Empty Snail Shells Research Articles

How can we measure resource quality when resources differ in many ways? Deconstructing shelter quality in a social fish.

Resource quality is an important concept in ecology and evolution that attempts to capture the fitness benefits a resource affords to an organism. Yet "quality" is a multivariate concept, potentially affected by many variables pertaining to the resource, its surroundings, and the resource chooser. Researchers often use a small number of proxy variables to simplify their estimation of resource quality, but without vetting their proxies against a wider set of potential quality estimators this approach risks overlooking potentially important characteristics that can explain patterns of resource use in their study systems. Here we used Neolamprologus multifasciatus, a group-living cichlid fish that utilizes empty snail shells as shelter resources, to examine how shells were used by, and partitioned among, group members in relation to a range of attributes, including shell size, intactness, texture, spatial position, and usage by heterospecifics. This approach generated a comprehensive picture of what characteristics contribute to the attractiveness and quality of each shell resource, confirming the importance of two previously proposed shell characteristics, size and intactness, but highlighting the influences of other unexplored variables, including shell spatial position and usage by heterospecifics. We also present a generally applicable "resource attractiveness index" as a means to estimate resource quality based on resource choice data. This index incorporates information from any number of resource characteristics and is of particular use when researchers wish to quantify resource value, but many characteristics jointly contribute to the value and attractiveness of the resource.

Genomic basis of Y-linked dwarfism in cichlids pursuing alternative reproductive tactics.

Sexually antagonistic selection, which favours different optima in males and females, is predicted to play an important role in the evolution of sex chromosomes. Body size is a sexually antagonistic trait in the shell-brooding cichlid fish Lamprologous callipterus, as "bourgeois" males must be large enough to carry empty snail shells to build nests whereas females must be small enough to fit into shells for breeding. In this species, there is also a second male morph: smaller "dwarf" males employ an alternative reproductive strategy by wriggling past spawning females into shells to fertilize eggs. L.callipterus male morphology is passed strictly from father to son, suggesting Y-linkage. However, sex chromosomes had not been previously identified in this species, and the genomic basis of size dimorphism was unknown. Here we used whole-genome sequencing to identify a 2.4-Mb sex-linked region on scaffold_23 with reduced coverage and single nucleotide polymorphism density in both male morphs compared to females. Within this sex region, distinct Y-haplotypes delineate the two male morphs, and candidate genes for body size (GHRHR, a known dwarfism gene) and sex determination (ADCYAP1R1) are in high linkage disequilibrium. Because differences in body size between females and males are under strong selection in L.callipterus, we hypothesize that sexual antagonism over body size initiated early events in sex chromosome evolution, followed by Y divergence to give rise to bourgeois and dwarf male reproductive strategies. Our results are consistent with the hypothesis that sexually antagonistic traits should be linked to young sex chromosomes.

The native snail shell–nesting beeOsmia conjuncta(Hymenoptera: Megachilidae) exploits a local abundance of exoticCepaeasnails (Stylommatophora: Helicidae), choosing empty shells by size, colour, and microhabitat

AbstractFor solitary bees, nesting substrate availability may limit population growth. Here, we investigate the nesting of a locally abundant beeOsmia conjunctaCresson (Hymenoptera: Megachilidae) in empty shells of exotic snails,Cepaeaspp. Held (Stylommatophora: Helicidae), in Hamilton, Ontario, Canada. A literature review determined thatO. conjunctawas sparsely distributed throughout its range, with a low relative abundance except for the Niagara Region of Ontario, Canada, three sites in Maryland, United States of America, and the focal study area in Hamilton, Ontario. The Hamilton field survey of five grassland bee communities foundO. conjunctato be the most abundant bee species. The average relative abundance was 18.7%, peaking at 87% on one spring day. A survey of 1088 empty snail shells was done at the site with the highestO. conjunctaabundance. It revealed an average of 10.87 empty shells per square metre with cocoons in 9.8% of the shells, averaging 1.6 cocoons per nest. The relation between shell traits and cocoon presence indicated thatO. conjunctapreferred nesting in shells with intermediate size and colour and avoided shells near shrubs. The presence ofCepaeasnails increasedO. conjunctapopulations with potential consequences to the local bee assemblage.

Palaearctic Osmia bees of the subgenera Allosmia and Neosmia (Megachilidae, Osmiini): biology, taxonomy and key to species

Osmia (Allosmia) and O. (Neosmia) are two closely related Palaearctic osmiine bee subgenera (Megachilidae, Megachilinae, Osmiini) comprising eleven and ten species, respectively. Analysis of female pollen loads revealed that the representatives of O. (Allosmia) and O. (Neosmia) are pollen generalists collecting pollen on up to 13 and 17 plant families, respectively. The species of both subgenera exclusively nest in empty snail shells and use chewed leaves as main nest building material, often in combination with fragments of mollusc shells and densely packed small particles, such as small stones, earth crumbs or plant fibers, which are used to barricade the nests. The taxonomic revision of the two subgenera revealed the existence of two undescribed species, O. (Allosmia) gemina spec. nov. from the southern Levant and O. (Neosmia) nigrocalcaribus spec. nov. from the Maghreb. Osmia fossoria Pérez, 1890 syn. nov. and O. decorata Morawitz, 1886 syn. nov. are newly synonymized with O. (Allosmia) lhotelleriei Pérez, 1887 and O. (Allosmia) melanura Morawitz, 1871, respectively, whereas the former synonymization of O. (Neosmia) rosea Friese, 1920 with O. scutispina Gribodo, 1894 is rejected. Osmia (Neosmia) purpurata Ducke, 1899 is regarded as a nomen dubium. Osmia (Allosmia) soror Pérez, 1896 and O. (Neosmia) secunda Peters, 1977 are treated as species rather than as subspecies of O. rufohirta Latreille, 1811 and O. tingitana Benoist, 1969, respectively, due to clear morphological gaps and partly sympatric occurrence. Identification keys for all species of the two subgenera are given.

The Evolution of Social Behaviour

The Evolution of Social Behaviour

Social and spatial conflict drive resident aggression toward outsiders in a group-living fish.

Group-living animals often experience within-group competition for resources like shelter and space, as well as for social status. Because of this conflict, residents may aggressively resist joining attempts by new members. Here, we asked whether different forms of competition mediate this response, specifically competition over 1) shelter, 2) spatial position within groups, and 3) social or sexual roles. We performed experiments on wild groups of Neolamprologus multifasciatus cichlids in Lake Tanganyika, either increasing or decreasing the number of shelters (empty snail shells) within their territories. We predicted that increases in resource abundance would reduce conflict and lower the aggression of residents toward presented conspecifics, while decreases in resources would increase aggression. We explored the effects of social conflict and spatial arrangement by introducing same or opposite sex conspecifics, at greater or lesser distances from resident subterritories. We found that changing the abundance of shells had no detectable effect on the responses of residents to presented conspecifics. Rather, aggression was strongly sex-dependent, with male residents almost exclusively aggressing presented males, and female residents almost exclusively aggressing presented females. For females, this aggression was influenced by the spatial distances between the presented conspecific and the resident female subterritory, with aggression scaling with proximity. In contrast, presentation distance did not influence resident males, which were aggressive to all presented males regardless of location. Overall, our results show that group residents respond to presented conspecifics differently depending on the type of competitive threat these potential joiners pose.

Palaearctic Osmia bees of the subgenera Hemiosmia, Tergosmia and Erythrosmia (Megachilidae, Osmiini): biology, taxonomy and key to species

Osmia (Hemiosmia), O. (Tergosmia) and O. (Erythrosmia) are three related Palaearctic osmiine bee subgenera (Megachilidae, Megachilinae, Osmiini) comprising eight, seven and four species, respectively. Analysis of female pollen loads and field observations revealed that the species of both O. (Hemiosmia) and O. (Tergosmia) have a distinct or even exclusive preference for Fabaceae as pollen hosts. Species of O. (Erythrosmia) also collect pollen on Fabaceae, but additionally frequently visit nototribic flowers of Lamiaceae and Antirrhineae (Plantaginaceae), which they exploit by rubbing a specialized facial pilosity over the raised anthers. The representatives of O. (Hemiosmia) and the basal members of O. (Tergosmia) build their brood cells in self-excavated burrows in the ground, whereas the more derived O. (Tergosmia) species as well as the O. (Erythrosmia) species place them in preexisting cavities above-ground, which are empty snail shells hidden under prominences of rocks or under stones in the latter subgenus. In O. (Hemiosmia) and O. (Tergosmia), the brood cells are entirely constructed from foreign material and either consist of chewed leaves ("leaf pulp") in the former subgenus or are distinctly three-layered with a thin layer of mud sandwiched between two layers of large pieces of petals in the latter subgenus. In O. (Erythrosmia), the brood cells within the shell whorls are delimited by one-layered partitions of leaf pulp and the nests are closed by a barricade of densely packed small particles followed by a double wall of leaf pulp at the shell opening. The taxonomic revision of the three subgenera revealed the existence of an undescribed species, O. (Hemiosmia) spinicoxa spec. nov., which occurs in southwestern Morocco. The subgenus O. (Ozbekosmia) Zanden, 1994 syn. nov. is newly put into synonymy with O. (Tergosmia) Warncke, 1988. Osmia (Tergosmia) rhodoensis arquata Warncke, 1988 syn. nov. and O. (Tergosmia) tergestensis ononidis Ferton, 1897 syn. nov. are no longer accepted as valid subspecies of O. (Tergosmia) rhodoensis (Zanden, 1983) and O. (Tergosmia) tergestensis Ducke, 1897, respectively. Identification keys for all species of the three subgenera are given.

Near-natural habitats near almond orchards with presence of empty gastropod shells are important for solitary shell-nesting bees and wasps

Solitary bees serve as important native pollinators of wild plants and crops. Several species of bees, which form strong populations in open habitats of European landscapes, nest in empty snail shells. In the present study, we focused on shell-nesting bees in near-natural habitats neighbouring almond orchards near the town Lleida in north-east Spain. We performed a series of manipulative experiments, which aimed to elucidate the importance of empty shells of four common large- and medium-sized gastropod species for nesting bees in near-natural and ruderal habitats surrounding the almond orchards. We also matched these data with findings of bees in naturally occurring shells at each examined site. All study sites consisted of habitats with naturally present shells and evidence of the natural presence of the study species of bees. In total, we recorded 15 nesting bee and wasp species and eight species of hymenopteran parasites in collected shells, one of which, the spider wasp Priocnemis propinqua, was recorded nesting in empty gastropod shells for the first time. In total, the deployed shells hosted eight nesting species and two species of parasitic Hymenoptera. The prevalence of nests in experimentally deployed shells was low, with only a single nest present near irrigated orchards. The abundance of nests was also low at sites where the naturally present shells were highly abundant. Combined, available evidence suggests the importance of near-natural habitats for the presence of shell-nesting bees and wasps in South-European landscapes. The abundance of shell nesting bees and wasps does not increase proportionally with the increase in empty snail shell abundance and other factors should be considered limiting at such sites.

Nest structure, pollen utilization and parasites associated with two west-Mediterranean bees (Hymenoptera, Apiformes, Megachilidae) nesting in empty snail shells

Around thirty species of European solitary bee species in the family Megachilidae nest in empty gastropod shells. We surveyed this group of bees in semi-natural sites adjacent to almond orchards near Lleida (north-eastern Spain) and collected 35Hoplitis fertoniand 58Osmia ferrugineanests in shells of six snail species. We describe the nest structure and report the identity of pollens collected by the two bee species. Both species adjust the number of brood cells to the size of the shell and occasionally build intercalary (empty) cells.H. fertoniuses clay andO. ferrugineachewed plant leaves for building cell partitions and nest plugs. Most nests of both species were built inSphincterochila candidissimashells. Analysis of the pollen of selected nests confirmed thatH. fertoniis oligolectic on Boraginaceae (in our study all pollen was fromLithodora fruticosa) andO. ferrugineais a polylectic species (collecting mostly pollen from Cistaceae, Fabaceae, and Lamiaceae in our study area). Nests ofH. fertoniwere parasitized by five species, the golden waspChrysura hybrida, the cuckoo beeDioxys moesta, the velvet antsStenomutilla collarisandStenomutilla hotentotta, and the bee-flyAnthrax aethiops; nests ofO. ferrugineawere parasitized by the sapygid waspSapyga quinquepunctataandA. aethiops. Except forC. hybridathese are newly recorded host-parasite associations. Our results confirm previous information and bring new findings on the ecology of both species.

Effects of grazing intensity, habitat area and connectivity on snail-shell nesting bees

The effects of habitat loss and fragmentation on wild bee populations are still not fully understood. Availability of resources, parasitism and habitat connectivity might influence population sizes of bees. Whereas floral resources for wild bees have been considered in research and conservation, nesting resources have been largely neglected as these are challenging to investigate. Snail shells are the exclusive nesting cavities for several Osmia species and provide a good - but so far unused - tool to study factors driving bee population dynamics. We investigated the effects of habitat area, connectivity and management of semi-natural grasslands on populations of snail-shell nesting bees. On 23 calcareous grasslands, we monitored snail shell colonization by providing empty snail shells as nesting resources, and recorded flower-visitor interactions.Five snail-shell nesting Osmia species were found on the grasslands, which made almost a quarter of recorded bee flower visits. Three species colonized the offered snail shells with high variation between study sites, Osmia species, and snail shell size. Habitat area had a positive effect on the population size of the habitat specialist Osmia aurulenta, whereas the more generalist species Osmia bicolor was positively influenced by habitat connectivity. Destruction rates of snail shells increased with sheep grazing intensity, leading to an estimated loss of more than a third of all bee nests. We conclude that large and connected habitats benefit bee populations in fragmented landscapes, while conservation management regimes should take into account potential negative effects of grazing on specific nesting resources of specialized bee species.

Experimental confirmation of empty snail shells as limiting resources for specialized bees and wasps

Observational studies have suggested that the presence of snail shells correlates with the presence of specialized adopters, particularly bees and wasps (Hymenoptera: Aculeata). However, should the empty shells be considered limiting resources once they are present in the respective habitat even if observational studies suggested low ratio of occupied relative to total shells? We performed a manipulative experiment, which consisted of the addition of marked snail shells of six terrestrial species that dominate the open and semi-open central European habitats to 21 sites with naturally present shells. We deployed empty shells for the spring and summer 2017, allowed their inhabitants to undergo a diapause during the follow-up winter period and complete metamorphosis. The specialized bee and wasp species abundantly occupied the provided shells and the occupancy rates were several times higher in experimentally provided shells compared to the naturally present shells. These differences in occupancy rates were higher at anthropogenic compared to natural sites and at sites with more limited availability of naturally present shells. In contrast, the few sites at which the examined resource was superabundant, both the naturally present and the experimentally provided shells were occupied only to a limited extent. The species composition of assemblages that occupied the experimentally provided and naturally present shells were similar, and the differences in species composition between the natural and anthropogenic sites resembled those exhibited in the naturally present shells. In conclusion, we experimentally confirmed that empty snail shells serve as limiting resources for specialized bees and wasps even at sites where the naturally present shells are perceived as abundant.

Morphology and Habitats of the Hermit-Crab-Associated Calyptraeid Gastropod Ergaea walshi.

Ergaea walshi, a gastropod with a markedly flat shell, often lives inside empty snail shells occupied by hermit crabs. We investigated its lifestyle, shell growth pattern, and habitat preference for host hermit crabs and host snail shells. Four hundred sixteen snail shells, including 363 shells with hermit crabs and 53 empty shells, were collected from intertidal zones of sandy and muddy flats around Kii Peninsula, Japan. The specimens comprised seven hermit crab species occupying 24 shell species; E. walshi was harbored in 13.2% of snail shells with hermit crabs and 17.0% of those without hermit crabs. Although no preference was detected for particular species of hermit crab or snail shell, E. walshi preferred to live inside of snail shells with wider apertures used by comparatively bigger hermit crabs. This suggests that the occurrence of E. walshi was influenced by host size rather than host species. When looking at growth patterns, we found that the attached shell portion of E. walshi continued to be enlarged horizontally, while growth in shell height slowed at approximately 5.0 mm. The conspicuously flattened shell of E. walshi is considered as a growth pattern for adapting to the narrow space within the snail shell occupied by hermit crabs. Consistent with this idea, our comparison of shell growth patterns in 23 calyptraeid species showed that shell of E. walshi is the flattest in this family.

The presence of high numbers of empty shells in anthropogenic habitats is insufficient to attract shell adopters among the insects

Abstract Terrestrial snail shells can be considered partial resources for insects, and as such, the expansion of numerous snail species to anthropogenic habitats makes them increasingly available. Here, we address for the first time a complete profile of insects that use empty terrestrial snail shells during the winter period in Central Europe. The specialisation for shells made by certain snail species was uncommon; however, a number of species showed significant preferences for certain shell types. We found that the presence of empty snail shells in anthropogenic habitats drives the presence of many empty snail shell adopters in these habitats. Nevertheless, the increased availability of snail shells proved to be insufficient for a transition of all the species of snail shell adopters from natural to anthropogenic habitats. The avoidance of anthropogenic habitats among snail shell adopters was particularly distinct in species that use them only as a winter retreat but which require additional feeding and breeding resources, such as the true bugs. The availability of snail shells is thus a pre‐requisite of the presence of specialised snail shell adopters but is not necessarily sufficient to establish their presence in the respective habitat.

Biology of Palaearctic Wainia bees of the subgenus Caposmia including a short review on snail shell nesting in osmiine bees (Hymenoptera, Megachilidae)

Wainia, a species-poor genus of osmiine bees, contains two Palaearctic species, whose biologies are unknown. In the present publication, we describe the nesting site and nest architecture ofW.sexsignataand analyse the pollen host spectra ofW.sexsignataandW.eremoplanaby microscopical analysis of larval faeces and female scopal contents.W.sexsignatanests in empty snail shells. It uses large petal pieces of yellow flowered Asteroideae to separate the linearly arranged brood cells and to construct a series of partitions between the outermost cell and the final nest closure, which consists of masticated green leaves.W.sexsignatais recorded as a new host of the meloid beetleSitarobrachysthoracica. Both PalaearcticWainiaspecies have a strong preference for Asteraceae as pollen hosts, but occasionally also collect pollen on other plant taxa. A literature survey revealed that nesting in empty snail shells is widespread among osmiine bees: obligate snail shell nesting is assumed to have independently evolved at least twelve times in their evolutionary history and to occur in at least 56 species belonging to 4 genera and 14 subgenera.

Colour variation of a shell-brooding cichlid fish from Lake Tanganyika

The dwarf morph of Telmatochromis temporalis uses empty snail shells as shelters and spawning sites. This morph varies in body colour from pale to dark within populations. Pale individuals are likely to be cryptic on sandy-brown backgrounds through colour matching, but adaptive significance of dark individuals is unknown. The present study proposes two hypotheses for the occurrence of dark individuals: crypsis in darkness inside shells (darkness matching) and crypsis in irregular patterns consisting of shell shadows on sandy-brown backgrounds (pattern matching). The former hypothesis is less likely to be the main factor evolving dark bodies, because body darkness was not correlated with the amount of time that the fish remained within shells. Body darkness was correlated with shell density in their habitats, supporting the latter hypothesis because dark individuals may be able to mingle with shell shadows effectively at sites where shells were abundant. This study also found that body darkness was environmentally induced; therefore, this fish may shift between these anti-predator tactics (crypsis of pale individuals on sandy-brown backgrounds through colour matching and crypsis of dark individuals in irregular shadow patterns through pattern matching) by changing body darkness in response to shell density.

Palaearctic Osmia bees of the subgenus Hoplosmia (Megachilidae, Osmiini): biology, taxonomy and key to species.

Hoplosmia, a subgenus of the osmiine bee genus Osmia (Megachilidae), comprises 21 species restricted to the Palaearctic region. Analysis of female pollen loads and field observations indicate that probably all O. (Hoplosmia) species are specialized on Asteraceae except for one pollen generalist species, which exhibits a preference for the pollen of Cistaceae. Among the Asteraceae specialists, differences exist with respect to the three main Asteraceae subfamilies exploited for pollen, with some species exclusively visiting Carduoideae, others exploiting only Asteroideae and Cichorioideae and again others collecting pollen on Asteroideae, Carduoideae and Cichorioideae. All O. (Hoplosmia) species build their brood cells within preexisting cavities: several species exclusively nest in empty snail shells, few species use small cavities in rock and stones and the remaining species colonize linear cavities in dead wood and plant stems or nest in abandoned burrows of other bees and wasps. Chewed leaves serve as material to construct brood cell partitions and nest plug except for two species, which use mud as nest building material. The taxonomic revision of O. (Hoplosmia) revealed the existence of an undescribed species, O. centaureae spec. nov., which occurs in a small area that ranges from the Dead Sea over the Jordan Valley to northernmost Israel. Due to clear morphological gaps and widely disjunct distribution with the nominotypical subspecies, O. pinguis carbo (Zanden 1974) is elevated to species rank. Based on morphology and biology, three species groups are recognized within Hoplosmia. Identification keys for all O. (Hoplosmia) species are given including the hitherto unknown male or female sex of three species.

Estimating factors influencing the detection probability of semiaquatic freshwater snails using quadrat survey methods

Developing effective monitoring methods for elusive, rare, or patchily distributed species requires extra considerations, such as imperfect detection. Although detection is frequently modeled, the opportunity to assess it empirically is rare, particularly for imperiled species. We used Pecos assiminea (Assiminea pecos), an endangered semiaquatic snail, as a case study to test detection and accuracy issues surrounding quadrat searches. Quadrats (9 × 20 cm; n = 12) were placed in suitable Pecos assiminea habitat and randomly assigned a treatment, defined as the number of empty snail shells (0, 3, 6, or 9). Ten observers rotated through each quadrat, conducting 5-min visual searches for shells. The probability of detecting a shell when present was 67.4 ± 3.0%, but it decreased with the increasing litter depth and fewer number of shells present. The mean (± SE) observer accuracy was 25.5 ± 4.3%. Accuracy was positively correlated to the number of shells in the quadrat and negatively correlated to the number of times a quadrat was searched. The results indicate quadrat surveys likely underrepresent true abundance, but accurately determine the presence or absence. Understanding detection and accuracy of elusive, rare, or imperiled species improves density estimates and aids in monitoring and conservation efforts.

Overwintering of Spiders in Terrestrial Molluscs Shells in the Wider Area of Vranov nad Topľou (Slovakia)

Overwintering of spiders in empty snail shells is a little studied topic and in Slovakia similar research has not been carried out yet. Collecting shells was carried out in the eastern part of Slovakia, in winter at the turn of 2012/2013 and 2013/2014. The study was conducted both at the steppe like and post-industrial habitats. A total of 1085 shells were collected with a spider occupation rate 10.69 %. All acquired 116 spiders were classified in ten families. The most abundant spider family was the Salticidae, with a dominant species Pellenes tripunctatus (Walckenaer, 1802). Significant records include Cheiracanthium montanum (L. Koch, 1877) which is considered as vulnerable and Sitticus penicillatus (Simon, 1875) with a lower risk thread according to Red List of Slovakian spiders.

Sperm-limited males save ejaculates for future matings when competing with superior rivals

Adjusting ejaculates to sperm competition can lead to sperm limitation. Particularly in polygynous species, males may face a trade-off between investing sperm in current or future mating opportunities. The optimal sperm allocation decision should depend on the relative intensity of sperm competition experienced in a mating sequence. Here we asked how males respond to this trade-off in polygynous fish with alternative male mating tactics, intense sperm competition and sperm limitation. Large bourgeois males of the shell-brooding cichlid Lamprologus callipterus build nests consisting of empty snail shells, in which females spawn and raise offspring. During spawning, nest males release ejaculates into the shell opening. Genetically distinct, parasitic dwarf males enter shells during spawning to fertilize the eggs from inside the shell. These dwarf males were previously shown to be superior sperm competitors to nest males. Here we showed that when spawning with several females simultaneously, nest males reduced the spawning duration for each clutch and the number of ejaculations per female tended to decrease, reflecting sperm limitation. Experimental exposure of nest males to sperm competition with dwarf males reduced the number and duration of ejaculations by roughly half. Hence, when exposed to competition with a superior rival, nest males did not increase their sperm expenditure as predicted by sperm competition risk models, but in fact saved sperm for future mating opportunities as predicted by sperm competition intensity theory. This seems to be adaptive because of the considerable sperm demands in this species, which is partly due to their high degree of polygyny.

Sex and breeding behaviour of the Sicilian snail-shell bee (Rhodanthidium siculum Spinola, 1838; Apoidea–Megachilidae): preliminary results

In spring 2014 and 2016, we studied Rhodanthidium siculum in dunes south of Syracuse (Sicily). The females search for one of the frequently empty snail shells (mostly of Theba pisana), checking their size. In a suitable snail shell, a mixture of sand and saliva is deposited in the navel. A severe struggle among several males gets started. At every opportunity, the males try to copulate with the females. Large males occupy the snail shell that is used by the female to bring in pollen and nectar (preferably from Galactites tomentosus, Centaurea sphaerocephala, Glebionis coronaria and Lotus creticus). The procedure of harvesting lasts about 2–3 h, sometimes several hours. During this period, the female is visited for copulation every 2 min (initially even more frequently), preferentially by one large male, but small males occasionally mate as well. After laying one or two eggs, the female closes the snail shell with pieces of seashells or snail shells, aggregated with sand and saliva. The female transports the closed snail shell to a safe site. Depending on weather conditions, this may take several days. The maximum distance of movement observed was about 10 m. Finally, the completed snail shell will be buried, most often beneath a plant. The burial follows a certain pattern of movement. Females ensure higher pollination efficiency than the males due to their higher flower constancy.