Enhancement of the response of rock crabs, Cancer irroratus, to prey odors following feeding experience

The rock crab, Cancer irroratus Say, uses chemically mediated learning in the search for food. Rock crabs are opportunistic benthic predators and scavengers. Observations indicate that although they eat a variety of items, they are more sensitive to, and prefer, odors of food items that they have been eating. We found that C. irroratus is more responsive to a familiar food source than to an unfamiliar one and can distinguish between the odors of two different prey after being fed one species for an extended time. Initial preferences for two mytilid bivalves, Mytilus edulis and Geukensia demissa, were determined in a Y-maze. Crabs were then fed only one of the mussel species for 28 days and retested, using sequential and simultaneous presentations, for their responses to familiar and unfamiliar prey odors. Crabs increased their responses to familiar prey odors, but not to unfamiliar odors. In foraging tests, crabs ate M. edulis more often regardless of the species to which they had been familiarized.     

Effect of meal type on specific dynamic action in the green shore crab,Carcinus maenas

The effect of meal type on specific dynamic action was investigated in the green shore crab, Carcinus maenas. When the crabs were offered a meal of fish, shrimp, or mussel of 3 % of their body mass the duration of the SDA response and thus the resultant SDA was lower for the mussel, compared with the shrimp or fish meals. In feeding behaviour experiments the crabs consumed almost twice as much mussel compared with fish or shrimp. When the animals were allowed to feed on each meal until satiated, the differences in the SDA response were abolished. The mussel was much softer (compression test) than the fish or shrimp meal, and meal texture is known to affect the SDA response in amphibians and reptiles. When the crabs were offered a meal of homogenized fish muscle or whole fish muscle, the SDA for the homogenized meal was approximately 35 % lower. This suggested that a significant portion of the SDA budget in decapod crustaceans may be related to mechanical digestion. This is not unexpected since the foregut is supplied by over forty muscles which control the cutting and grinding movements of the gastric mill apparatus. There were slight, but significant differences in protein, lipid, moisture and total energy content of each meal type. Three prepared meals that were high in either protein, lipid or carbohydrate were offered to the crabs to determine if the nutrient content was also a contributing factor to the observed differences in the SDA. The crabs did not eat the prepared meals as readily as the natural food items and as they are messy feeders there was a large variation in the amount of food eaten. The lack of significant differences in the SDA response as a function of nutrient content was likely due to differences in amount of food eaten, which is a major factor determining the SDA response. The differences in SDA when consuming natural food items were likely due to a combination of the costs of mechanical digestion, variation in nutrient content and food preference: determining how each of these factors contributes to the overall SDA budget remains a pressing question for comparative physiologists.     

Hermit crabs, humans and Mozambique mangroves

There is a complex interrelationship between upper shore hermit crabs (such as Coenobita sp. and Clibanarius sp.), coastal human populations and mangrove forests in Mozambique. The abundance, activity, shell selection and behaviour of three species of hermit crab are related to the level of mangrove cover. With increased density of mangrove trees, the study species of hermit crab changed in abundance, tended to become diurnal, spent more time feeding and were clustered in larger groups when doing so, and selected longer spired shells. All five of the same variables are also linked to the proximity and activity of humans through both direct and indirect actions. Direct effects included a tendency to nocturnal activity with proximity to human activity; indirect effects included increased and more clumped food supplies, and shell middens from intertidal harvesting and deforestation. Mangroves are important to local human populations as well as to hermit crabs, for a wide variety of (similar) reasons. Mangroves provide storm shelter, fisheries and fishery nursery grounds for adjacent human settlements, but they also harbour mosquito populations and their removal provides valuable building materials and fuel. Hermit crabs may be useful (indirectly) to coastal human populations by being a source of food to certain commercial species, and by quickly consuming rotting/discarded food and faeces (thereby reducing disease and pests). They can also cause minor problems to coastal human populations because they use shells of (fisheries) target mollusc species and can be more abundant than the living molluscs, thereby slowing down effective hand collection through confusion over identification. The mixture of positive and negative attributes that the three groups impart to each other in the Quirimba Archipelago, northern Mozambique, is discussed.     

红树林湿地相手蟹科动物摄食生态研究进展

相手蟹科物种是红树林湿地的主要底栖动物类群,在生态系统中起着重要的作用。我国大陆地区目前已记录的相手蟹科物种数量为12种,低于其它红树林地区(国内常用的采样方法会造成螃蟹物种数量和密度的低估),其中褶痕相手蟹(Sesarma plicata)、无齿相手蟹(S.deaani)和双齿相手蟹(S.bidens)等是常见种。红树植物叶片是相手蟹科动物的主要食物来源,相手蟹科动物通过地表摄食和洞穴贮存的形为消耗了大量的红树植物凋落叶,使这些凋落叶的有机质和营养元素得以保留在生态系统内,在凋落叶的周转和物质归还方面起到重要的作用。它们同时也摄食红树植物的繁殖体并且对不同物种的繁殖体具有摄食偏好,这可能对一些红树物种的植被更新能力和红树植被群落结构产生影响。相手蟹科动物对不同物种和不同状态的红树叶片也存在摄食偏好,通常对腐烂的叶片摄食偏好较强;螃蟹的摄食偏好与叶片的营养成份、粗纤维和单宁含量以及C/N比等性质有关;但在恶劣的野外环境下,螃蟹则会表现出随机性摄食。目前关于相手蟹科动物生态学作用的认识仍不充分,例如它们的种群大小和对凋落物的转化作用等,有待于进一步研究。     

Red-gartered Coot Fulica armillata feeding on the grapsid crab Cyrtograpsus angulatus: advantages and disadvantages of an unusual food resource

The behaviour of Red-gartered Coots feeding on an unusual food source was examined at Mar Chiquita Coastal Lagoon, Argentina. The grapsid crab Cyrtograpsus angulatus made up all observed prey items, and 61% were small. Both handling and foraging duration increased with the size of captured crabs, but foraging efficiency decreased. Crab availability affected both the dive duration of the Coots and their foraging decisions with regard to prey-size selection. Two species of gull were observed kleptoparasitizing Coots, especially when the Coot was handling medium or large crabs. Feeding by Coots on Cyrtograpsus angulatus has not been previously documented and may be a feeding innovation. Our estimations suggest that Coots were foraging optimally, since smaller crabs were more energetically profitable.     

Feeding activity of Callinectes ornatus Ordway, 1863 and Callinectes danae Smith, 1869 (Crustacea, Brachyura, Portunidae) in Ubatuba, SP, Brazil

The feeding activity along the day cycle and the time consumed for extracellular digestion were evaluated in the portunids C. ornatus and C. danae. Swimming crabs were obtained from trawling in Ubatuba bay, São Paulo, Brazil, during both the rainy and dry seasons. In each season, daily scheduled samples were taken at dawn (±6 h), noon (±12 h), dusk (±18 h) and midnight (±24 h). All individuals were dissected and the degree of stomach replenishment was recorded. In order to estimate the time elapsed for extracellular digestion, crabs were fed, and groups were dissected at 30 min intervals to check the conditions of their stomachs. In general, both species show a higher feeding activity during periods of lower light intensity, as evidenced by an increased percentage of full stomachs in dusk and midnight samples. The obtained results support higher feeding activity at night in these species and indicate short time for extracellular digestion, not exceeding 8 h. Nevertheless, full stomachs were recorded in all sampling schedules. In this case, it should be considered that elimination of certain food items such as fish bones, mollusk shells and carapace fragments of crustaceans could take more time than other items. Additionally, some crab species could require a cycle of cell replacement in the midgut gland epithelium until they can take their next meal.     

Specializations and limitations in the utilization of food resources by the carp,Cyprinus carpio: a study of oral food processing

Synopsis The wide variety of aquatic food is considered to be instrumental for the diversification in fish species. Yet their abilities and inabilities of handling food are poorly known. For these reasons the food processing and feeding repertoire of the adult carp, Cyprinus carpio, fed on a variety of food types, were analyzed by light and X-ray cinematography of the head parts and by electromyography of the head and body muscles during feeding. Nine stereotyped movement patterns (particulate intake, gulping, rinsing, spitting, selective retention of food, transport, crushing, grinding and deglutition) compose the feeding process, their sequence and frequency were adjusted to the type of food. Following quantitative morphological analysis at macroscopic, light- and electronmicroscopical level, the relations between the functioning and architecture of the feeding apparatus were established. The structure and dimensions of the mouth opening, the protrusible upper jaw, the slit-shaped pharyngeal cavity, the palatal and postlingual organ, the branchial sieve, the pharyngeal masticatory apparatus and the distribution of taste buds, mucous cells and muscle fibers along the oropharyngeal surface were the directive structural characters used for estimating the abilities in food processing. The specializations for utilizing food items and its limitations, derived from structural and functional data, are compared with diet data found in the literature in order to evaluate the relative position of the carp in competition for food in the aquatic environment. It is established that the ‘omnivorous’ carp is specialized in effective handling of several categories of aquatic food, even when these are mixed with non-food (bottom invertebrates <4% SL in diameter) since the palatal organ enables the carp to separate food from non-food. This includes very hard-skinned food items, processed with the powerful pharyngeal jaws of the fish, and to a lesser extent zooplankton (>250 μm). The carp is at the same time very limited in processing long and struggling prey (e.g. fish) as well as vegetable matter, due to the lack of oral teeth and the specialized morphology of its pharyngeal chewing apparatus. These feeding abilities agree with diet data from literature. The reported herbivorism of carp illustrates its opportunism in feeding behaviour. Specialization in feeding is discussed and the necessity to take into account the total series of post-capture feeding actions for a more complete view on trophic specialization. Food intake and the intra-oral food processing of carp are bound to the structures of its sensory, central processing and effector apparatus and to the plasticity in their functioning. These together determine its feeding efficiency in exploiting the available aquatic food resources. Next to ethological and ecological studies functional morphology is another important tool to explain the trophic interactions of fish.     

Effects of testosterone upon feeding in male mice.

Food intake and pattern of feeding were recorded in intact and castrate male mice. Intact adults are larger and eat more than pre-pubertal castrates, due to rapid growth at puberty. Castration in adulthood has little effect upon food intake, but feeding becomes concentrated into large, infrequent meals. The change in meal size appears primary and is reversed by testosterone. During rapid growth at puberty, both meal size and frequency are high. It is suggested that testosterone affects feeding in two ways. Rapid weight gain at puberty leads to high meal size and frequency, but in adulthood meal size is depressed. Meal size in adults is further depressed by stimuli from females, and it is argued in this case that testosterone affects a general mechanism for altering feeding when other activities have a high priority.     

Effects of feeding frequency on nestling begging and digestion

Nestling begging has the potential to provide parents with honest information about both short- and long-term nutritional needs, yet the importance of previous feeding experience remains largely untested in empirical studies. We examined the effect of two experimental feeding rates on nestling begging in Southern Grey Shrikes Lanius meridionalis using differences in load size to equalize the total volume of food received. There was variation in the pattern of begging behaviour between six pairs of siblings during a hand-feeding trial, although individuals maintained a similar begging intensity throughout a 9-h feeding period. Both treatment groups showed elevated begging responses during a terminal deprivation period, but nestlings fed small food items at frequent intervals demonstrated higher begging responses after a period of deprivation than did siblings fed large food items infrequently. As nestlings fed frequently with small food items had greater levels of undigested protein present in their faeces than birds fed large items infrequently, we suggest experimentally induced variation in digestive efficiency may account for the observed differences in begging behaviour. The possible role of learning, the adaptive significance of trade-offs between feeding rate and digestive efficiency, and a possible conflict of interests between parents and offspring are discussed.     

Food and feeding of three Citharinus species in Lake Kainji, Nigeria

This paper provides information on the food and feeding habits of three Citharinus species in Lake Kainji. A total number of 2357 specimens made up of 1628 Citharinus citharus , 705 C. distichodoides and 24 C, latus were examined. The majority of the fish caught by gill-netting had empty stomachs unlike those caught by cast-netting and electrofishing. The main food items were planktonic and epipelic blue-green and green algae. All the three species had similar feeding habits and also fed on the same items; their food varied with the seasons. Citharinus spp. have high gut length/standard length ratios which are adaptations to the microphagous and planktivorous feeding habits.     

Morphological divergence between three Arctic charr morphs – the significance of the deep‐water environment

Morphological divergence was evident among three sympatric morphs of Arctic charr (Salvelinus alpinus (L.)) that are ecologically diverged along the shallow‐, deep‐water resource axis in a subarctic postglacial lake (Norway). The two deep‐water (profundal) spawning morphs, a benthivore (PB‐morph) and a piscivore (PP‐morph), have evolved under identical abiotic conditions with constant low light and temperature levels in their deep‐water habitat, and were morphologically most similar. However, they differed in important head traits (e.g., eye and mouth size) related to their different diet specializations. The small‐sized PB‐morph had a paedomorphic appearance with a blunt head shape, large eyes, and a deep body shape adapted to their profundal lifestyle feeding on submerged benthos from soft, deep‐water sediments. The PP‐morph had a robust head, large mouth with numerous teeth, and an elongated body shape strongly related to their piscivorous behavior. The littoral spawning omnivore morph (LO‐morph) predominantly utilizes the shallow benthic–pelagic habitat and food resources. Compared to the deep‐water morphs, the LO‐morph had smaller head relative to body size. The LO‐morph exhibited traits typical for both shallow‐water benthic feeding (e.g., large body depths and small eyes) and planktivorous feeding in the pelagic habitat (e.g., streamlined body shape and small mouth). The development of morphological differences within the same deep‐water habitat for the PB‐ and PP‐morphs highlights the potential of biotic factors and ecological interactions to promote further divergence in the evolution of polymorphism in a tentative incipient speciation process. The diversity of deep‐water charr in this study represents a novelty in the Arctic charr polymorphism as a truly deep‐water piscivore morph has to our knowledge not been described elsewhere.     

Nutritional composition of the diet of the northern yellow-cheeked crested gibbon (<Emphasis Type="Italic">Nomascus annamensis</Emphasis>) in northeastern Cambodia

This study measured the nutritional composition of foods consumed by the northern yellow-cheeked crested gibbon (Nomascus annamensis) in northeastern Cambodia. One group of N. annamensis was studied, and focal animal sampling was used to observe their feeding behavior. The study was conducted for 4 months (January–April 2015) in the dry season and 69 foods were collected for nutritional analyses. N. annamensis fed on 37 plant species, but only seven species made up more than 80% of feeding time. N. annamensis spent the majority of their time feeding on fruit (60.36%), followed by young leaves (22.60%), flowers (13.74%), and mature leaves (3.30%). Fruit had the highest concentrations of total non-structural carbohydrates, while young leaves had the highest concentration of crude protein compared to other food items. All food items had similar concentrations of lipids, except young leaves, which had lower levels than fruit and flowers. All plant parts consumed by N. annamensis had similar amounts of neutral detergent fiber, acid detergent fiber, and acid detergent lignin. Thirty-two percent of all food items contained condensed tannins, and they had a similar level of it. All foods also had similar energy contents. This study adds to our knowledge of the feeding ecology of N. annamensis by providing baseline data on the make-up of the foods they eat and may contribute to captive feeding programs, ultimately assisting the conservation of this species.     

A tadpole-induced polyphenism in the salamander Hynobius retardatus

Abstract.— Larvae of the salamander Hynobius retardatus have two distinct morphs: normal and broad-headed, cannibal morphs. We performed three experiments to differentiate among the following hypotheses: The broad-headed morph is induced to allow: (1) feeding on nutritious conspecifics; (2) exclusion of strong competitors for food or space; or (3) feeding on large, tough prey when smaller prey items are unavailable. When newly hatched larvae were reared with a heterospecific, Rana pirica (an anuran amphibian) tadpoles, the broad-headed morph was induced more frequently compared with those reared with conspecifics. The phenotype expressed depended on the size of the tadpoles: The broad-headed morph occurred more frequently with small and the normal morph with large tadpoles. Metamorphosis occurred sooner in larvae fed conspecifics compared with those fed heterospecific tadpoles, and the mean growth rate of larvae fed conspecifics was significantly faster than that of those fed tadpoles, suggesting that the heterospecific tadpoles were less nutritive than the conspecifics. These results do not support the hypotheses that the broad-headed morph evolved for consuming conspecifics because of their better balance of nutrients or for excluding strong competitors for food or space. We tentatively conclude that the morph evolved to eat large, tough prey, including both conspecifics and heterospecific tadpoles. Because H. retardatus usually spawns very early in the spring in small ponds partially covered with ice and snow, newly hatched larvae may starve from the lack of proper food owing to extremely low water temperatures. Thus, the broad-headed morph of H. retardatus may represent a cold-habitat adaptation to overcome the severe circumstance when the only food items available are relatively large conspecifics or heterospecific tadpoles.     

Food segregation in three species of portunid crabs

Gastric mills of 2,619 specimens of portunid crabs(Portunus pelagicus, P. sanguinolentus andCharybdis feriatus) in Tolo Harbour, Hong Kongwere examined. Of these, 1,498 containedrecognizable food items. A total of 34 food items wereidentified, 10 of which (29.4%) were common to thethree crab species. The principal food items werebivalves, decapods, fish and algae. However, thepercent frequency of occurrence of these food itemsdiffered significantly (ANOVA, p < 0.001) among thecrabs. In addition to these principalfood items, differences were noted in other dietarycategories: P. pelagicusand P. sanguinolentus fed on the brittle starOphiura kinbergi, whereas P.sanguinolentus and C. feriatus fed on detritusand gastropods. The diet of P. sanguinolentuswas found to be more similar to C. feriatus thanto P. pelagicus. Exploitation of different fooditems and selective feeding on common prey may reduceinter-specific competition for natural food resourcesand allow the three portunid crabs to co-exist in thesame habitat. This revised version was published online in July 2006 with corrections to the Cover Date.     

Ontogenetic changes in diet and related morphological adaptations in Ocypode gaudichaudii

There are conflicting reports as to whether Ocypode gaudichaudii individuals switch from carnivory as juveniles to deposit‐feeding primarily on diatoms as adults, or whether they expand diet range and become omnivorous with maturity. At the onset of deposit‐feeding, crabs develop specialized claws with truncated ends that they use to shovel sediment during foraging. Eighty‐eight crabs were collected from Culebra Island (Republic of Panama) to study how the diet of this crab shifts with changes in claw shape, mouthpart proportions, and setation, as well as gastric mill width. Forty‐four crabs had identifiable material in their foreguts: 30 had animal material, 12 had diatoms, and two had a mix of both. There were no differences between the gastric mill, mandibles, and the proximal endites of the first maxillipeds of predators and deposit‐feeders, but extra rows of plumose setae were present on the second maxilliped of deposit‐feeding crabs with carapace length (CL) >10.6 mm. All individuals with CL <12.3 mm and non‐truncated claws ate animals, but those with larger CL and truncated claws had animal, diatom, or mixed diets; hence, claw truncation does not restrict the crab's diet to diatoms but, instead, broadens the diet to include both animals and diatoms. Perhaps this is a strategy to balance the economics of foraging on animals and diatoms on medium to low‐energy beaches that lack the larger invertebrates that adults of other species of ghost crabs eat. More generally, our study shows that specialized feeding structures need not imply a narrow or specialist diet.     

Selective Prey Delivery to Incubating Females by Dominant Males, but not Helpers, in the Cooperatively Breeding Green Woodhoopoe

In cooperatively breeding bird species, one of the most conspicuous helping behaviours is the provisioning of food. Many studies have considered the feeding of nestlings, but far fewer have examined feeding of incubating females, and none have looked at the types of prey delivered at this stage. Here I show that green woodhoopoe ( Phoeniculus purpureus ) group members selectively feed incubating females with certain prey items: the diet delivered to incubating females contains a higher proportion of caterpillars, centipedes and cockroaches than that eaten by the provisioning adults themselves. The prey items selectively delivered are the largest in the diet and so might be provided in an effort to enhance the breeding female's condition, thus minimizing the time that she spends off the nest and so increasing hatching success. Intriguingly, it is only breeding males that show this adjustment in provisioning diet; helpers of both sexes simply provide the incubating female with the same proportions of different prey items that they eat when self-feeding. My results therefore offer the first evidence that members of cooperative groups may not all follow the same provisioning rules, and they also emphasize the need for studies to consider the incubation stage in just as much detail as the nestling phase if we are to understand fully the complexities of cooperative societies.     

Nutritional Ecology of <Emphasis Type="Italic">Ateles chamek</Emphasis> in lowland Bolivia: How Macronutrient Balancing Influences Food Choices

All free-living animals must make choices regarding which foods to eat, with the choices influencing their health and fitness. An important goal in nutritional ecology is therefore to understand what governs animals’ diet selection. Despite large variation in the availability of different food items, Peruvian spider monkeys (Ateles chamek) maintain a relatively stable daily protein intake, but allow total energy intake to vary as a function of the composition of available food items. This is referred to as protein-dominated macronutrient balancing. Here we assess the influence of this nutritional strategy on daily and seasonal nutritional intakes, estimate the nutritional value of different foods, and interpret unusual food choices. We conducted continuous all-day observations of focal spider monkeys inhabiting a semideciduous forest in Bolivia. We recorded feeding events, collected foods, and analyzed their nutrient content. By using the Geometric Framework for nutrition, we show that individuals reached their daily end-point in nutrient space —balance between protein and nonprotein energy intake— by consuming nutritionally balanced foods or by alternating between nutritionally complementary foods. The macronutritionally balanced figs of Ficus boliviana were their primary staple food and therefore dominated their overall nutritional intake. Our results also demonstrate that spider monkeys consumed a diverse array of ripe fruits to overcome periods of fig scarcity rather than vice versa; they could obtain sufficient protein on a diet of pure fruit; and unripe figs constituted a nutritionally rewarding and reliable food resource. We hope that the approaches taken and the conclusions reached in this study will catalyze further inquiries into the nutritional ecology of frugivorous primates.     

Morphological specializations in heterocongrinae (Anguilliformes: Congridae) related to burrowing and feeding

The remarkable lifestyle of heterocongrines has drawn the attention of many authors in the past, though no or little attention has been paid to the morphology of the tail and the head of these species. In order to examine the true nature of possible morphological specializations of the head and tail and their relation to their tail-first burrowing habit and/or feeding mode, a detailed myological and osteological study of Heteroconger hassi and Heteroconger longissimus was performed. The osteological similarities of the cranial skeleton between H. hassi and H. longissimus are striking. Most of the cranial muscles show no variation in presence, insertion or origin between these two species except for the adductor mandibulae complex, the adductor hyomandibulae and the intermandibularis. The adductor mandibulae complex is small, compared to that of other anguilliform species, and is probably related to their suction-dominated feeding mode and a diet, comprising mainly small, soft prey items. Heterocongrinae have undergone several morphological specializations in the tail for their tail-first burrowing lifestyle. The skeleton and musculature of the tail of H. hassi and H. longissimus are similar. In both species the caudal skeleton is highly reduced and fortified, forming a firm, pointed burrowing tool. Intrinsic caudal musculature is reduced and some muscles (interradials, supracarinalis) are even absent.     

Seasonal changes in food resource distribution and feeding sites selected by Japanese macaques on Koshima Islet,Japan

Feeding sites of Japanese macaques (Macaca fuscata) change according to seasonal fluctuations in food resource distribution. To examine what characteristics of food items affect feeding site selection, I describe herein the seasonal changes in food items, feeding sites, and food resource distributions of this species. Feeding behavior of monkeys and their food resource distributions were investigated on Koshima Islet, southern Japan, for four study periods (i.e., seasons) in 2002. Monkeys showed large variations in their diet between seasons. To weigh the relative influence of the distribution and abundance of food items on feeding site selection in each season, multiple regression analyses were performed by 100 m × 100 m grid. In the analyses, feeding time was a dependent variable and the abundance of staple food items, for which feeding time was over 5% in each season, in each grid square was an independent variable. There was no correlation between the resource distribution of most food items and the distribution of feeding time by monkeys in each season. Monkeys spent more feeding time where multiple staple food items were available. Food items that affected feeding site selection by monkeys had the following three characteristics: (1) clumped distribution, (2) seasonal availability, and (3) fruit. This suggests that monkeys are likely to select feeding sites to consume food items whose availability is limited temporally and spatially, which may enable them to simultaneously use other widely distributed, abundant food items efficiently.     

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.