Sexual dimorphism in shell size of the land snail Leptopoma perlucidum (Caenogastropoda: Cyclophoridae).

Sexual dimorphism in the shell size and shape of land snails has been less explored compared to that of other marine and freshwater snail taxa. This study examined the differences in shell size and shape across both sexes of Leptopoma perlucidum land snails. We collected 84 land snails of both sexes from two isolated populations on two islands off Borneo. A total of five shell size variables were measured: (1) shell height, (2) shell width, (3) shell spire height, (4) aperture height, and (5) aperture width. We performed frequentist and Bayesian t-tests to determine if there was a significant difference between the two sexes of L. perlucidum on each of the five shell measurements. Additionally, the shell shape was quantified based on nine landmark points using the geometric morphometric approach. We used generalised Procrustes and principal component analyses to test the effects of sex and location on shell shape. The results showed that female shells were larger than male shells across all five measurements (all with p-values < 0.05), but particularly in regards to shell height and shell width. Future taxonomic studies looking to resolve the Leptopoma species' status should consider the variability of shell size caused by sexual dimorphism.

Exploring the predation of large land snails using preyed shell remains from rock anvil sites in a tropical limestone rainforest in Malaysia.

The study of prey-predator interactions between land snails and birds offers important insights into evolutionary and ecological relationships. Here, we report a case study of rock anvils presumably used by the birds Myophonuscaeruleus and Enicurusruficapillus in a cave cavity of a limestone hill in Malaysia. We did not detect any other species in the plots and, therefore, based on our short study duration, we cannot rule out the possibility that other species, such as mammals, preyed on the snails. The predated shell remains of four land snails namely, Hemiplecta sp., Cyclophorusperdixperdix, Amphidromusatricallosusperakensis and Cyclophorussemisulcatus, were found around rock anvils in the nine plots. Finally, we discussed the potential and the limitations of using shell remains of preyed land snails for behavioural, ecological and evolutionary studies between land snails and their predators.

Distribution and abundance of the land snail Pollicaria elephas (Gastropoda: Pupinidae) in limestone habitats in Perak, Malaysia.

This study aimed to reveal the habitat variables that determine the distribution and abundance of the land snail Pollicaria elephas in limestone habitats in Perak, Malaysia. Seventeen plots were selected on a limestone hill to determine the effect of environmental variables on the abundance of this land snail. The environmental variables we considered included habitat (canopy cover and leaf litter thickness), topography (elevation, aspect, ruggedness, and slope), microclimate (soil temperature, air temperature, and humidity), and vegetation (abundance of respective vascular plant species). The correlation analyses suggested that the snails' abundance was positively correlated with the abundance of the four vascular plant species: Diospyros toposia var. toposoides, Croton cascarilloides, Kibatalia laurifolia, and Mallotus peltatus. Plots with lower soil temperatures had more snails than plots with higher soil temperatures. Our results show that plots in the southern part of the limestone hill, in which P. elephas were absent, were similar in habitat, topography, microclimate, and vegetation to the plots in the northern part of the limestone hill, where specimens were mostly present. The absence of this species in suitable habitats may be due to their low dispersal ability rather than adverse environmental conditions.

Land snails and slugs of Bau limestone hills, Sarawak (Malaysia, Borneo), with the descriptions of 13 new species.

This study presents a list of land snails and slugs found on limestone hills in the District of Bau, the state of Sarawak in Malaysian Borneo. Systematic and random sampling for land snails was conducted at eight limestone outcrops, namely, Gunung Stulang, Padang Pan, Gunung Kapor, Gunung Lobang Angin, Gunung Doya, Gunung Batu, Bukit Sekunyit and Gunung Sebayat. A total of 122 land snail species was documented with photographs of each species. Of the 122 species collected, 13 are new to science, namely, Acmella bauensis sp. nov., Japonia bauensis sp. nov., Plectostoma margaretchanae sp. nov., Microcystina arabii sp. nov., Microcystina atoni sp. nov., Microcystina paripari sp. nov., Microcystina lirata sp. nov., Microcystina oswaldbrakeni sp. nov., Microcystina kilat sp. nov., Philalanka jambusanensis sp. nov., Everettia microrhytida sp. nov., Everettia minuta sp. nov., and Paralaoma sarawakensis sp. nov.

A molecular phylogeny of Geotrochus and Trochomorpha species (Gastropoda: Trochomorphidae) in Sabah, Malaysia reveals convergent evolution of shell morphology driven by environmental influences.

There are currently eleven Geotrochus and four Trochomorpha species in Sabah. The primary diagnostic character that separates the two genera is the intensity of sculpture on the shell upper surface. All Trochomorpha species have a coarse nodular sculpture while Geotrochus species has a non-nodular sculpture or smooth shell. However, it is known that shell characters are often evolutionary labile with high plasticity in response to environmental factors. Hence, identifying the phylogenetic and ecological determinants for the shell characters will shed light on the shell-based taxonomy. This study aims to estimate the phylogenetic relationship between Geotrochus and Trochomorpha species in Sabah based in two mitochondrial genes (COI, 16S) and one nuclear gene (ITS) and also to examine the influence of temperature, elevation and annual precipitation on the coarseness of shell upper surface sculpture and shell sizes of the species of both genera. Additionally, we also investigated the phylogenetic signal of the shell characters. The phylogenetic analysis showed that Geotrochus and Trochomorpha species are not reciprocally monophyletic. The phylogenetic signal test suggested that shell size and upper surface sculpture are homoplastic, and these shell traits are strongly influenced by elevation and annual precipitation, particularly at the cloud zone of Mount Kinabalu. The highland species of both genera have a coarser shell surface than lowland species. The shell and aperture width decrease with increasing elevation and annual precipitation. In the view of finding above, the current taxonomy of Geotrochus and Trochmorpha in this region and elsewhere that based on shell characters need to be revised with sufficient specimens throughout the distribution range of the two genera.

A comparative evaluation of thermal camera and visual counting methods for primate census in a riparian forest at the Lower Kinabatangan Wildlife Sanctuary (LKWS), Malaysian Borneo.

A number of primate census techniques have been developed over the past half-century, each of which have advantages and disadvantages in terms of resources required by researchers (e.g., time and costs), availability of technologies, and effectiveness in different habitat types. This study aims to explore the effectiveness of a thermal imaging technique to estimate the group size of different primate species populations in a degraded riparian forest in the Lower Kinabatangan Wildlife Sanctuary (LKWS), Sabah. We compared this survey technique to the conventional visual counting method along the riverbank. For 38 days, a total of 138 primate groups were observed by thermal camera and visually throughout the study. Optimal conditions for the thermal camera were clear weather, not more than 100 m distance from the observer to the targeted area, boat speed ranging between 5 and 12 km/h, and early morning between 04:30 and 05:30 am. The limitations of the thermal cameras include the inability to identify individual species, sexes, age classes, and also to discern between animals closely aggregated (i.e., mothers with attached infants). Despite these limitations with the thermal camera technique, 1.78 times more primates were detected than counting by eye (p < 0.001), showing the potential benefit of using thermal cameras as an important tool in primate surveys. Nevertheless, ground truthing must be conducted immediately after, or simultaneously during, the thermal survey to verify the species of animals observed on the thermal imagery.

Molecular phylogenetics and evolutionary history of the endemic land snail genus Everettia in northern Borneo.

Borneo has gone through dramatic changes in geology and topography from the early Eocene until the early Pliocene and experienced climatic cycling during the Pleistocene. However, how these changes have shaped the present-day patterns of high diversity and complex distribution are still poorly understood. In this study, we use integrative approaches by estimating phylogenetic relationships, divergence time, and current and past niche suitability for the Bornean endemic land snail genus Everettia to provide additional insight into the evolutionary history of this genus in northern Borneo in the light of the geological vicariance events and climatic fluctuations in the Pleistocene. Our results show that northern Borneo Everettia species belong to two deeply divergent lineages: one contains the species that inhabit high elevation at the central mountain range, while the other contains lowland species. Species diversification in these lineages has taken place before the Pliocene. Climate changes during the Pleistocene did not play a significant role in species diversification but could have shaped contemporary species distribution patterns. Our results also show that the species-rich highland habitats have acted as interglacial refugia for highland species. This study of a relatively sedentary invertebrate supports and enhances the growing understanding of the evolutionary history of Borneo. Species diversification in Everettia is caused by geological vicariance events between the early Miocene and the Pliocene, and the distribution patterns were subsequently determined by climatic fluctuations in the Pleistocene.

Investigating leaf beetles (Coleoptera, Chrysomelidae) on the west coast islands of Sabah via checklist-taking and DNA barcoding.

Sabah is a province of Malaysia located on the northern part of the island of Borneo. Most of the leaf beetle fauna studies from this region conducted over the past 15 years have focussed on the mainland habitats while the leaf beetle fauna from island habitats (ca. 500 islands) have largely been overlooked. This study looks into the leaf beetle fauna of 13 small satellite islands off the west coast of Sabah. All specimens were first sorted into morpho-species operational taxonomic unit (OTU) before being identified to species rank where possible based on morphological characters and species names assigned when the specimens fitted the description of species in the literature. We collected 75 OTUs from 35 genera and five subfamilies according to morphology, 12 of which were identifiable to species level. In addition, the DNA barcode for each OTU was cross checked with records in GenBank and Barcoding of Life Data system (BOLD) to verify their identity. The number of species recorded was reduced from 12 species and 63 OTUs (total 75 OTUs) to 12 species and 56 OTUs (total 68 OTUs) after removal of the colour polymorphic species based on DNA barcode analyses. Pulau Gaya has the highest species richness and Pulau Sulug has the lowest species richness. A total of 64 Barcode Index Numbers consisting of 101 DNA barcodes were obtained from the 12 leaf beetle species and 48 OTUs. Based on the DNA barcode analyses, it was possible to confirm several polymorphic OTUs and cryptic species. The mean intraspecific and interspecific genetic divergence were determined as 0.77% and 16.11%, respectively. DNA barcodes of this study show a low similarity with records in GenBank and BOLD, highlighting the lack of representation and the urgency of studying leaf beetles from this region. The study provides the first documentation of leaf beetle fauna from island habitats of Sabah and the first DNA barcoding data for leaf beetles from this part of the world, with the next steps being larger scale sampling over a wider geographical scale for a better understanding of tropical arthropod diversity.

Land snail fauna in Gunung Kuang Limestone Hill, Perak, Malaysia and its conservation implications (Mollusca, Gastropoda).

This paper presents the first land snail species checklist for Gunung Kuang (Kuang Hill), a limestone hill located next to Gunung Kanthan that is recognised as one of the most important limestone hills for its diverse land snail fauna in Kinta Valley. Samplings were carried out at five plots in Gunung Kuang. This survey documented 47 land snail species, in which six species were identified as unique to Gunung Kuang. Approximately half of the land snails from Gunung Kanthan were found in Gunung Kuang. In addition, one of six unique species from Gunung Kanthan was also found in Gunung Kuang. These rich land snail species in Gunung Kuang are similar to other hills in Kinta Valley, but it is relatively lesser than the adjacent Gunung Kanthan. In view of Gunung Kuang's unique land snail species, and its location closest to disturbed Gunung Kanthan, Gunung Kuang should be considered in the conservation management plan for Gunung Kanthan.

Cryptic diversity: Two morphologically similar species of invasive apple snail in Peninsular Malaysia.

Invasive snails in the genus Pomacea have spread across Southeast Asia including Peninsular Malaysia. Their effects on natural and agricultural wetlands are appreciable, but species-specific effects are less clear because of morphological similarity among the species. Our objective was to establish diagnostic characteristics of Pomacea species in Malaysia using genetic and morphological criteria. The mitochondrial COI gene of 52 adult snails from eight localities in Peninsular Malaysia was amplified, sequenced, and analysed to verify species and phylogenetic relationships. Shells were compared using geometric morphometric and covariance analyses. Two monophyletic taxa, P. canaliculata and P. maculata, occurred in our samples. The mean ratio of shell height: aperture height (P = 0.042) and shell height: shell width (P = 0.007) was smaller in P. maculata. P. maculata co-occurred with P. canaliculata in five localities, but samples from three localities contained only P. canaliculata. This study is the first to confirm the presence of two of the most invasive species of Pomacea in Peninsular Malaysia using a molecular technique. P. canaliculata appears to be the more widespread species. Despite statistical differences, both quantitative and qualitative morphological characteristics demonstrated much interspecific overlap and intraspecific variability; thus, shell morphology alone cannot reliably verify species identity. Molecular techniques for distinguishing between these two highly invasive Pomacea species are needed to understand their specific ecological niches and to develop effective protocols for their management.

Land snails of Leptopoma Pfeiffer, 1847 in Sabah, Northern Borneo (Caenogastropoda: Cyclophoridae): an analysis of molecular phylogeny and geographical variations in shell form.

Leptopoma is a species rich genus with approximately 100 species documented. Species-level identification in this group has been based on shell morphology and colouration, as well as some anatomical features based on small sample sizes. However, the implications of the inter- and intra-species variations in shell form to the taxonomy of Leptopoma species and the congruency of its current shell based taxonomy with its molecular phylogeny are still unclear. There are four Leptopoma species found in Sabah, Borneo, and their taxonomy status remains uncertain due to substantial variation in shell forms. This study focuses on the phylogenetic relationships and geographical variation in shell form of three Leptopoma species from Sabah. The phylogenetic relationship of these species was first estimated by performing Maximum Likelihood and Bayesian analysis based on mitochondrial genes (16S rDNA and COI) and nuclear gene (ITS-1). Then, a total of six quantitative shell characters (i.e., shell height, shell width, aperture height, aperture width, shell spire height, and ratio of shell height to width) and three qualitative shell characters (i.e., shell colour patterns, spiral ridges, and dark apertural band) of the specimens were mapped across the phylogenetic tree and tested for phylogenetic signals. Data on shell characters of Leptopoma sericatum and Leptopoma pellucidum from two different locations (i.e., Balambangan Island and Kinabatangan) where both species occurred sympatrically were then obtained to examine the geographical variations in shell form. The molecular phylogenetic analyses suggested that each of the three Leptopoma species was monophyletic and indicated congruence with only one of the shell characters (i.e., shell spiral ridges) in the current morphological-based classification. Although the geographical variation analyses suggested some of the shell characters indicating inter-species differences between the two Leptopoma species, these also pointed to intra-species differences between populations from different locations. This study on Leptopoma species is based on small sample size and the findings appear only applicable to Leptopoma species in Sabah. Nevertheless, we anticipate this study to be a starting point for more detailed investigations to include the other still little-known (ca. 100) Leptopoma species and highlights a need to assess variations in shell characters before they could be used in species classification.

Inferring microevolution from museum collections and resampling: lessons learned from Cepaea.

Natural history collections are an important and largely untapped source of long-term data on evolutionary changes in wild populations. Here, we utilize three large geo-referenced sets of samples of the common European land-snail Cepaea nemoralis stored in the collection of Naturalis Biodiversity Center in Leiden, the Netherlands. Resampling of these populations allowed us to gain insight into changes occurring over 95, 69, and 50 years. Cepaea nemoralis is polymorphic for the colour and banding of the shell; the mode of inheritance of these patterns is known, and the polymorphism is under both thermal and predatory selection. At two sites the general direction of changes was towards lighter shells (yellow and less heavily banded), which is consistent with predictions based on on-going climatic change. At one site no directional changes were detected. At all sites there were significant shifts in morph frequencies between years, and our study contributes to the recognition that short-term changes in the states of populations often exceed long-term trends. Our interpretation was limited by the few time points available in the studied collections. We therefore stress the need for natural history collections to routinely collect large samples of common species, to allow much more reliable hind-casting of evolutionary responses to environmental change.

A review of the land snail genus Alycaeus (Gastropoda, Alycaeidae) in Peninsular Malaysia.

A total of 11 species and 1 subspecies of Alycaeus were recognised in Peninsular Malaysia prior to this study. However, these taxonomic descriptions of Alycaeus taxa were based on limited numbers of examined materials, where a whole spectrum of morphological variations were not accounted for and diagnoses were often provided without sufficient comparison between congeners from across the peninsula. We reviewed Peninsular Malaysian Alycaeus through the examination of 5137 specimens in 522 collection lots from all major museum collections and literature sources. Based on these examined materials, we utilised a more comprehensive revised set of 39 shell and operculum characters, as well as living animal colour to describe all Alycaeus species in this paper. We also noted their habitat and ecology, as well as updated the distribution of each species. Of the 12 previously described taxa, 10 are reconfirmed as present on Peninsular Malaysia (Alycaeus balingensis, Alycaeus carinata, Alycaeus conformis, Alycaeus gibbosulus, Alycaeus kapayanensis, Alycaeus kelantanensis, Alycaeus liratulus, Alycaeus perakensis perakensis, Alycaeus perakensis altispirus and Alycaeus thieroti) and 2 are confirmed as absent from the peninsula (Alycaeus jagori and Alycaeus pyramidalis). A new record of Alycaeus robeleni is reported for Peninsular Malaysia. One species, Chamalycaeus jousseaumei is confirmed as present on the peninsula and is reassigned to Alycaeus. The subspecies Alycaeus perakensis altispirus Möllendorff, 1902, is elevated to species. Examined Peninsular Malaysian materials that do not fit previously recognised species are described as new species. A total of 11 new species are proposed (Alycaeus selangoriensissp. n., Alycaeus costacrassasp. n., Alycaeus ikanensissp. n., Alycaeus alticolasp. n., Alycaeus charasensissp. n., Alycaeus kurauensissp. n., Alycaeus regalissp. n., Alycaeus virgogravidasp. n., Alycaeus senyumensissp. n., Alycaeus expansussp. n., Alycaeus clementsisp. n.). Overall, 23 species of Alycaeus are now recognised in Peninsular Malaysia.

A checklist of land snails from the west coast islands of Sabah, Borneo (Mollusca, Gastropoda).

Sabah, situated in one of the world's biodiversity hotspots, has the largest number of islands in Malaysia with more than 500 of various sizes and degrees of isolation. However, information on the islands' biodiversity is limited. This study provides an up-to-date checklist of land snail species found on 24 west coast islands in Sabah. A total of 67 species (nearly 20% of the total number of land snail species in the state) representing 37 genera and 19 families is enumerated based on systematic field surveys of 133 sampling plots, BORNEENSIS database records and species checklists published between 2000 and 2016. The number of species on the islands ranges from four to 29. Labuan Island has the highest number of species (29), followed by Tiga Island (25), Mantanani Besar Island (24) and Gaya Island (23). However, the populations of some land snail species may have declined as several previously recorded species on the islands were not found in a recent systematic field sampling. This checklist is provided as a baseline inventory for future island land snail studies and to better inform biodiversity conservation plans of marine parks and other islands on the Sabah west coast.

A preliminary checklist of the freshwater snails of Sabah (Malaysian Borneo) deposited in the BORNEENSIS collection, Universiti Malaysia Sabah.

Sabah, a Malaysian state at the north-eastern tip of Borneo, is situated in one of the Earth's biodiversity hotspots yet its freshwater gastropod diversity remains poorly known. An annotated checklist of the freshwater gastropods is presented, based on specimens deposited in the BORNEENSIS collection of the Institute for Tropical Biology and Conservation at Universiti Malaysia Sabah, Malaysia. A KMZ file is also provided, which acts as a repository of digital images and complete collection data of all examined material, so that it can be shared and adapted to facilitate future research.

Diversity and biogeography of land snails (Mollusca, Gastropoda) in the limestone hills of Perak, Peninsular Malaysia.

Limestone hills are now gaining global conservation attention as hotspots for short-range endemic species. Levels of land snail endemism can be high at limestone hills, especially at hill clusters that are geographically isolated. In the State of Perak, Peninsular Malaysia, limestone hills have been opportunistically surveyed for land snails in the past, but the majority have yet to be surveyed. To address this knowledge gap, we systematically surveyed the terrestrial malacofauna of 12 limestone hills that, based on our opinion, are a representation of the limestone land snail assemblages within the State. Our inventory yielded high sampling completeness (>85%). We found 122 species of land snails, of which 34 species were unique to one of the surveyed hills. We identified 30 species that are potentially new to science. The number of land snail species recorded at each hill ranged between 39 and 63 species. Four of the sampled limestone hills namely, Prk 01 G. Tempurung, Prk 55 G. Pondok, Prk 47 Kanthan, and Prk 64 Bt Kepala Gajah, have high levels of species richness and unique species, representing 91% of the total species recorded in this study. We identified two clusters of limestone hills in central Perak with distinct differences in land snail species composition - a northern hill cluster on elevated granite bedrock and southern hill cluster in a low-lying valley surrounded by alluvial soils. As limestone hills continue to be quarried to meet the cement demand, the four identified limestone hills, along with other hills from the two clusters, warrant urgent conservation attention in order to maintain high species diversity within Perak's terrestrial malacofauna.

A Method for Quantifying, Visualising, and Analysing Gastropod Shell Form.

Quantitative analysis of organismal form is an important component for almost every branch of biology. Although generally considered an easily-measurable structure, the quantification of gastropod shell form is still a challenge because many shells lack homologous structures and have a spiral form that is difficult to capture with linear measurements. In view of this, we adopt the idea of theoretical modelling of shell form, in which the shell form is the product of aperture ontogeny profiles in terms of aperture growth trajectory that is quantified as curvature and torsion, and of aperture form that is represented by size and shape. We develop a workflow for the analysis of shell forms based on the aperture ontogeny profile, starting from the procedure of data preparation (retopologising the shell model), via data acquisition (calculation of aperture growth trajectory, aperture form and ontogeny axis), and data presentation (qualitative comparison between shell forms) and ending with data analysis (quantitative comparison between shell forms). We evaluate our methods on representative shells of the genera Opisthostoma and Plectostoma, which exhibit great variability in shell form. The outcome suggests that our method is a robust, reproducible, and versatile approach for the analysis of shell form. Finally, we propose several potential applications of our methods in functional morphology, theoretical modelling, taxonomy, and evolutionary biology.

Additions to the knowledge of the land snails of Sabah (Malaysia, Borneo), including 48 new species.

We present reviews of the Sabah (Malaysia, on the island of Borneo) species of the following problematical genera of land snails (Mollusca, Gastropoda): Acmella and Anaglyphula (Caenogastropoda: Assimineidae); Ditropopsis (Caenogastropoda: Cyclophoridae); Microcystina (Pulmonata: Ariophantidae); Philalanka and Thysanota (Pulmonata: Endodontidae); Kaliella, Rahula, (Pulmonata: Euconulidae); Trochomorpha and Geotrochus (Pulmonata: Trochomorphidae). Next to this, we describe new species in previously revised genera, such as Diplommatina (Diplommatinidae); Georissa (Hydrocenidae); as well as some new species of genera not revised previously, such as Japonia (Cyclophoridae); Durgella and Dyakia (Ariophantidae); Amphidromus, and Trachia (Camaenidae); Paralaoma (Punctidae); Curvella (Subulinidae). All descriptions are based on the morphology of the shells. We distinguish the following 48 new species: Acmella cyrtoglyphe, Acmella umbilicata, Acmella ovoidea, Acmella nana, Acmella subcancellata, Acmella striata, and Anaglyphula sauroderma (Assimineidae); Ditropopsis davisoni, Ditropopsis trachychilus, Ditropopsis constricta, Ditropopsis tyloacron, Ditropopsis cincta, and Japonia anceps (Cyclophoridae); Diplommatina bidentata and Diplommatina tylocheilos (Diplommatinidae); Georissa leucococca and Georissa nephrostoma (Hydrocenidae); Durgella densestriata, Dyakia chlorosoma, Microcystina microrhynchus, Microcystina callifera, Microcystina striatula, Microcystina planiuscula, and Microcystina physotrochus (Ariophantidae); Amphidromus psephos and Trachia serpentinitica (Camaenidae); Philalanka tambunanensis, Philalanka obscura, Philalanka anomphala, Philalanka rugulosa, and Philalanka malimgunung (Endodontidae); Kaliella eurytrochus, Kaliella sublaxa, Kaliella phacomorpha, Kaliella punctata, Kaliella microsoma, Rahula delopleura, (Euconulidae); Paralaoma angusta (Punctidae); Curvella hadrotes (Subulinidae); Trochomorpha trachus, Trochomorpha haptoderma, Trochomorpha thelecoryphe, Geotrochus oedobasis, Geotrochus spilokeiria, Geotrochus scolops, Geotrochus kitteli, Geotrochus subscalaris, and Geotrochus meristorhachis (Trochomorphidae).

On growth and form of irregular coiled-shell of a terrestrial snail: Plectostoma concinnum (Fulton, 1901) (Mollusca: Caenogastropoda: Diplommatinidae).

The molluscan shell can be viewed as a petrified representation of the organism's ontogeny and thus can be used as a record of changes in form during growth. However, little empirical data is available on the actual growth and form of shells, as these are hard to quantify and examine simultaneously. To address these issues, we studied the growth and form of a land snail that has an irregularly coiled and heavily ornamented shell-Plectostoma concinnum. The growth data were collected in a natural growth experiment and the actual form changes of the aperture during shell ontogeny were quantified. We used an ontogeny axis that allows data of growth and form to be analysed simultaneously. Then, we examined the association between the growth and the form during three different whorl growing phases, namely, the regular coiled spire phase, the transitional constriction phase, and the distortedly-coiled tuba phase. In addition, we also explored the association between growth rate and the switching between whorl growing mode and rib growing mode. As a result, we show how the changes in the aperture ontogeny profiles in terms of aperture shape, size and growth trajectory, and the changes in growth rates, are associated with the different shell forms at different parts of the shell ontogeny. These associations suggest plausible constraints that underlie the three different shell ontogeny phases and the two different growth modes. We found that the mechanism behind the irregularly coiled-shell is the rotational changes of the animal's body and mantle edge with respect to the previously secreted shell. Overall, we propose that future study should focus on the role of the mantle and the columellar muscular system in the determination of shell form.

Association between shell morphology of micro-land snails (genus Plectostoma) and their predator's predatory behaviour.

Predator-prey interactions are among the main ecological interactions that shape the diversity of biological form. In many cases, the evolution of the mollusc shell form is presumably driven by predation. However, the adaptive significance of several uncommon, yet striking, shell traits of land snails are still poorly known. These include the distorted coiled "tuba" and the protruded radial ribs that can be found in micro-landsnails of the genus Plectostoma. Here, we experimentally tested whether these shell traits may act as defensive adaptations against predators. We characterised and quantified the possible anti-predation behaviour and shell traits of Plectostoma snails both in terms of their properties and efficiencies in defending against the Atopos slug predatory strategies, namely, shell-apertural entry and shell-drilling. The results showed that Atopos slugs would first attack the snail by shell-apertural entry, and, should this fail, shift to the energetically more costly shell-drilling strategy. We found that the shell tuba of Plectostoma snails is an effective defensive trait against shell-apertural entry attack. None of the snail traits, such as resting behaviour, shell thickness, shell tuba shape, shell rib density and intensity can fully protect the snail from the slug's shell-drilling attack. However, these traits could increase the predation costs to the slug. Further analysis on the shell traits revealed that the lack of effectiveness in these anti-predation shell traits may be caused by a functional trade-off between shell traits under selection of two different predatory strategies.