行是最有利于滴灌棉花田间水氮管理的模式.     

Phylogeny and biogeography of the Vitrinidae (Gastropoda: Stylommatophora)

The phylogeny of the Vitrinidae is reconstructed in a cladistic analysis based on characters of the genitalia, the copulation behaviour and the radula. The genera with an atrial stimulator turned out to be the earliest branches of the Vitrinidae, whereas the genera with a glandula amatoria form a monophyletic, taxonomically apomorphic group. The differences between the proposed phylogeny and previous hypotheses are discussed. The ancestral areas of the Vitrinidae and its sister group, the limacoid slugs Boettgerillidae–Limacidae–Agriolimacidae, are estimated using weighted ancestral area analysis. The Vitrinidae and the limacoid slugs might have originated by a vicariance event between Central Europe and the Near East. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 134 , 347–358.     

The morphology and taxonomy of Womersleya monanthos, an endemic species and genus of Delesseriaceae (Ceramiales, Rhodophyta) from southeastern Australia

Womersleya monanthos (J. Agardh) Papenfuss is typically an epiphyte of larger brown and red algae that are common in drift along the southeastern coasts of Australia. A hitherto little-known member of the Phycodrys group of the Nitophylloideae, its reproductive features have been studied in detail and its taxonomic position clarified. Blades are polystromatic throughout and lack veins or nerves, with blades originating from apical cells of primary and second-order cell rows. Intercalary cell divisions take place in primary cell rows and all other branch orders, with third-order laterals arising both abaxially and adaxially on cells of second-order rows. Fertile central cells bear procarps on pericentral cells on both sides of the blade, the procarps consisting of two 4-celled carpogonial branches and a single central group of sterile cells that enlarge and persist at the distal end of a bicampanulate fusion cell at maturity. Spermatangia and tetrasporangia form in circular subapical sori on both sides of the blade or in marginal lobes or proliferations. After comparing it to other members of the Phycodrys group, we conclude that Womersleya is a monotypic genus well distinguished from other genera and with probable closest affinities to the Northern Hemisphere Polyneura, Erythroglossum and Sorella, as well as the Australian endemic, Crassilingua.     

The relation between excitation-contraction coupling and fine structure of a molluscan muscle,the radular retractor of the whelk,Buccinum undatum

The Buccinum radula is of the rachiglossate type with two outer rows of fierce hook-like attack teeth and a medial row of straight sharp-pointed shredding teeth. Individual cells of the radular retractor muscle are 10–12 m in diameter and separated at the closest by gaps of only 40 nm, providing areas of potential electrical contact. The cell membranes are heavily invested with long finger-like invaginations, associated with sarcoplasmic reticular cisternae, and surface caveolae; the latter are associated with the numerous dense body membrane attachment plaques found in this muscle. The radular retractor muscle possesses a significant sarcoplasmic reticulum of peripheral cisternae and deeper vesicles associated with mitochondria. The surface caveolae may result from myofilament force exerted via attachment plaques at the cell membrane, while deeper invaginations may constitute a rudimentary transverse tubular system to relay surface depolarization to associated sarcoplasmic reticular cisternae inducing calcium release to effect excitation-contraction coupling. The radular retractor muscle possesses the usual thick paramyosin and thin actin myofilaments, the latter associated with dense bodies and attachment plaques presumably to transduce force to the cell membrane. The mitochondria are unusually large and packed into dense central clusters surrounded by large deposits of glycogen granules. The nerve endings on the radular retractor muscle fibres show four different types of transmitter vesicle, presumably related to the four kinds of agonist action in this muscle, cholinergic, serotonergic, peptidergic and purinergic. All nerve endings have mixed vesicle populations, clear evidence of co-transmission. In this muscle we see a modification of usual smooth muscle structure to effect fast sustained contractions, an ultrastructural configuration functionally designed for the muscle's central role in the feeding cycle.Abbreviations ABRM anterior byssus retractor muscle - EC coupling excitation-contraction coupling - RP radular protractor muscle - RR radular retractor muscle - SR sarcoplasmic reticulum - T-system transverse tubular system     

Seasonal aberrant radular formation in Thais bronni (Dunker) and T. clavigera (Küster) (Gastropoda : Muricidae)

Radulae of Thais bronni (Dunker) and T. clavigera (Küster) were examined at Mukaishima Island for a period of 2 yr, 1982 to 1984. Radulae of both species are similar in morphology, both having the basic pentacuspid rachidian plan. Sexual dimorphism of the radula was not observed, but rachidian tooth changes morphologically in different growth stages. Seasonal conditions affect the size and shape of the radula; in winter it is clearly malformed and strikingly thin. These aberrant parts of the radula comprised some dozens of rows, in which only several extremely thin rows exist. Results of experiments using T. clavigera under different water temperature conditions showed that the radula is rarely produced below 10 °C and that rate of radular production and replacement increases with increase in temperature. These results suggest that in the field the radula of these species is replaced entirely 2–2.5 times per year and 10–15 times during the life of the animal.     

Ammoniten mit Kieferapparat und Radula aus Lias-Geschieben

Jaws and radula have been found in two adult microconch specimens of the Liassic ammonite speciesEleganticeras elegantulum (Young & Bird, 1828) from glacial drift boulders near Hamburg. They are more similar to those of recent dibranchiates than ofNautilus, the radula having seven longitudinal rows of teeth instead of thirteen in recentNautilus. Since bothOwens classification of the cephalopods into Dibranchiata and Tetrabranchiata according to the number of gills (which cannot be observed in fossil cephalopods) andSchwarz’ division into Ecto- and Endocochlia according to the (assumed) relative position of body and shell are rather unfortunate, the radula is suggested as a sound basis of classification. It is acceptable to paleo- as well as to neozoologists. As yet, the known evidence is sufficient for a twofold division only: Class Cephalopoda Subclass Lateradulata: Nautiloidea Subclass Angusteradulata: Ammonoidea Coleoidea     

Revision of three camaenid and one bradybaenid species (Gastropoda,Stylommatophora) from China based on morphological and molecular data,with description of a new bradybaenid subspecies from Inner Mongolia,China

We have revised the taxonomy of three camaenid and one bradybaenid species from China and described one new subspecies of the genus Bradybaena (Family Bradybaenidae) from Inner Mongolia, China. The genitalia of three Satsuma (Family Camaenidae) species S. mellea stenozona (Moellendorff, 1884), S. meridionalis (Moellendorff, 1884), comb. n. and S. uncopila (Heude, 1882), comb. n. assigned to the genus Bradybaena previously,lack a dart sac and mucous glands. Moreover, the molecular phylogeny has revealed close relationships between the three species and the genus Satsuma. Two species, S. stenozona (Moellendorff, 1884) from Fuzhou and Ganesella citrina Zilch, 1940 from Wuyi Mountain, are considered as synonymous and should be a subspecies of S. mellea mellea (Pfeiffer, 1866) because of the morphological and molecular similarities. Meanwhile, the other two are placed in the genus Satsuma: S. meridionalis (Moellendorff, 1884), comb. n. and S. uncopila (Heude, 1882), comb. n. G. virgo Pilsbry, 1927 differs from species of the genera Ganesella and Satsuma not only in its shell, but also in anatomical characters, such as having a dart sac and mucous gland, and lacking a flagellum. Additionally, phylogenetic analyses highly support the sister relationship with other Bradybaena species. Thus, placement of G. virgo Pilsbry, 1927 in the genus Bradybaena issuggested.     

A peculiar new species in the genus Landouria Godwin-Austen, 1918 from China (Gastropoda: Heterobranchia: Stylommatophora: Camaenidae)

Landouria omphalostoma n. sp. is described from northern Yunnan, China. The new species is placed in the genus Landouria Godwin-Austen, 1918 based on the presence of tuberculated flagellum and swollen basal part in the bursa copulatrix, and on the absence of dart sacs and mucous glands. L. omphalostoma n. sp. is characterized by a horizontal aperture of the shell which is unique in the genus. This feature resembles those of only two other camaenid/bradybaenid taxa, species of the Chinese genus Pseudaspasita Möllendorff, 1902 and species of the Japanese genus Coelorus Pilsbry, 1900. This is the first verified record of the genus in China and the easternmost record of Landouria. This paper provides the taxonomic ground work for further studies of the repeated evolution of a horizontal aperture.