Thermal tolerance,evaporative water loss,air-water oxygen consumption and zonation of intertidal prosobranchs: a new synthesis

Duration of emergence increases with tidal height on rocky shores therefore, emergence adaptations in intertidal species such as littorine and other prosobranch gastropods have been considered correlated with zonation patterns; temperature tolerance, desiccation resistance and aerial respiration rate all commonly assumed to increase progressively with increasing zonation level. Such direct correlations are rarely observed in nature. Maximal aerial gas exchange occurs in mid-shore, not high shore species. Temperature tolerance and desiccation resistance do not increase directly with shore height. Thus, hypotheses regarding physiological correlates of zonation require revaluation. A new hypothesis is presented that the high tide mark presents a single major physiological barrier on rocky shores. Above it, snails experience prolonged emergence and extensive desiccation; below it, predictable submergence and rehydration with each tidal cycle. Thus, desiccation stress is minimal below the high tide mark and maximal above it. Therefore, species restricted below high tide (the eulittoral zone) should display markedly different adaptive strategies to emergence than those above it (the eulittoral fringe). A review of the literature indicated that adaptations in eulittoral species are dominated by those allowing maintenance of activity and foraging in air including: evaporative cooling; low thermal tolerance; elevated aerial O₂ uptake rates; and high capacity for radiant heat absorption. Such adaptations exacerbate evaporative water loss. In contrast, species restricted to the eulittoral fringe display adaptive strategies that minimize desiccation and prolong survival of emergence including: foot withdrawal, preventing heat conduction from the substratum; aestivation in air; elevated thermal tolerance reducing necessity for evaporative cooling; position maintenance by cementation to the substratum and increased capacity for heat dissipation. In order to test of this hypothesis the upper thermal limits, tissue and substratum temperatures on emergence in direct sunlight and evaporative water loss and tissue temperatures on emergence in 40 °C were evaluated for specimens of six species of eulittoral and eulittoral fringe gastropods from a granite shore on Princess Royal Harbour near Albany, Western Australia. The results were consistant with adaptation to the proposed desiccation barrier at high tide. The eulittoral species, Austrocochlea constricta, Austrocochlea concamerata, Nerita atramentosa and Lepsiella vinosa, displayed adaptations dominated by maintenance of activity and foraging during emergence while the eulittoral fringe littorine species, Bembicium vittatum and Nodilittorina unifasciata displayed adaptations dominated by minization of activity and evaporative water loss during emergence. The evolution of adaptations allowing tolerance of prolonged desiccation have allowed littorine species to dominate high intertidal rocky shore gastropod faunas throughout the world's oceans.     

Cold,freezing, and desiccation tolerance of the limpet Acmaea digitalis (Eschscholtz)

Cold, freezing, and desiccation tolerance were examined in the limpet, Acmaea digitalis (Eschscholtz). Fifty percent of the experimental population survived freezing at temperatures between ?10 ° and ?12 °C for at least 24 hr. In this temperature range, 60–80 % of the body water was frozen. The LD₅₀ for water loss by desiccation was 76.5% ± 1.3_δ. At 70–80% body water loss, solutes were concentrated between 350 and 500%. Some limpets failed to survive immersion in 450% seawater for 6 hr (100% scawater = salinity of 31‰), suggesting that much of the damage from freezing and desiccation was from an increased solute concentration. No glycerol could be detected in extracts of the foot and the visceral mass of A. digitalis during winter, confirming similar results in studies carried out on other intertidal molluscs.     

Movement patterns of the sub-Antarctic pulmonate limpet Kerguelenella lateralis (Gould) (Mollusca: Gastropoda)

Summary Kerguelenella lateralis produces distinct spots on microalgal-coated rocks in the high intertidal zone at Kerguelen. Spots are the result of microalgal removal, corresponding in outline to the limpet's shell. Although limpet spots could be associated with homing behavior, K. lateralis does not exhibit this behavior. Limpets in low-and high-intertidal areas show high levels of sitespecific emigration and immigration. Rates of site-emigration (No. limpets leaving/day) are positively related with limpet density indicating that local limpet densities may influence population movements. Distances of displacement and rates of emigration and immigration are highly variable and do not differ in low-and high-intertidal sites. Lack of homing in K. lateralis may reflect characteristics of its sub-Antarctic environment: low levels of predation, few competitors, and protection from desiccation and wave action through occupation of cracks in the rocky substratum.     

The tenacity of the limpet, Patella vulgata L.: An experimental approach

It has been shown that adhesion of the limpet, Patella vulgata L. is influenced by both physical and physiological factors. The tenacity is sensitive to surface properties of the substratum, varying inversely with the contact angle which water makes with a substratum. This can be explained in terms of thermodynamics. Surface roughness also affects tenacity and this is explained in the same manner. Different angles of detachment were tested and it was clearly shown that when a strong peeling component was introduced, a much reduced force was needed to detach a limpet. Contrary to a normal pull, when a shear pull is exerted the force is not proportional to the surface area of the foot. It has also been shown that the speed of separation affects the measured tenacity; there is a speed at which tenacity will be maximum. The effect of water temperature on tenacity has been tested, tenacity increasing with rising temperature (7, 13, 20 °C). At the higher temperatures limpets are able to contract the foot muscles more powerfully, indicating that increased foot rigidity increases tenacity. By measuring the tenacity of limpets left out of water for different periods of time it has been shown that desiccation has no effect on tenacity, but a change from aquatic to aerial respiration increases tenacity. Tenacity has also been measured when the limpets have been subjected to a reduction in metabolic rate. The effect of both anoxia and narcotization shows that reduced muscle tonus, especially in the foot, results in decreased tenacity. These results further demonstrate that foot rigidity is essential for efficient adhesion. Eimpets from different habitats (exposed and sheltered) and vertical distribution (high and low level on shore) exhibited no differences in tenacity. During locomotion limpets leave a mucous trail, most of the mucus being confined to the edge of the trail. Water is incorporated anteriorly under each new locomotory wave and these pockets of water are used to release the mucus from the substratum during locomotion. It is concluded from this study that limpet adhesion can be explained solely by the tackiness of the pedal mucus, tack being due to the stored elastic energy within the mucous layer itself.     

The influence of desiccation and predation on vertical size gradients in populations of the gastropod Oxystele variegata (Anton) on an exposed rocky shore

Summary Oxystele variegata (Anton.) exhibits a vertical size gradient contrary to the model proposed by Vermeij (1972) for low/mid intertidal species, as shell size increases in an upshore direction. Settlement occurs in the lowest zones and juveniles are restricted to the lower shore by conditions of desiccation higher up the beach. Juveniles suffer rapid water loss due to a relatively large opercular surface area and circumference and have a much lower resistance to water loss than adults. This leads to high mortality under conditions of low humidities, and juveniles caged at the top of the balanoid zone, where adults normally occur, die within a few days. As animals increase in size their resistance to desiccation rises allowing them to migrate upshore. This is a response to high rates of predation by the whelk Burnupena delalandii in the lower balanoid zone. Predation is so intense as to override the advantages of higher food availability which lead to a greater body weight for adults protected by cages on the lower shore.     

Predator-induced defenses in a salt-marsh gastropod

Predator-induced defenses are among the most ecologically important forms of phenotypic plasticity. Although predation and induced defenses are well documented in rocky-intertidal systems, they have received less attention in soft-bottom communities. Shell-crushing predators are common in soft-bottom, vegetated habitats, which often exhibit substantial spatial heterogeneity in predation intensity. We examined variations in shell morphology of the salt-marsh periwinkle, Littoraria irrorata, among marsh microhabitats in the northern Gulf of Mexico that vary in their accessibility to predatory blue crabs, Callinectes sapidus. Littoraria from high-predation sites exhibited more extensively calcified apertural lips and narrower apertural openings relative to snails from low-predation sites. Thick apertural lips generally increased the handling time required by Callinectes to breach Littoraria shells in laboratory experiments, although the method of shell entry used by crabs was dependent on the crab:snail size ratio. Apertural-lip thickness was not related to past predation events in field-collected snails. Snails exposed to water treated with the effluent of Callinectes and crushed conspecifics produced significantly thicker apertural lips than controls, with a response time and morphological extent comparable to that of their rocky-shore counterparts. This study underscores the widespread occurrence of predator-induced plasticity in marine gastropods and emphasizes its role in soft-bottom, vegetated marine habitats, where shell-crushing predation can be as prevalent a selective force as in the rocky intertidal.     

The functional significance of aperture form in gastropods

Aperture form of marine prosobranch gastropods has evolved under the influence of a number of different selective forces, including: generation of shell form; protection from predation; accommodation of the foot during clamping behavior: and accommodation of water currents in and out of the mantle cavity. Aperture form correlates positively with foot shape in most gastropods and foot shape, in turn, correlates moderately well with substrate preference. Almost all gastropods that have non-round apertures elongate the aperture parallel to the foot so that water currenth tend to flow anteriorly to posteriorly. Fresh-water pulmonates have responded to somewhat different stresses. They exhibit clamping behavior and thus show correspondence between foot shape and aperture shape. They show less apertural strengthening as crab (or crayfish) predation is less of a factor and presumably because calcium carbonate is less available. They also lack anterior-posterior apertural elongation due to the absence of water currents through their mantle cavity. Due to the absence of mantle cavity water currents and clamping behavior, terrestrial gastropods do not show the apertural modifications associated with these two factors. In addition. few adaptations of apertural form are present to resist predation. Instead, many of the apertural modifications of terrestrial pulmonates seem to be concer-ned with the problems of water loss during estivation.     

Shell form,behaviour, and tolerance to water movement in the limpet Patina pellucida (L.) (Gastropoda: Prosobranchia)

The diotocardian gastropod Patina pellucida (L.) is a macro-herbivore common upon the fronds of laminarian algae. Considerations of shell shape, adhesive tenacity, and current tolerance, together with calculations of expected hydrodynamic drag, indicate that this limpet is effectively streamlined. Smaller individuals are relatively more resistant to dislodgement than are larger ones. Experimental animals of 6.0–13.5 mm shell length resisted constant flows of up to 0.9-1.3 m s^?1. An orientation, in which the longitudinal axis of a limpet is aligned parallel to the direction of the prevailing flow, is induced by currents faster than ≈ 0.5 m s^?1, and is an adaptation to conditions of directional water streaming. P. pellucida appears to show a seasonal change in orientatory preference, individuals tending to point distally with respect to the algal lamina in spring and proximally in autumn: this change may be associated with the annual growth cycle of the host plant.     

Changes in chloroplast morphology of different parenchyma cells in leaves of Haberlea rhodopensis Friv. during desiccation and following rehydration

The size, shape, and number of chloroplasts in the palisade and spongy parenchyma layers of Haberlea rhodopensis leaves changed significantly during desiccation and following rehydration. The chloroplasts became smaller and more rounded during desiccation, and aggregated in the middle of the cell. The size and number of chloroplasts in the palisade parenchyma cells were higher than in spongy parenchyma. The good correlation observed between the size or number of chloroplasts and the cross-sectional area of mesophyll cells, the cross-sectional width of the leaf and its water content suggested that the palisade cells were more responsive to water availability than the spongy cells. Changes in chloroplast number during desiccation and rehydration process are characteristic features for desiccation-tolerant plants (especially in homoiochlorophyllous strategy).     

Reproduction and larval development of the limpet <Emphasis Type="Italic">Lottia persona</Emphasis> (Rathke, 1833) (Gastropoda: Lottiidae)

The reproduction and larval development of the sea limpet Lottia persona (Rathke, 1833) were investigated in vitro for the first time. The limpets breed in late July; they have external fertilization and a pelagic lecithotrophic type of development. The larval shell is transparent, symmetrical, and bottle-shaped, with well-pronounced lateral fossae and a large rounded operculum. The protoconch sculpture is characterized by broad wavy lines and radial ribs at the dorsal side. Ventrally, the lines become narrow and are directed perpendicular to those of the dorsal and lateral regions. Development from fertilization to settling lasts 3 days at a water temperature of 19–20°C.     

The relationship between position on shore and shell ornamentation in two size-dependent morphotypes of Littorina striata,with an estimate of evaporative water loss in these morphotypes and in Melarhaphe neritoides

Shells of small (< 7 mm) Littorina striata are frequently nodulose, but shells of larger individuals are striate. Nodulose L. striata dominated the littoral fringe of a black basalt Azorean shore where daytime rock temperatures rose significantly higher than nearby shores of different rock composition or colour. There was no evidence of intraspecific size-partitioning on the latter shores, where the numbers of striate and nodulose L. striata were approximately equal between high eulittoral (low-shore) and high littoral fringe (high-shore) localities. The prevalence of small L. striata in the littoral fringe is opposite to that usually characteristic of the Littorinidae, where the largest individuals usually occupy the higher positions on the shore. It is hypothesized that small L. striata attain a resting posture better able to minimize heat absorption from the substratum than attained by larger individuals. Smaller individuals also take advantage of both posture and a nodulose shell surface to re-radiate absorbed incident radiant thermal energy more effectively to the atmosphere by convection. Thus, small, nodulose L. striata are especially well adapted to occupy geologically young basaltic rocks commonly found fringing islands of the mid-Atlantic. The rate of evaporative water loss was determined for Melarhaphe neritoides and striate and nodulose L. striata for approximately 11 days emersion. All three groups are exceptionally capable of controlling evaporative water loss. Total percent evaporative water loss by nodulose L. striata (17.9%) was significantly greater than that lost by either striate L. striata (14.1%) or M. neritoides (13.5%) but, among 15 species for which evaporative water loss has been determined by similar methodology, M. neritoides and striate L. striata are the most capable of conserving body water during 11 days of emersion.     

Ascaris sp.: water loss during desiccation of embryonating eggs

The ability of embryonating eggs of Ascaris lumbricoides to avoid desiccation by reducing the loss of water through the egg shell was investigated. When exposed to desiccation the eggs lost water at a rate dependent upon the relative humidity and ambient temperature, eventually resulting in the collapse of the eggs and the death of the enclosed embryo. The eggs are small with a large surface to volume ratio. A low permeability to gaseous exchange thus restricts water loss while still ensuring an adequate supply of oxygen for embryonic development. Relative humidity did not appear to affect the rate of development. In eggs exposed to desiccation at various constant temperatures, the rate of water loss increased as an exponential function of increasing temperature. When eggs were exposed to various temperatures before exposure to desiccation at 22 C, the rate of water loss increased as a function of increasing pretreatment temperature. After exposure to 63–65 C, the ability of the egg shell to slow down the loss of water was destroyed. These phenomena suggest that there is not a simple “critical” or “transition” temperature, but a gradual melting of the complex mixture of components forming the lipid layer.     

Defensive responses of gastropods to the predatory flatworms Freemania litoricola (Heath and McGregor) and notoplana acticola (Boone)

Three species of gastropods were tested in the laboraory and in the field for responses to contact with the polyclad flatworms Freemania litoricola (Heath and McGregor) and Notoplana acticola (Boone). The three limpets differed considerably in their responses to Freemania litoricola All specimens of Notoacmea scutum (Rathke) responded in some way to this flatworm, mostly by running. Fewer than half of the Collisella digitalis (Rathke) responded to Freemania litoricola, and these responded by mushrooming - elevating the shell several millimeters off the substratum. Collisella scabra (Gould) did not respond any differently to contact with Freemania litoricola than to contact with a control probe. Differences in the responses of these limpets to Freemania litoricola are discussed in relation to the characteristic intertidal distributions of this flatworm and the limpets. In contrast to Freemania litoricola, the polyclad Notoplana acticola typically did not elicit defensive responses. This difference is related to the diets of the two flatworms. Freemania litoricola was found to be a predator on limpets, whereas no evidence was found that Notoplana acticola eats limpets.     

Alginate production by Pseudomonas putida creates a hydrated microenvironment and contributes to biofilm architecture and stress tolerance under water-limiting conditions

Biofilms exist in a variety of habitats that are routinely or periodically not saturated with water, and residents must integrate cues on water abundance (matric stress) or osmolarity (solute stress) into lifestyle strategies. Here we examine this hypothesis by assessing the extent to which alginate production by Pseudomonas putida strain mt-2 and by other fluorescent pseudomonads occurs in response to water limitations and how the presence of alginate in turn influences biofilm development and stress tolerance. Total exopolysaccharide (EPS) and alginate production increased with increasing matric, but not solute, stress severity, and alginate was a significant component, but not the major component, of EPS. Alginate influenced biofilm architecture, resulting in biofilms that were taller, covered less surface area, and had a thicker EPS layer at the air interface than those formed by an mt-2 algD mutant under water-limiting conditions, properties that could contribute to less evaporative water loss. We examined this possibility and show that alginate reduces the extent of water loss from biofilm residents by using a biosensor to quantify the water potential of individual cells and by measuring the extent of dehydration-mediated changes in fatty acid composition following a matric or solute stress shock. Alginate deficiency decreased survival of desiccation not only by P. putida but also by Pseudomonas aeruginosa PAO1 and Pseudomonas syringae pv. syringae B728a. Our findings suggest that in response to water-limiting conditions, pseudomonads produce alginate, which influences biofilm development and EPS physiochemical properties. Collectively these responses may facilitate the maintenance of a hydrated microenvironment, protecting residents from desiccation stress and increasing survival.     

Grazing patterns in Siphonaria gigas (Mollusca,Pulmonata) on the rocky Pacific coast of Panama

The pulmonate limpet Siphonaria gigas is the most abundant molluscan grazer in the mid zone on rocky, wave-exposed shores of the Pacific coast of Panama. Erect macroalgae and sessile invertebrates are rare; crustose algae cover ～90% of the rock. The relative abundance of a common blue-green algal crust (Schizothrix calcicola?) is negatively correlated with Siphonaria's abundance. Large-scale removals of the limpet cause rapid increases in percent cover of Schizothrix and concomitant decreases in other crusts, but no changes in the abundance of erect algae or sessile invertebrates. Removing Siphonaria also (1) increases recruitment of crustose algae and barnacles onto new rock and plexiglass substrata, and (2) decreases the abundance of a calcified form of Schizothrix. Harsh conditions during daytime low tides and foraging by fishes at high tide control the microdistribution of most of this region's mobile and sessile benthic organisms. Wave action and substratum heterogeneity modify these constraints: Siphonaria is rare or absent in sheltered areas, especially on homogeneous surfaces, and is most abundant at wave-exposed sites. However, at extremely wave-beaten sites, Siphonaria and other benthic consumers are rare and ineffective. Crustose algae are reduced in abundance and space is dominated by erect macroalgae and/or barnacles. These normally rare species can outcompete crusts only when thermal or desiccation stress and the effects of benthic consumers and fishes are drastically reduced.     

Shell Morphological Variation of Littoraria angulifera among and within Mangroves in NE Brazil

The shell morphological variation of the periwinkle Littoraria angulifera (Lamarck, 1822) was studied in tropical northeast Brazilian mangroves. This area was selected because mangroves in different stages of regeneration, and thus different tree heights can be found. We evaluated whether differences in solar radiation due to differences in tree height influenced the distribution and shell morphology of L. angulifera, and carried out an experiment to test if individuals collected from mangroves with different tree heights differed in their resistance to desiccation. We also analysed if there were differences in L. angulifera shell length and shape between tidal levels within a mangrove. Finally, we tested if increased habitat complexity due to the presence of oysters could influence L. angulifera shell length and shape in different-statured mangroves. We predicted that the oysters could reduce desiccation stress on periwinkles in small-statured mangroves but have no effect in taller ones. Shell length and shape varied among mangroves, although a large variation within mangroves was also recorded. Shell proportionality (shell length:width ratio) increased with shell length, and this relationship differed among mangroves; however, no differences were found in a subsequent year. Individuals from small-statured mangroves survived longer than those from taller mangroves in the desiccation experiment, with a weak correlation between shell proportionality and loss of mass. The presence of oysters had no apparent influence on shell morphology that could be correlated with reduced desiccation stress. The patterns found varied greatly both at small and large spatial scales, suggesting that future studies should evaluate phenotypic and genetic variation at the same time to properly understand variation in L. angulifera shell morphology.     

Selective site segregation in Patelloida (Chiazamea) Pygmaea (Dunker) and P.(C.) Lampanicola Habe (gastropoda: Patellacea) on a Hong Kong shore

Although originally described as a separate species, Patelloida (Chiazacmea) lampanicola Habe has subsequently been regarded as a “form” of P. (C.) pygmaea (Dunker). The conical shape of the shell in the former was assumed to result from the settlement and growth of some individuals of the population upon shells of intertidal Batillaria (Potamididae).This paper reports upon an analysis of a population of limpets resident on a small sand flat in Hong Kong and it is concluded that Patelloida (C.) lampanicola is distinct from P. (C.) pygmaea. This conclusion is based upon differences in radula teeth structure, shell morphometrics and behaviour. Furthermore it has been demonstrated that the earlier confusion surrounding these species results from a very nice example of selective site segregation. P. (C.) pygmaea inhabits stones and empty shells (in this region of its total range) embedded in the sand but can occasionally does reside, up to a length of 4.2 mm, upon Batillaria shells. Thereafter the flattened form of the limpet shell, on a round substratum, presumably makes such individuals more susceptible to either predation or dislodgement. The high-coned Patelloida (C.) lampanicola, on the other hand, almost exclusively colonizes living Batillaria (particularly B. zonalis and B. multiformis) shells but can, with little or no significant alteration in form, also colonize stones. Patelloida (C.) lampanicola has a positive behavioural response towards Batillaria and is clearly living in symbiotic association with the potamid. The benefits accrued by both are discussed.Thus, the limited hard substratum of a primarily soft shore environment has been segregated by these two limpets; one occupies the stones and oyster shells, the other the dense cover of epifaunal potamid snails.     

Geographical variation of heat sensitivity and water loss rates in the tropical lizard,Anolis gundlachi

• 1.1. Sensitivity to high temperatures and desiccating conditions were separately measured in four samples of Anolis gundlachi collected over an altitudinal transect and one sample of lowland A. cristatellus.
• 2.2. Critical thermal maxima (CTM) of gundlachi samples varied inversely in perfect rank order with elevation from a low of 33°C at 850m to a high of 36°C at 270m. The data suggest a possible genetic basis of the variation. No other measures of heat sensitivity exhibited significant altitudinal variation in gundlachi.
• 3.3. Rates of evaporative water loss among gundlachi samples exhibited the same pattern of population differentiation as did the CTMs. Survival times also varied among samples, but vital limits of desiccation did not.
• 4.4. A. cristatellus was invariably more heat and desiccation resistant than its montane counterpart, A. gundlachi.

     

Intraspecific trait cospecialization of constitutive and inducible morphological defences in a marine snail from habitats with different predation risk

1.?Studies examining the integration of constitutive and inducible aspects of multivariate defensive phenotypes are rare. 2.?I asked whether marine snails (Nucella lamellosa) from habitats with and without abundant predatory crabs differed in constitutive and inducible aspects of defensive shell morphology. 3.?I examined multivariate shell shape development of snails from each habitat in the presence and absence of waterborne cues from feeding crabs (Cancer productus). I also examined the influence of constitutive and inducible shell morphology on resistance to crushing. 4.?Regardless of the presence of crabs, snails from high-risk (HR) habitats developed rotund, short-spired shells, while snails from low-risk habitats developed elongate shells, tall-spired shells, indicating among-habitat divergence in constitutive shell shape. Moreover, allometry analyses indicated that constitutive developmental patterns underlying this variation also differed between habitats. However, snails from HR habitats showed greater plasticity for apertural lip thickness and apertural area in the presence of crab cues, indicating among-habitat variation in defence inducibility. 5.?Both shell shape and apertural lip thickness contributed to shell strength suggesting that constitutive shell shape development and inducible lip thickening have evolved jointly to form an effective defence in habitats where predation risk is high.     

Aspects of the population ecology of Lucinomaborealis (L.) (Bivalvia) in Raunefjorden,Western Norway

2194 Lucinoma borealis (L.) were obtained from 99 quantitative (0.2 m²) samples of substratum (stratified random sampling) collected between November 1975 and October 1976 in the archipelago of Eggholmane ≈20 km south of Bergen. The depth range of the 20900 m² area was 0.3–13.3 m, and the bottom sediment (fine to very coarse sand) was rich in calcium carbonate. The highest density was found in the deep (≈ 8–13 m) sub-areas, with fine sediment (median grain-size diameter ≈ 0.3–0.5 mm). L. borealis was the most abundant bivalve species in the area studied (59.6%, by number). The average density was 22.2 (C.I. ± 5.4) ind. · 0.2 m ^?2, and the biomass was 690 (C.I. ± 116) mg AFDW · 0.2 m ^?2. The average size of animals found in shallow water (coarse sediment), with low abundance, was markedly larger than that of those from deeper water. Adults from shallow water consisted of 49.5% females, but in deep water the corresponding figure was only 38.6%.According to the data, spawning in deep water occurred in June, August–September, and possibly also in January–February. The spawning period in shallow water was more diffuse and uncertain. The egg diameter of 123.75 (C.I. ± 1.59) μm, indicates a lecithotrophic development. Maturation occurred at a shell length of 10–13 mm, and the largest animal found was 34.3mm long. The shell of L. borealis grows allometrically.