Functional aspects of shell geometry in some British land snails

Allometric relationships between the area of the shell mouth and live body weight are examined in 19 species of British land snails. Within species, the rate of increase of mouth area on weight is usually less than the isometric expectation, in spite of the logarithmic spiral pattern of growth in most species. It is suggested that this deviation is due mainly to changes in density with size. Two species which conform to isometric expectation alter the geometry of the shell as they pow.
Between species, the rate of increase in mouth area on weight in adults is much greater than isometric expectation, and the range of mouth areas at standard weight is considerable. These deviations are almost entirely accounted for by differences in shell geometry between large and small species, the former having higher rates of whorl expansion and smaller or non–existent umbilicuses.
The range of loading (weight per unit mouth area) on resting adult snails is thus much less than would be expected. It is suggested that large species are limited in the range of possible shell geometries by the need to minimize loading, while in small species other forces such as desiccation and predation may also be important: the range of geometries is generally larger. Observations on ecology and behaviour tend to support these conclusions.     

Variation in Vertebral Number in Galaxiid Fishes (Teleostei: Galaxiidae): A Legacy of Life History, Latitude and Length

Mean vertebral counts among species of Galaxiidae vary curvilinearly with fish size – large species have more vertebrae (pleomerism). The relationship with size breaks down among adults of diadromous species. There is mostly no relationship between vertebral count and body shape, although an inverse relationship between count and body depth in a series of divergent small galaxiids may reflect a shift in swimming mode. Diadromous populations have more vertebrae than non-diadromous (landlocked) conspecifics (around the same number as related non-diadromous species). Diadromous species have more vertebrae than related non-diadromous species and exhibit rising vertebral number with increasing latitude (Jordan's rule applies). Regressions of vertebral number against latitude differ among conspecific populations from different regions, perhaps reflecting different relationships between latitude and local climate. Differences in counts between diadromous and non-diadromous species are not simply a direct influence of environment on ontogeny, as embryonic development of species/populations undertaking the two distinct life history strategies occurs in similar, freshwater habitats; differences seem likely to be evolved, and to reflect an advantage of higher vertebral counts in marine environments, where diadromous species live as juveniles. This conclusion is supported by correlation between vertebral counts and size of juveniles of diadromous species at return from the sea, suggesting selection for vertebral number during marine life. Overall, vertebral count is thus influenced by a complex amalgam of fish size, life history, and environment. The explanation for pleomerism may well be related to the scaling relationship between length and cross-sectional area (area rises by the square of length increase) leading to increasing stiffness with growth – compensated for in larger species by having more vertebrae. This could also be driven by scaling in the surface area of the articular facets of the vertebrae themselves.     

Causes of variation in wing loading among Drosophila species

We examined influences on wing and body size in 11 species (12 strains) of Drosophila. Six measures of wing length and width were closely correlated with wing area and suggested little variation in wing shape among the species. Among ten species wing loading, an important factor in flight costs and manoeuvrability, increased as body mass increased at a rate consistent with expectations from allometric scaling of wing area and body mass to body length. Intraspecific variation in wing loading showed similar relationships to body mass. Density and temperature during larval development influenced wing loading through general allometric relations of body size and wing area. Temperature during the pupal stage, but not during wing hardening after eclosion, influenced wing area independently of body size. Wing area increased as growth temperature decreased. Individuals reared at cooler temperatures thus compensated for a potential allometric increase in wing loading by differentially enlarging the wing area during pupal development.     

Phenotypic homogeneity of two intertidal snails across a wave exposure gradient in South Australia

Dislodgement by the large drag forces imparted by breaking waves is an important cause of mortality for intertidal snails. The risk of drag-induced dislodgement can be reduced with: (1) a smaller shell of lower maximum projected surface area (MPSA); (2) a streamlined shell shape characterized by a squatter shell; and/or (3) greater adhesive strength attained through a larger foot area or increased foot tenacity. Snails on exposed coasts tend to express traits that increase dislodgement resistance. Such habitat-specific differences could result from direct selection against poorly adapted phenotypes on exposed shores but may reflect gastropod adaptation to high wave action achieved through phenotypic plasticity or genetic polymorphism. With this in mind, we examined the size, shape and adhesive strength of populations of two gastropod species, Austrocochlea constricta (Lamarck) and Nerita atramentosa (Reeve), from two adjacent shores representing extremes in wave exposure. Over a 5 day period, maximum wave forces were more than 10 times greater on the exposed than sheltered shore. Size-frequency distributions indicate that a predator consuming snails within the 1.3-1.8 cm length range regulates sheltered shore populations of both snail species. Although morphological scaling considerations suggest that drag forces should not place physical limits on the size of these gastropods, exposed shore populations of both snails were small relative to the maximum size documented for these species. Therefore, selective forces at the exposed site might favour smaller individuals with increased access to microhabitat refuges. Unexpectedly, however, neither snail species exhibited between-shore differences in shape, foot area or foot tenacity, which are likely to have adaptive explanations. Hence, it is possible that these snails are incapable of adaptive developmental responses to high wave action. Instead, the homogeneous and wave-exposed nature of Australia's southern coastline may have favoured the evolution of generalist strategies in these species.     

The intrinsic rates of increase of insects of different sizes

ABSTRACT.

• 1 A negative relationship between intrinsic rate of increase, r, and body size has only clearly been shown using data for species drawn from a number of phyla and covering several orders of magnitude in size. Analyses for more closely related species are equivocal.
• 2 Data for ninety-one species of insects, from nine orders, were used to examine the correlation between intrinsic rate of increase and size.
• 3 Intrinsic rate of increase was negatively correlated with both length and weight across orders, but no relationship could be shown within orders.
• 4 Generation times were positively related to body size, but there was no relationship between net reproductive rate (R_Q) and size.
• 5 These results support the hypothesis that documented relationships between species size and colonization success in insects could be a consequence of the scaling of intrinsic rate of increase with size.

     

A Middle Devonian symbiotic relationship involving a gastropod, a trepostomatous bryozoan, and an inferred secondary occupant

Morris, P. J., Linsley, R. M. & Cottrell. J. F. 1991 01 15: A Middle Devonian symbiotic relationship involving a gastropod, a trepostomatous bryozoan. and an inferred sceondary occupant. Lethaia , Vol. 24. pp. 55–67. Oslo. ISSN 0024–1164.
The high-spired gastropod Palaeozygopleuru (Loxonematacea) of the Hamilton Group (Middle Devonian) in New York State is often entrusted by the trepostomatous bryozoan Leptotrypella (Hetero-trypidae). The form of this enerustation leads us to infer a secondary occupant that dwelled in empty shells of Palaeozygopleura while the bryozoan grew upon them. Encrusted specimens usually have an open aperture, thin or no enerustation on the apertural side of the gastropod shell. and thick encrustation on the abapertural side. While the aperture is invariably open. the columellar and parictal lips of the aperture, and the apertural face of the first whorl are usually encrusted. While the gastropod is alive, this area rests upon the dorsal surface of the foot and the remainder of the shell rests upon the substratc. These encrustation patterns suggest that an occupant of the shell. other than the gastropexl. prevented overgrowth of the aperture and oriented the shell aperture-down. This allowed ahapertural growth of the bryozoan. The presence of thin encrustation on the apertural side of the shell is problematical. It requires either the presence of a secondary occupant capable of holding a thinly encrusted shell off the substrate, or the ability of the juvenile bryozoan colony to extend onto portions of the shell that were in contact with the substrate. The presence of a secondary occupant. such as a sipunculan worm. capable of lifting the shell, but usually resting it aperture-down on the substrate seems the most suitable explanation for the nature of the encrustation. Gastropoda. bryozoa. symbiosis. Devoniun. hermit crab, Sipuncula     

Allometry of gill dimensions in some British and American decapod crustacea

The gill areas and their component measurements of 16 species of North American crab and seven species of British decapod crustaceans have been analysed in relation to body mass using the method of logarithmic transformation.
A wide range of relationships was found, each of which is typical for a given species. The slope, b , of the log/log regression lines varied from 0·5 to 1·0, the lower values being most commonly found in the Macrura. For the North American species the average slope is about 0·8 whereas for British brachyurans the relationship was close to linear (b=0·97).
The analysis shows that the increase in gill surface body size is mainly due to the increasing area for individual platelets or gill lamellae.
Comparison of weight-specific gill areas for animals of the same body weight suggests that the most active species have larger gill areas. For some of these species the values (900mm²/g) approximate to those of active fish.
As plots for interspecific relationships derived from average values for many individual species have slopes which are not typical for any of the component species, it is concluded that caution must be exercized when interpreting such interspecific plots in Allometric studies.     

The effects of exposure and predation on the shell of two British winkles

Changes in shell size and shell shape of the two British winkles Littorina nigrolineata and L. rudis were studied in relation to exposure and to crab-size. In both species, shells from exposed shores are smaller and more globose than those from sheltered shores. Also, in rudis of exposed shores the mouth is relatively wider. In shores of equally sheltered conditions, shells are bigger at those localities where crabs are large than at those localities where they are small. The largest shells are found in those localities where it is extremely sheltered, and the crabs are very large.
It is argued that on exposed shores, small shells are favoured because they have more possibilities than large ones to shelter in crevices and in barnacle interspaces, from the impact of winds and waves. A globose shell could accommodate more foot muscle and thus enable a stronger adherence to the rock; and an increased mouth diameter would increase the area of foot adherence to the rock. On sheltered shores, on the other hand, large, narrow-mouthed shells are favoured because they discourage crab predation, large crabs being abundant mainly on sheltered shores.
The possible significance of shell size and shape in relation to zonation is discussed, in view of the different predatory and physical conditions which prevail in different zones of the shore, and the different shell specializations which these conditions would require.     

Mechanics of locomotion of dogs (Canis familiaris) and sheep (Ovis aries)

Records have been made of the forces exerted on the ground by dogs and a sheep, in walking, trotting, cantering and slow galloping. Film has been taken simultaneously. The difference between walking and trotting was much less marked for the sheep than for the dogs.
Step length and stride length increase as speed increases. They are expressed as functions of the Froude number.
The vertical component of the force exerted by a foot on the ground shows two main maxima in walking, except in the case of the fore feet of sheep. In this case and in other gaits there is only one main maximum. The vertical movements of the fore and hind quarters which occurred in examples of each gait have been calculated from the force records.
The force exerted by a foot on the ground changes direction in the course of a step so as to remain more or less in line with a point fixed relative to the animal, but dorsal to its back.
The force records show impact disturbances in the first 003 sec of contact of each foot with the ground.
The point of application of the force on the sole of a foot tends to move posteriorly as the force increases.
The results are discussed in relation to a theoretical account of the mechanics of locomotion on legs.     

Variations in litter size and reproductive effort within and between some populations of Lacerta vivipara

We studied the relationships between litter size, litter weight, newborn weight, relative clutch mass and the female snout-vent length in some Lacerta vivipara populations over a period of three years.
Litter size and litter weight were positively correlated with female snout-vent length in all the populations for all the years, as in most other lizard species. Relative clutch mass generally increased with female size, though correlations appear not to be very tight.
Considering the two best studied populations suggests that montane females invest less in reproduction than lowland ones.
The main reproductive traits of the species appeared highly variable between as well as within the different populations hitherto studied.
We argue that current theory about lizard reproductive strategy requires, first to work out a good estimate of reproductive effort, and second to get more information about the relations between the species and their environmental, biotic and abiotic conditions.     

Use of tooth impressions to identify and age live Proechimys guyannensis and P. cuvieri (Rodentia: Echimyidae)

A technique whereby impressions of the cheekteeth were used to identify and age live Proechimys guyannensis and P. cuvieri is presented and tested. Based on tooth eruption and occlusal surface wear, 15 age classes were defined. Used in concert with a body measurement, measurements of the length of the toothrow and age class allowed almost all live individuals of the two species to be distinguished. The relationship between mean cheektooth length and hind foot length was most useful for distinguishing relatively old individuals, whereas the relationship between hind foot length or head-plus-body length and toothrow age class was most useful for distinguishing younger individuals. Mean eye lens weight of P. guyannensis increased with toothrow age class, suggesting that the toothrow age classes successfully measured relative age. Individuals of P. guyannensis in lower age classes had juvenile or subadult pelage and did not show signs of reproductive activity, whereas individuals in higher age classes had adult pelage and approximately 50% showed signs of reproductive activity. In the central Amazon, P. cuvieri was found to be proportionally more abundant relative to P. guyannensis in early-successional and edge-dominated habitats.     

Adaptive design of locomotion and foot form in prosobranch gastropods

A comparison of the locomotor types, speed, tenacity, and foot form of nearly 300 species in 52 families of marine prosobranchs has revealed that foot size and shape and even subtle variations of locomotion affect the speed and strength of adhesion to the substratum.Gastropods inhabiting soft substrata move primarily by pedal cilia or by discontinuous locomotion in which shell and foot move alternately. Both types of movement are accompanied by low tenacity. A specialized type of discontinuous locomotion, namely, leaping, surpasses all other methods of movement in speed. Species with ciliary locomotion have a very large foot while those with discontinuous movement have an exceedingly small foot relative to shell size.The majority of prosobranchs inhabit hard substrata, move by continuous pedal muscular gliding, and have moderately high tenacity during movement. Arhythmic pedal locomotion yields lower maximum speeds and tenacities than do rhythmic pedal waves. Foot size and shape relative to shell length in species with arhythmic locomotion vary from very short and broad to long and narrow. Studies of transects at several temperate and tropical marine littoral stations showed that these species are confined to low littoral or sublittoral habitats that are sheltered from heavy wave action. High speed and tenacity are simultaneously attained only by species with rhythmic pedal waves.Speed and tenacity do not represent competing selective pressures on the size and shape of the foot. Speed increases among species as the foot approaches or exceeds shell length and is highest if the foot is also broad; the greatest tenacities are attained by species with a long, broad foot whose dimensions do not exceed that of the shell. The optimal shape for both high tenacity and speed is a broad foot somewhat shorter than the shell; neither speed nor tenacity are much compromised by this form. In general, only species with rhythmic pedal waves whose foot size and shape approximate the optimal form for high tenacity and speed are found in habitats exposed to much wave action. Long rhythmic waves, moving a large proportion of foot area at once, are in theory energetically more economical than small, very rapid waves resulting in the same overall speed, but experiments showed that tenacity is significantly reduced in gastropods which increase speed by enlarging the waves. The optimal wave pattern of a species should be a balance between the demand for speed with the least expenditure of energy, favored by a pattern of many large waves at once, and the demand for tenacity, favored by a pattern of few and small waves.Retrograde ditaxic waves of elongation are the most common pattern encountered among prosobranchs, and are associated with a large range of foot sizes and shapes. Such waves are at least one third as long as the foot, while direct waves and other waves of compression are frequently much smaller. The range of foot forms of species with waves of compression is restricted, tending to be optimal for high tenacity or to be long and narrow. Waves of compression appear to be a specialization with the potential for maintaining high tenacity even at high speeds since the waves can be very small, and for giving superior speed since they can travel very rapidly.     

Expression of recombinant goldfish glutamic acid decarboxylase 65 and evidence for differential pH and PLP responsiveness compared to the human enzyme

Variability of taurine (2-aminoethane sulfonic acid) was studied as a function of size in the mussel Mytilus galloprovincialis and tissue specificity. Isometric and/or allometric relationships were established with regard to total soft mass of the mussels between 20 and 60 mm shell length. Relative amounts of taurine dropped significantly with increasing mass of whole soft tissues with an allometric coefficient value of -0.15. The inverse relationship between taurine and increasing size of mussels was confirmed for gill epithelium and labial palp (allometric coefficient values of -0.16 and -0.10, respectively), tissues that, in turn, represented isometric functions with regard to total soft mass. Although relative amounts of taurine were significantly different in digestive gland, mantle and foot, relationships with increasing size of mussels remained unchanged in these tissues. Gill area of mussels was related to soft mass with an allometric coefficient of 0.70 by 2D Image Analysis, but increased to 0.85 when introducing a third dimension, i.e. gill thickness. Results are discussed according to gill structure analysis and taurine functionality.     

Ecological divergence and talar morphology in gorillas

Gorillas occupy a variety of habitats from the west coast to eastern central Africa. These habitats differ considerably in altitude, which has a pronounced effect on forest ecology. Although all gorillas are obligate terrestrial knuckle‐walking quadrupeds, those that live in lowland habitats eat fruits and climb more often than do those living in highland habitats. Here we test the hypothesis that gorilla talus morphology falls along a morphocline that tracks locomotor function related to a more inverted or everted foot set. This proposed morphocline predicts that gorillas living in lowland habitats may have a talocrural joint configured to facilitate a more medially oriented foot during climbing, suggesting that they may be more adaptively committed to arboreality than gorillas living in highland habitats. To quantify the relative set of the foot in gorillas, we chose two three‐dimensional measurements of the talocrural joint: mediolateral curvature of the trochlea and relative surface area of the lateral malleolus. Our results show that, in comparison to their eastern counterparts, western gorillas have talar features that reflect a more medially directed sole of the foot. This morphology likely facilitates foot placement in a wider range of positions and minimization of shearing stresses across the joint when the foot is loaded on more curved or vertically oriented substrates as occurs during climbing and other arboreal behaviors. In contrast, eastern gorilla talar morphology is consistent with habitual placement of the foot with the sole directed more inferiorly, suggesting more effective loading during plantigrade push‐off on terrestrial substrates. Am J Phys Anthropol 153:526–541, 2014. © 2013 Wiley Periodicals, Inc.     

Life history variation in Pisidium (Bivalvia: Pisidiidae)

Intraspecific variation in the maximum shell length of Pisidium is small in comparison with variation in the life span of the clams, from 4 mo to 4 yr. The use of shell length as a measure of size is complicated by large intraspecific variation in the weight-length relationship, a possible reflection of resource availability on the weight of the soft parts. Maximum embryo length is relatively constant (1 mm) in the majority of species but litter size is variable (1 to 40), generally increasing with parent size. The time of egg-laying is determined by the size of the clam, season, and previous episode of reproduction, while the timing of the release of embryos, depending also on embryonic growth rate, is additionally affected by temperature and oxygen conditions. In brief, the maximum adult and embryo lengths are relatively conservative traits while growth rates vary by an order of magnitude.
Reproductive effort, when measured by the ratio of litter weight to parent weight, may increase or decrease with parent length and age. Both semelparous and iteroparous populations are known in several species. We suggest that if maximum adult length can be reached by the first reproductive season (in a favourable environment) the population is semelparous, otherwise (in an unfavourable environment) the population is iteroparous. Gravid clams in semelparous populations continue to grow, and thus assimilation cannot be meaningfully partitioned into reproduction, somatic growth and maintenance.
Along a depth gradient from 8 to 65 m, maximum and mature shell lengths, embryo length and litter size varies significantly in Pisidium conventus subpopulations not distinguishable electrophoretically. Large clams in deep water produce large embryos but small litters, which is contrary to the general trend of increasing litter size with parent length. Low food availability that decreases juvenile growth rate may restrict litter size in this case.     

Influence of gait parameters on the loading of the lower limb

The ground reaction force which acts on the foot during normal walking consists of the sum of two components: the support of the weight of the body and the acceleration of the body. The relationships between the initial loading rate of the lower limb (ignoring the contribution of the heelstrike transient) and the general gait parameters — cadence, stride length, and velocity — have been examined. Plots of the resultant ground reaction force were used to calculate the loading rate of the limb. A sample of 13 normal male subjects, aged from 18 to 63 years, walked at five different self-selected speeds. Velocity showed the highest correlation with loading rate (r = 0.95) and stride length the lowest (r = 0.85). The relationship between cadence and loading rate was non-linear.     

Size and shape of the mandibular condyle in primates

The relationships between the size of the articular surface of the mandibular condyle and masticatory muscle size, tooth size, diet, and biomechanical variables associated with mastication were studied by taking 12 measurements on skulls of 253 adult female anthropoid primates, including three to ten specimens from each of 32 species. In regressions of condylar length, width, or area against body weight, logarithmic transformations substantially improve the fit of the equations compared with untransformed data. There is a strong relationship between condylar measurements and body weight, with all correlations being .94 or higher. The slopes of the allometric regressions of length, width, and area of the condylar head indicate slight positive allometry with body size. Folivorous primates have smaller condyles than frugivorous primates, and colobines have smaller condyles than cebids, cercopithecines, or hominoids. When colobines are eliminated, the differences between frugivores and folivores are not significant. However, the two species with the relatively largest condyles are Pongo pygmaeus and Cercocebus torquatus, suggesting that there may be a relationship between unusually large condylar dimensions and the ability to crack hard nuts between the teeth. Cranial features having strong positive correlations with condylar dimensions include facial prognathism, maxillary incisor size, maxillary postcanine area, mandibular ramus breadth, and temporal fossa area. These data are interpreted as indicating that relatively large condyles are associated with relatively large masticatory muscles, relatively inefficient mandibular biomechanics, and a large dentition. These relationships support the growing evidence that the temporomandibular joint is a stress-bearing joint in normal function.     

Cortical Structure of Hallucal Metatarsals and Locomotor Adaptations in Hominoids

Diaphyseal morphology of long bones, in part, reflects in vivo loads experienced during the lifetime of an individual. The first metatarsal, as a cornerstone structure of the foot, presumably expresses diaphyseal morphology that reflects loading history of the foot during stance phase of gait. Human feet differ substantially from those of other apes in terms of loading histories when comparing the path of the center of pressure during stance phase, which reflects different weight transfer mechanisms. Here we use a novel approach for quantifying continuous thickness and cross-sectional geometric properties of long bones in order to test explicit hypotheses about loading histories and diaphyseal structure of adult chimpanzee, gorilla, and human first metatarsals. For each hallucal metatarsal, 17 cross sections were extracted at regularly-spaced intervals (2.5% length) between 25% and 65% length. Cortical thickness in cross sections was measured in one degree radially-arranged increments, while second moments of area were measured about neutral axes also in one degree radially-arranged increments. Standardized thicknesses and second moments of area were visualized using false color maps, while penalized discriminant analyses were used to evaluate quantitative species differences. Humans systematically exhibit the thinnest diaphyseal cortices, yet the greatest diaphyseal rigidities, particularly in dorsoplantar regions. Shifts in orientation of maximum second moments of area along the diaphysis also distinguish human hallucal metatarsals from those of chimpanzees and gorillas. Diaphyseal structure reflects different loading regimes, often in predictable ways, with human versus non-human differences probably resulting both from the use of arboreal substrates by non-human apes and by differing spatial relationships between hallux position and orientation of the substrate reaction resultant during stance. The novel morphological approach employed in this study offers the potential for transformative insights into form-function relationships in additional long bones, including those of extinct organisms (e.g., fossils).     

Structure and habits of the 'living fossil' pelecypod Neotrigonia

Neotrigonia (Molluscs, Pelecypoda). the only surviving genus of the once diverse and widespread Trigoniacea, was collected live from depths of 6–80 m on sandy mud bottoms off southeastern Australia. The two species collected. Neotrigonia gemma Iredale, 1924 and Neotrigonia margaritacea (Lamarck), 1804, are structurally similar and differ mainly in size and coloration.
Neotrigonia has such primitive features as an unfused mantle and filibranchiate ctenidia. Its foot is large, active and muscular and the ctenidial ciliary pattern resembles that of the Unionacea. The shell is non-streamlined and has pronounced radial omamentation.
Neotrigonia lives infaunally near the sediment surface, filter feeds. and is capable of fairly rapid burrowing. The burrowing facility, conferred by the foot, is unexpected in light of shell morphology. This suggests that shell morphology alone is not necessarily a reliable guide to the life habits of fossil trigoniaceans.     

Variations in body weight, wing length and condition of Mallard Anas platyrhynchos platyrhynchos and their relationship to environmental changes

This paper describes the analysis of over 15,000 weights of Mallard caught for ringing at Borough Fen Decoy, Northamptonshire, during a 13 year period. Weights are corrected to give condition indices by using wing length to give an indication of body size. The weight of pectoral muscles varies with body weight but as the fat stores increase in weight the proportion of body weight made up of flight muscles decreases. Body weight corrected for size gives a better indication of the bird's condition (defined as its fitness for its present and future needs) than the relative weight of the pectoral muscles or the shape of the breast.
The relationships between food availability and weather, and condition are examined. Only occasionally can a drop in condition following hard weather be detected; the birds normally emigrate to the coast after a few days of adverse conditions. There is a relationship between the condition of Mallard in autumn and the amount of grain left on their stubble feeding grounds after harvest. The implications of variations in winter condition on survival and breeding success are briefly discussed.