Biology and functional morphology of a new species of endolithic Bryopa (Bivalvia: Anomalodesmata: Clavagelloidea) from Japan and a comparison with fossil species of Stirpulina and other Clavagellidae

Abstract. A new species of Clavagellidae, Bryopa aligamenta, from Okinawa, Japan, is described. The species is endolithic in living corals, with the left valve cemented to the crypt wall, as in all clavagellids. The free right valve exhibits an unusual growth pattern, with commarginal lines seemingly arising from the posterior valve margin and extending towards the anterior. This results from: (i) progressive anterior erosion of the umbones, probably as a consequence of the boring process; (ii) the apparent migration posteriorly, as the umbones are eroded, of the dorso‐ventral growth axis of the shell; and (iii) enhanced posterior inter‐commarginal growth. Unlike other clavagellid genera and species, however, there is no discernible primary ligament, at least in the adult. It is possible, however, that if a juvenile ligament were present (as in B. lata), it too would be lost as a consequence of antero‐dorsal erosion during boring. To retain valve alignment in the absence of a primary ligament, and possibly upon reaching an adult size, the mantle lays down alternating layers of calcium carbonate and proteinaceous periostracum onto the interior surface of the shell to thicken it, most noticeably marginally and, especially, posteriorly. The two valves are united dorsally, therefore, by thin layers of periostracum that probably exert a minimal opening force. B. aligamenta is, however, further characterised by large adductor, pallial, and siphonal retractor muscles so that the entire animal is encased tightly within an internally strengthened shell within a crypt. Movement must be minimal, blood being pumped into pallial haemocoels to push open the valves and extend the siphons. Despite a suggestion to the contrary, Bryopa is retained in the Clavagellidae, its unusual growth processes resulting from an endolithic life style within living corals. The fossil clavagellid Stirpulina bacillus, from the Pliocene/Pleistocene of Palermo, Sicily, Italy, was, unlike Bryopa aligamenta and other clavagellids, endobenthic, with a long adventitious tube and anterior watering pot superficially similar to species of Penicillidae, another family of the Clavagelloidea. Furthermore, as in all clavagellids only the left valve is fused into the fabric of the tube, the right being free within it. In all penicillids, both valves are fused into the fabric of their tubes. The watering pots of the fossil S. coronata, S. vicentina, and S. bacillus, moreover, are formed in a different manner to that of penicillids, by progressive encasement of the right valve inside the tube. In penicillids, the tube is secreted in a single event from the general mantle surface and the incorporation of both valves into its fabric. The constituent genera of the Clavagellidae thus constitute an example of parallel evolution with members of the Penicillidae.     

A cadaver unearthed: the anatomy of the Japanese living fossil Stirpulina ramosa (Bivalvia: Anomalodesmata: Clavagellidae) – the unique specimen in the collections of Emperor Shōwa

The only extant, preserved and complete specimen of Stirpulina ramosa has been discovered in the collections of the late Emperor Shōwa (Hirohito) and is described herein. Hitherto, only shells and the adventitious tubes of this species have been described. This study of the anatomy complements previous ones and adds further information as to how the adventitious tube is formed: that is, by the laying down of an organic, periostracum‐like, template that is then biomineralized internally to form the tube and is externally plastered with the clasts that constitute the enclosing burrow wall. Such secretions are produced by pallial lobes that unite on the right side to create the characteristic sutured pleat in this region of the tube. The periostracum enclosing the so‐interred animal is then secreted against the internal template of the tube. The shell of S. ramosa and, as a consequence, the musculature, is greatly disfigured as a result of the incorporation of the left valve into the fabric of the adventitious tube. This valve can, however, continue to grow within the tube, especially posteriorly. The right valve remains free inside the tube but grows only a little more anteriorly. The anterior mantle, with a minute pedal gape, is greatly thickened and secretes the watering pot component of the tube. In many anatomical respects, for example simultaneous hermaphroditism, S. ramosa still reflects the basic anomalodesmatan plan and such modifications as there are from the clavagellid form relate principally to the structure, formation and thus functioning of the adventitious tube. This study of S. ramosa has allowed the full spectrum of clavagellid adaptive radiation to be analysed and an evolutionary picture created which suggests that species of Clavagella/Dacosta and Stirpulina are Mesozoic (Late Cretaceous) remnants. Conversely, species of Bryopa and Dianadema are more modern, Late Oligocene and Palaeocene (Cenozoic), respectively, and possibly evolved in association with the emergence of the Indo‐West Pacific centre of coral diversity, with a postulated average age of just 30 Myr. © 2013 The Linnean Society of London     

Tube construction in the watering pot shell Brechites vaginiferus (Bivalvia; Anomalodesmata; Clavagelloidea)

The bizarre watering pot shells of the clavagellid bivalve Brechites comprise a calcareous tube encrusted frequently with sand grains and other debris, the anterior end of which terminates in a convex perforated plate (the ‘watering pot’). It has not proved easy to understand how such extreme morphologies are produced. Previously published models have proposed that the tube and ‘watering pot’ are formed separately, outside the periostracum, and fuse later. Here we present the results of a detailed study of the structure and repair of the tubes of Brechites vaginiferus which suggest that these models are not correct. Critical observations include the fact that the external surface of the tube and ‘watering pot’ are covered by a thin organic film, on to the inner surface of which the highly organized aragonite crystals are secreted. There is no evidence of a suture between the tube and the ‘watering pot’ or that the periostracum of the juvenile shell passes through the wall of the tube. Live individuals of B. vaginiferus are able to repair substantial holes in the tube or ‘watering pot’ by laying down a new organic film followed by subsequent calcareous layers. Brechites vaginiferus displays Type C mantle fusion, with the result that the whole animal is encased by a continuous ring of mantle and periostracum, thereby making it possible to secrete a continuous ‘ring’ of shell material. On the basis of these observations we suggest that watering pot shells are not extra‐periostracal but are the product of simple modification of ‘normal’ shell‐secreting mechanisms.     

The functional morphology of Mactrinula reevesii (Bivalvia: Mactroidea) in Hong Kong: adaptations for a deposit-feeding lifestyle

The shallow subtidal mactrid bivalve Mactrinula reevesii is a deposit-feeder in the southern and south-eastern oceanic waters of Hong Kong. Buried obliquely, large quantities of fine sediment are taken into the mantle cavity and sorted on enormous labial palps. The small ctenidia probably have little value in collecting material, amounts taken in being too large. The mid gut is long and complexly folded inside the visceral mass. It is also capable of distension, although superficial visceral muscles maintain internal tonus. The rectum is narrow and creates compact faecal pellets.
Most interest is in the ventral mantle margin which is, posterior to the pedal gape and the base of the inhalant siphon, united by a sheet of cuticle. There is no fourth pallial aperture. There are, however, two pairs of flaps extending along the posterior third of the internal ventral mantle surface. These arch over left and right mantle rejection tracts which transfer unwanted material to the base of the inhalant siphon for final expulsion. The mantle flaps prevent such material from being returned to the anterior end of the mantle cavity, for palp reprocessing, when new material arrives. They, thus, maximize sorting efficiency by separating unsorted from sorted and rejected material.
Other mactrids have similar mantle flaps which they use in different ways, including the channelling of unwanted material to a fourth pallial aperture for expulsion, as in Lutraria lutraria. The Mactridae have thus evolved a unique method of increasing the efficiency of pseudofaecal waste rejection which has thereby facilitated the deposit-feeding lifestyle by the diverse representatives of this family.     

Anatomy of Afropomus balanoideus (Mollusca,Gastropoda, Ampullariidae) and its implications for phylogeny and ecology

Summary The hitherto unknown anatomy of Afropomus balanoideus is described. Its systematic position within the Ampullariidae is indicated by the possession of the following autapomorphic characters of the Ampullariidae: presence of a lung sac, an ampulla-shaped aorta anterior, a bipartite copulatory organ of pallial origin, a left-sided zygoneur nervous system, a bipartite kidney, a lamellate osphradium, labial tentacles, an ingestion and an egestion sipho and a radula with strong teeth. The pallial vas deferens is completely closed, while the pallial oviduct is anatomically open in its distal part but functionally closed, because it is embedded in the surrounding mantle tissue. The copulatory organ is of pallial origin and innervated by the right pleural ganglion. The plesiomorphic states of some characters indicate that Afropomus balanoideus might represent the adelphotaxon of the remaining Ampullariidae. The morphological structures show that this species is adapted to an amphibious mode of life and to aestivation, as has been shown for other species of Ampullariidae.     

New freshwater mussels (Bivalvia,Unionida) with potential trigonioidid and hyriid affinities from the Early Cretaceous of Brazil

Two new taxa of freshwater mussels (Bivalvia: Unionida) from the Aptian Crato Formation of the Araripe Basin, NE Brazil, are described. The fossil bivalves are confined to 30- to 130-cm-thick bioturbated mudstones overlying the fossil-rich laminated limestones of the Crato Formation Konservat-Lagerstätte. Individuals are often preserved with closed or splayed articulated valves, some of them potentially in life position, forming an autochthonous to parautochthonous assemblage. Monginellopsis bellaradiata nov. gen., nov. sp. shares key characters with the Trigonioidoidea: (i) the anterior pedal retractor muscle scar is clearly separated from the anterior adductor muscle scar; (ii) the shell has fold-like radial ribs on the posterior half; (iii) a right valve anterior tooth has a striated facet. Araripenaia elliptica nov. gen., nov. sp. is the most abundant and widely distributed unionid of the Crato Formation. Its ornament of anterior inverted V-shaped riblets, and central and posterior radial and sub-radial riblets resembles modern and fossil Hyriidae from the Americas, but also Trigonioidoidea from Eurasia. Its dentition of two smooth anterior pseudocardinals and two smooth posterior laterals in each valve provides no further clues for systematic assignment; muscle scars are not preserved. Assignment to the Hyriidae would make Araripenaia the oldest member of this family known from South America. Moreover, this bivalve assemblage of trigonioidoidids, hyriids, and previously reported silesunionoids suggests palaeobiogeographic links to other areas in both Gondwana and Laurasia.     

A dynamic model for the evolution of sabrecat predatory bite mechanics

The ability of sabretoothed felids to achieve sufficiently high bite forces for predation at extreme gape angles has been the subject of decades of debate. Previous studies have indicated that bite forces in derived sabretoothed felids would have been low, but that they were probably augmented by head depressing muscles. However, bite mechanics is a dynamic process, and mechanical properties change with changes in gape angles. In this study, I present the first comprehensive model of bite mechanics, vector angles, and forces about the temporomandibular joint at gape angles from occlusion to maximal inferred gape in sabretoothed felids. Primitive sabrecats (Machairodus, Paramachairodus) appear broadly comparable to extant large felids (Panthera, Puma), but derived sabrecats in the groups Homotherini (Amphimachairodus, Homotherium, Xenosmilus) and Smilodontini (Megantereon, Smilodon) are often substantially different from either of the former. The ability of the mandibular adductors to generate torque changes with gape angle, indicating that previous models fail to capture potentially important differences in bite function. Inferred muscle sizes and the angles of effective torque from individual adductor fibres in derived sabrecats are different from those of primitive sabrecats and extant large felids, but they had evolved a number of compensatory adaptations for maximizing force output at the canine and carnassial, primarily changes in muscle fibre angles and more compact crania. Inferred outforces at the canines and carnassials were comparable amongst all groups at low gape angles, but at extreme gape angles outforces would have been low, supporting previous hypotheses of head flexor contribution during initial stages of the killing bite in sabrecats. Mandibular adduction in extant carnivores is a complicated pattern of differences in twitch tension and electromyographical activity at different gape angles, and inference of maximal isotonic bite forces from reconstructed mandibular adductor sizes in fossils will give estimates primarily suitable for comparative purposes. Potentially, derived sabrecats could have evolved differences from extant felids in adductor histochemistry or pinnation angle of individual fibres. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 220–242.     

A new species of watering pot shell (Bivalvia: Anomalodesmata: Clavagelloidea) from the Miocene of the Murray Basin,South Australia

Abstract: A new species of penicillid watering pot shell, Kendrickiana coquinacola sp. nov., is described from the middle Miocene (Balcombian) Bryant Creek Formation of the Murray Basin, South Australia. The new species differs from the extant K. veitchi in its smaller size, much shorter posterior tube, fewer tubules in the anterior watering pot structure, absence of the pedal slit, discontinuous dorsolateral bands of pitted muscle scars on the internal surface of the anterior bulb and habit of cementing itself to the shells in its surrounding environment. The fossil record of Kendrickiana is reviewed. The record from the Dry Creek Sands is discounted, while a record for the extant K. veitchi from the earliest Pleistocene of York Peninsula is added. K. coquinacola indicates the highly derived anatomy of the genus evolved over a 10‐Ma period from the late Oligocene through the early Miocene.     

The functional morphology of the abyssal Limopsis cristata (Arcoida: Limopsidae) with a discussion on the evolution of the more advanced bivalve foot

The bivalve Limopsis cristata pursues a semi‐endobenthic life in abyssal soft sediments. It attaches to particles by up to three byssal threads and filter feeds by inhaling water from posterior and anterior directions. Because of partial burial, however, only the latter is functionally significant. Complex ciliary currents in the mantle cavity concerned with the rejection of unwanted particles keep most material out of the simplified intestine. It is generally considered that the ligament is the constraining force in arcoid evolution. This may be true in part, but the lack of pallial fusions and the retention of anterior and posterior inhalant flows are more powerful limits to radiation in the Arcoida. In the deep sea, the Limopsidae has radiated into many micro‐niches through micro‐morphological adaptations. Loss of the arcoid ‘heel’ has resulted in the union of the separate rejectory currents of the visceral mass and foot, creating a single discharge point in more advanced bivalves. This greatly simplifies the rejectory roles of the visceral mass and foot and is thus of functional and selective advantage.     

Ingestion of particulate matter by the outer surface cells of the mollusc mantle

A study of the ingestion of particulate matter from the pallial space located between the shell and the outer surface of the mantle of Isognomon alatus and Pinctada radiata was undertaken with the aid of the electron microscope. For this purpose colloidal thorium dioxide (thorotrast) was introduced into the pallial space for periods of 1–5 days after which time the mantle was excised and prepared for examination with the electron microscope. After 24 hours thorotrast micelles were observed in the pallial space, on the surface of the microvilli, in small pinocytotic vesicles between the bases of the microvilli, in vacuoles undergoing coalescence and finally in large dense bodies (lysosomes). Amoebocytes in the pallial space also participate in the removal of particulate matter in a manner similar to that described for the surface epithelium. During active ingestion the Golgi apparatus changes from a vesicular to a lamellar form. The method of ingestion observed in the surface epithelia and the amoebocytes is similar to the ingestion of protein and other particulate material reported for a variety of vertebrate tissues.     

Extraction and Identification of the Pigment in the Adductor Muscle Scar of Pacific Oyster Crassostrea gigas

In this study, UV (ultraviolet) and IR (infrared radiation) spectral analysis were integrated to identify the pigment in the adductor muscle scar of the Pacific oyster Crassostrea gigas. The pigment was extracted from the adductor muscle scars of cleaned oyster shells that were pulverized, hydrolyzed in hot hydrochloric acid, purified with diethyl ether, and dissolved in 0.01 mL/L NaOH. The maximum absorption of the pigment in the UV absorption spectrum within the range of 190–500 nm was observed between 210–220 nm. The UV absorbance decreased with increasing wavelength which was consistent with the UV spectral absorption characteristics of melanin. In addition, Fourier transform infrared spectroscopy scanning revealed characteristic absorption peaks that emerged near 3440 cm^-1 and 1630 cm^-1, which was consistent with infrared scanning features of eumelanin (a type of melanin). This study has demonstrated for the first time that the pigment in the adductor muscle scar of the Pacific oyster is melanin, hinting that the adductor muscle could be another organ pigmenting the mollusc shell with melanin other than mantle.     

Changes in gape frequency, siphon activity and thermal response in the freshwater bivalves Anodonta cygnea and Margaritifera falcata

Physiologically-driven rhythms in bivalve molluscs are predictedto vary as a function of metabolic rate and temperature, incontrast to genetically predisposed biological clocks. Theserhythms can be evaluated using long-term video monitoring techniquesunder controlled conditions in laboratory aquaria. The bivalvesAnodonta cygnea and Margaritifera falcata were used to evaluatethe effect of temperature on rhythms in gape and the formationof siphons at the mantle edge. Frequency and duration of shellclosure vary with temperature in both species, but with differentresponses. Mean duration of intervals of valve closure decreasesas temperature rises in both species, and is consistent withphysiological limitation by increased biological oxygen demand.For A. cygnea, cumulative gape duration peaks at 25°C, withless time spent closed than at any other temperature, but increasingtemperatures correspond to an increase in gape frequency witha strong increase observed at 31°C. In contrast, frequencyof adduction and valve closure peak at 25°C in M. falcata,and continuous gaping is observed above 29.5°C. This physiologicalstress is consistent with evidence from sclerochronologically-calibratedstable isotope studies of shells, where growth breaks in manymarine taxa coincide with maximum temperatures above 31°Cas derived for ¹⁸O_carbonate. The results of this study suggestthat these growth breaks may be due to physiological limitationsin oxygen uptake and metabolic activity, rather than being adirect consequence of elevated temperature alone. (Received 17 March 2008; accepted 3 October 2008)     

Functional morphology of the Hippuritidae

The hippuritid shell consists of a small operculiform left valve capping a larger cylindrico-conical valve. There are three shell layers: (1) An outer calcitic fibrillar prismatic layer. (2) A crossed-lamellar aragonitic middle layer. (3) A complex crossed-lamellar aragonitic innet layer. There are also traces of myostracal prismatic aragonite which mark former adductor positions, the attachment site of the body mass, which hung largely from within the left valve, and the pallial curtain of the right valve alone. It =ms that water was drawn through the pores on the outer surface of the left valve, into the radial canals which lie within its outer layer. It then flowed outwards from the canal apertures, passing over the broad and radially crenulated right mantle margin, whereon food particles were trapped. The particles were sorted and passed inwards anterieventrally onto the ctenidia and/or palps, which carried them to the mouth. Faeces were ejected via the dorsalmost of two oscules in the left valve, and pseudofaeces via the ventralmost oscule. The form of the adductor muscles and their orientations upon myophores projecting from the left valve suggest immovable valves only separated by a minute gape. These modifications complemented the atrophy of the normal bivalve internal feeding and respiratory current system. La coquille des Hippuritidae comprend we valve gauche operculiforme et une valve droite cylindro-conique. Le test est formé de trois couches: 1. Une couche externe constituée de fibres prismatiques de calcite; 2. Une couche moyenne aragonitique à structure lamellaire entrecroisée; 3. Une couche interne aragonitique à structure lamellaire entrecroisée com-plexe. Les zones d'insertion des muscles adducteurs, de la masse viscérale et des muscles palléaux, montrent en outre des traces de primes myostracaux d'aragonite. Les courants d'eau inhalants pénéraient par les pores de la valve gauche et cheminaient dans les canaux radiaux qui parcourent la couche externe, pour déboucher à la cornissure des deux valves; où les particules alimentaires étaient retenues. Les particules étaient triées et acheminées vers la région antéro-ventrale de la cavité générale en direction des branchieslou des palpes et de h bouche. Les fèces étaient rejetés par l'oscule dorsal de la valve gauche, les pseudo-faxs par l'oscule ventral. La forme courte des muscles adducteurs et leur orientation sur les apophyses myophores de la valve gauche suggère que les valves étaient immobiles et ne laissaient entre elles qu'un faible intervalle. Ces modifications vont de pair avec I'atrophie du système digestif et respiratoire.     

Comparative taphonomy of three bivalve species from a mass shell accumulation in the intertidal regime of North Sea tidal flats

This study comprises a comparative taphonomic analysis of three endobenthic bivalves (Mya arenaria, Cerastoderma edule, and Macoma balthica) derived from a mass accumulation of mainly vertically packed shells from the “Wurster Watt” in Lower Saxony’s Wadden Sea, German Bight. Bulk samples from two transects were analyzed with respect to taxonomic composition, left/right valve presence, counts and weight percentages of taxa, and size-frequency distributions. Taphonomic features including abrasion, fragmentation, encrustation and bioerosion were subjected to a semi-quantitative analysis. Taphonomic results show significant differences with respect to bivalve taxa as well as between transects. Mya arenaria, a large endobenthic bivalve, shows the greatest amount of fragmentation and is often encrusted by balanids and bryozoans. The smaller and more robust Cerastoderma edule has the greatest values for bioerosion especially by the polychaete Polydora ciliata. Macoma balthica, which has the thinnest valves, shows the highest values for abrasion, but low values for the rest of the measured taphonomic features. Taphonomic differences between the taxa, samples and transects are related to (1) the origin of the bivalves (from living populations or exposed colonization horizons), (2) the different size and morphology of the valves (themselves related to mode of life), (3) the taphonomic trajectories of the different bivalves, as well as (4) the varying depositional environment of the two transects.     

Coral preference behaviour by planktotrophic larvae of Spirobranchus giganteus corniculatus (Serpulidae: Polychaeta)

Planktonic larvae of the serpulid polychaete Spirobranchus giganteus, an obligate associate of live coral, were tested for preferences for materials diffusing from natural substrates. Choices offered were Acropora prolifera, a very abundant coral on the Heron Island reef flat; Palauastraea ramosa, a less abundant coral, dead coral rubble and a glass tube as a control. Larval life is approximately 12 days. The larvae tested were 1–4 days old. Adults of S. giganteus occur commonly on A. prolifera and much less frequently on P. ramosa. Experiments were designed to prevent contact between larvae and substrate. Larvae preferred A. prolifera over P. ramosa, rubble and the control. There was no preference expressed between the control and P. ramosa or the control and rubble. A preference by young S. giganteus larvae for a substance diffusing from coral, acting together with a known positive phototaxis, may be adaptive in that it may help to maintain larvae in surface waters over the reef and in the vicinity of a specific coral until they are old enough to settle.     

Monolayer formation and DNA synthesis of the outer epithelial cells from pearl oyster mantle in coculture with amebocytes

Summary In vitro experiments were conducted to clarify the involvement of the epithelium-amebocyte interaction in epithelial regeneration of bivalves. The outer epithelia of the pallial mantle of the pearl oyster, Pinctada fucata martensii, were separated in cell sheets from the inner connective tissue layers by digestion with Dispase. Clumps of the separated mantle epithelia were inoculated onto the amebocyte layers prepared on the bottom of culture dishes and maintained at 20° C in 5% CO₂:95% air for 1 wk. Balanced salt solution with 0.03% (wt/vol) glucose was used as a culture medium. The epithelial cells adhered to the amebocyte layers within 24 h, changed their shape from cuboidal to squamous, and migrated and formed monolayer sheets within 3 d. Electron microscopy confirmed maintenance of epithelial polarity and cell to cell junction in the sheets; 6 d after the inoculation, 5-bromo-2′-deoxyuridine was added to the culture at 30 μM. After labeling for 24 h, the cultures were fixed and stained with anti 5-bromo-2′-deoxyuridine antibody. Cells with immunoreactive nuclei were clearly observed in the epithelial cell sheets, indicating active DNA synthesis in the epithelial sheets. Thus, cocultured with amebocytes, the outer epithelial cells from pallial mantle tissue formed a monolayer sheet and started DNA synthesis. The morphological features of the mantle outer epithelial cells are analogous to those described for the in vivo cutaneous wound healing process, suggesting that the epithelium-amebocyte interaction is important in the regeneration of epithelium in bivalves.     

A novel approach to parasite population genetics: Experimental infection reveals geographic differentiation,recombination and host‐mediated population structure in Pasteuria ramosa,a bacterial parasite of Daphnia

The population structure of parasites is central to the ecology and evolution of host‐parasite systems. Here, we investigate the population genetics of Pasteuria ramosa, a bacterial parasite of Daphnia. We used natural P. ramosa spore banks from the sediments of two geographically well‐separated ponds to experimentally infect a panel of Daphnia magna host clones whose resistance phenotypes were previously known. In this way, we were able to assess the population structure of P. ramosa based on geography, host resistance phenotype and host genotype. Overall, genetic diversity of P. ramosa was high, and nearly all infected D. magna hosted more than one parasite haplotype. On the basis of the observation of recombinant haplotypes and relatively low levels of linkage disequilibrium, we conclude that P. ramosa engages in substantial recombination. Isolates were strongly differentiated by pond, indicating that gene flow is spatially restricted. Pasteuria ramosa isolates within one pond were segregated completely based on the resistance phenotype of the host—a result that, to our knowledge, has not been previously reported for a nonhuman parasite. To assess the comparability of experimental infections with natural P. ramosa isolates, we examined the population structure of naturally infected D. magna native to one of the two source ponds. We found that experimental and natural infections of the same host resistance phenotype from the same source pond were indistinguishable, indicating that experimental infections provide a means to representatively sample the diversity of P. ramosa while reducing the sampling bias often associated with studies of parasite epidemics. These results expand our knowledge of this model parasite, provide important context for the large existing body of research on this system and will guide the design of future studies of this host‐parasite system.     

Direct innervation of the mantle edge gland by neurosecretory axons in Helisoma duryi (Mollusca: Pulmonata)

Summary The mantle edge gland of Helisoma duryi is innervated by neurosecretory axons from the pallial nerves. Synaptoid contacts occur between axons and gland cells, and there is ultrastructural evidence for the release of neurosecretory material. The mantle edge gland contributes to the deposition of periostracum during shell formation, and direct neurosecretory innervation may control shell growth and regeneration.Supported by a National Research Council of Canada Grant (A-4673) and Negotiated Grant D-61     

Partial Root Drying Effects on Biomass Production in Brassica napus and the Significance of Root Responses

Partial root drying (PRD) has been shown to stimulate stomatal-closure response and improve water-use efficiency and thus biomass production and grain yield under water deficiency. While most studies focus on above-ground responses to PRD, we examined how root responses contributed to effects of partial root drying. In particular, in two experiments with oilseed rape (Brassica napus L.) we investigated whether roots were able to forage for patchily distributed water, and how this affected plant growth compared with uniform watering and alternate watering (in which different parts of the roots receive water alternately). The first pot experiment was carried out in the greenhouse and the second outside under a rain-shelter in which also the watering amount was varied. The results indicate that B. napus roots were able to forage for fixed water patches by selective root placement. In the first experiment with small plants, root foraging was equally effective as enhanced water-use efficiency under alternate watering. Both treatments resulted in about 10% higher shoot biomass compared with uniform watering. Alternate watering generally outperformed uniform watering in the second experiment, but the success depended on the time of harvest and the water supply level. Measurements indicated that only the alternate watering regime effectively reduced stomatal conductance, but lead to a higher shoot biomass only under more severe (50%) rather than under milder water deficiency (70% of a well watered control). Water deficiency strongly reduced leaf initiation rates and leaf sizes in B. napus, but for a given level of water supply the supply pattern (uniform control, fixed patchy or alternate watering) hardly influenced these growth parameters. Although also in the second experiment, the plants selectively placed their roots in the wet parts of the pot, root foraging was not as effective as in the first experiment. Possible reasons for these discrepancies are discussed as well as their implications for the application of PRD effects for crop growth.