Euchlorocystis gen. nov. and Densicystis gen. nov., Two New Genera of Oocystaceae Algae from High‐altitude Semi‐saline Habitat (Trebouxiophyceae,Chlorophyta)

The Oocystaceae family is generally considered to contain common freshwater eukaryotic microalgae, and few are reported living in semi‐saline habitats. Our latest ecological survey in Qinghai Lake and Angzicuo Lake, both large, closed, high‐altitude, semi‐saline lakes located on the Qinghai‐Tibet plateau in China, revealed Oocystaceae species as a dominant group among plankton. Since limited knowledge exists about semi‐saline species in the Oocystaceae family, a taxonomical study was carried out using morphological and phylogenetic methods. Using this approach, four new strains of Oocystaceae were identified and successfully cultured in the lab. Molecular results correlated with morphological characters and resolved these species into at least three genera. A new genus, Euchlorocystis, with type species Euchlorocystis subsalina, is described here as having the distinctive morphology of multiple pyrenoids per chloroplast among Oocystaceae, and an independent phylogenetic position at the base of the Oocystaceae. Similarly, the genus Densicystis, with type species Densicystis glomerata, is newly proposed here as having a unique colony morphology of dozens or hundreds of little cells tightly embedded in ellipsoid to round mucilage masses. Oocystis marina, originally described from the Baltic Sea, was also identified in Qinghai Lake and Angzicuo Lake and phylogenetically positioned in the semi‐saline clade of the Oocystaceae. The result that a marine species was detected in the closed inland lakes implies a further need to reevaluate the origins of these species.     

Bryozoans are returning home: recolonization of freshwater ecosystems inferred from phylogenetic relationships

Bryozoans are aquatic invertebrates that inhabit all types of aquatic ecosystems. They are small animals that form large colonies by asexual budding. Colonies can reach the size of several tens of centimeters, while individual units within a colony are the size of a few millimeters. Each individual within a colony works as a separate zooid and is genetically identical to each other individual within the same colony. Most freshwater species of bryozoans belong to the Phylactolaemata class, while several species that tolerate brackish water belong to the Gymnolaemata class. Tissue samples for this study were collected in the rivers of Adriatic and Danube basin and in the wetland areas in the continental part of Croatia (Europe). Freshwater and brackish taxons of bryozoans were genetically analyzed for the purpose of creating phylogenetic relationships between freshwater and brackish taxons of the Phylactolaemata and Gymnolaemata classes and determining the role of brackish species in colonizing freshwater and marine ecosystems. Phylogenetic relationships inferred on the genes for 18S rRNA, 28S rRNA, COI, and ITS2 region confirmed Phylactolaemata bryozoans as radix bryozoan group. Phylogenetic analysis proved Phylactolaemata bryozoan's close relations with taxons from Phoronida phylum as well as the separation of the Lophopodidae family from other families within the Plumatellida genus. Comparative analysis of existing knowledge about the phylogeny of bryozoans and the expansion of known evolutionary hypotheses is proposed with the model of settlement of marine and freshwater ecosystems by the bryozoans group during their evolutionary past. In this case study, brackish bryozoan taxons represent a link for this ecological phylogenetic hypothesis. Comparison of brackish bryozoan species Lophopus crystallinus and Conopeum seurati confirmed a dual colonization of freshwater ecosystems throughout evolution of this group of animals.     

Extreme,continuous variation in an island snail: local diversification and association of shell form with the current environment

On Rosemary Island, a small continental island (11 km²) in the Dampier Archipelago, Western Australia, snails of the genus Rhagada have extremely diverse morphologies. Their shells vary remarkably in size and shape, with the latter ranging from globose to keeled‐flat, spanning the range of variation in the entire genus. Based primarily on variation in shell morphology, five distinct species are currently recognized. However, a study of 103 populations has revealed continuity of shell form within a very closely‐related group. A phylogenetic analysis of specimens from Rosemary Island, and other islands in the Dampier Archipelago, indicates that much of the morphological variation has evolved on the island, from within a monophyletic group. Within the island, snails with distinct shell morphologies could not be distinguished based on variation in mitochondrial DNA or their reproductive anatomy. The shell variation is geographically structured over a very fine scale, with clines linking the extreme forms over distances less than 200 m. Although there is no evidence that the different forms have evolved in isolation or as a consequence of drift, there is a strong association between keeled‐flat shells and rocky habitats, suggesting that shell shape may be of adaptive significance. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 756–769.     

Bioerosion of gastropod shells: with emphasis on effects of coralline algal cover and shell microstructure

Organisms boring into fifty nine species of gastropod shells on reefs around Guam were the bryozoan Penetrantia clionoides; the acrothoracian barnacles Cryptophialus coronorphorus, Cryptophialus zulloi and Lithoglyptis mitis; the foraminifer Cymbaloporella tabellaeformis, the polydorid Polydora sp. and seven species of clionid sponge. Evidence that crustose coralline algae interfere with settlement of larvae of acrothoracian barnacles, clionid sponges, and boring polychaetes came from two sources: (1) low intensity of boring in limpet shells, a potentially penetrable substrate that remains largely free of borings by virtue of becoming fully covered with coralline algae at a young age and (2) the extremely low levels of boring in the algal ridge, a massive area of carbonate almost entirely covered by a layer of living crustose corallines. There was a strong negative correlation between microstructural hardness and infestation by acrothoracian barnacles and no correlation in the case of the other borers. It is suggested that this points to a mechanical rather than a chemical method of boring by the barnacles. The periostracum, a layer of organic material reputedly a natural inhibitor of boring organisms, was bored by acrothoracican barnacles and by the bryozoan. The intensity of acrothoracican borings is shown to have no correlation with the length of the gastropod shell.     

Cryptic yet colorful: anatomy and relationships of a new genus of Cerithiidae (Caenogastropoda,Cerithioidea) from coral reef drop‐offs

Cerithium koperbergi is a rare gastropod of the family Cerithiidae from the tropical Indo‐West Pacific. The species has a small, unusual shell and often inhabits deeper water, fore‐reef habitats that are atypical for the genus. Anatomical investigations reveal that it possesses a combination of features heretofore considered diagnostic of two main cerithiid subfamilies: Cerithiinae and Bittiinae. While the shell is bittiine, the animal lacks mesopodial pedal glands and possesses a seminal receptacle (vs. a spermatophore bursa) in the lateral lamina of the oviduct, which are considered to be cerithiine features. Re‐evaluation of the anatomy of Bittium reticulatum, the type species of Bittium, indicates the defining anatomical difference in oviduct anatomy between the two subfamilies does not stand up to closer scrutiny. Partial mitochondrial cytochrome c oxidase I (COI) sequences support the interpretation that C. koperbergi is a species complex around the western Pacific rim comprising three divergent mitochondrial lineages. Bayesian analysis of partial mitochondrial COI and 16S rRNA sequences confirm the placement of the C. koperbergi complex within a monophyletic Bittiinae, despite the apparent absence of a unifying anatomical feature. Species in the C. koperbergi complex are here united in Pictorium nov. gen. and two species are described as new. It is hypothesized that features of the midgut may be diagnostic of the Bittiinae, but more comparative data are needed.     

Influence of shell morphometry,microstructure, and thermal conductivity on thermoregulation in two tropical intertidal snails

Tropical intertidal organisms tolerate large fluctuations in temperature and high desiccation rates when exposed during low tide. In order to withstand the short‐term heat stress, intertidal organisms adopt behavioral responses to maximize their survival. Our previous research showed that tropical littorinids found at the upper and lower intertidal shores in Singapore exhibited different behavioral adaptations during low tide. Most of the upper‐shore Echinolittorina malaccana kept a flat orientation, with the aperture against the substrate and the long axis of the shell towards the sun, whereas a majority of the lower‐shore individuals of Echinolittorina vidua stood with the edge of the aperture perpendicular to the substrate on the rocky shore during low tide. This prompted analyses of the shells of these two species to determine whether the differences in the shell morphometry, microstructure, and thermal conductivity of shells of E. malaccana and E. vidua were associated with their respective behavioral responses to thermal stress. Analyses of shell morphometry and thermal conductivity showed that shells of E. malaccana were more likely to minimize heat gain, despite having a higher thermal conductivity on the outer surface, due to their light‐gray, elongated shell. By contrast, the dark‐colored, globose shells of E. vidua probably gain heat more readily through solar radiation. Scanning electron microscopy images of the shells of both littorinid species further revealed that they have cross‐lamellar structure; however, only individuals of E. vidua showed the presence of disjointed rod layers and a pigmented inner shell surface. Individuals of E. malaccana had a rough outer shell surface with holes that inter‐connect to form water‐trapping channels that probably aid cooling. Individuals of E. vidua, however, had a smooth outer surface with rows of kidney‐shaped depressions as microsculptures which probably help to stabilize shell shape. In both Echinolittorina species, behavioral responses were used to overcome thermal stress during low tide that was associated with shell morphometry and shell thermal conductivity. Such combined adaptations increase survivability of the littorinids at their respective tidal levels.     

Phenotypic plasticity in gastropod shell remodeling

A laboratory experiment was conducted with the marine gastropod Littorina littorea from three different sites in Long Island Sound to test for inducible responses to the non‐native predatory crab Hemigrapsus sanguineus. Individuals of L. littorea did not exhibit decreased shell growth in response to increased predation risk, which differs from the response seen in other species of Littorina. However, snails did exhibit slower growth in response to reduced food availability, as seen in other gastropods. Surprisingly, snails from two sites exhibited differences between the crab‐exposed and reduced food‐availability treatments in how they remodeled the thickness of different microstructural layers of shell material in apical parts of the shell. Although individuals of L. littorea did not show the typical shell thickening response seen in other gastropods, they did show differences among populations in remodeling their shell. This underlies the importance of examining how the shell is constructed and remodeled in studies of inducible defenses in gastropods, as well as the importance of examining animals from multiple populations.     

A new boring bryozoan from the Middle Jurassic of the Moscow Region and its micro-CT research

A new boring ctenostome bryozoan without calcareous skeleton, Orbignyopora opulenta sp. nov. (Vesiculariida, Eurystomata), is described from deposits of the Middle Callovian of the Moscow Region. A pioneering micro-ct examination of a colony of this species reveals its diagnostic features hidden in the shell substratum: shape, size, and distinctive features in the budding of zooids and stolons, and also tubular processes (tubulets) developed on the frontal surface of autozooids. The diagnosis of the genus Orbignyopora is added by dimorphism of autozooids (A-zooids and B-zooids) and spindly kenozooids.     

Taphonomy and palaeoecology of Hauterivian–Barremian nerineoid shell beds from the Neuquén Basin,west‐central Argentina

Nerineoid shell beds are described for the first time from Lower Cretaceous deposits of southern South America. These come from carbonates near the top of the Agrio Formation in southern Mendoza Province, west‐central Argentina. To envisage the origin of the nerineoid shell beds, a taphonomic study was carried out, which indicated that these represent within‐habitat time‐averaged, primary sedimentological concentrations with a secondary biogenic imprint related to a relatively high local production of nerineoid shells. The associated palaeoenvironments were studied through a facies analysis of the carbonate succession including the shell beds. The carbonates were deposited in a homoclinal ramp system and depict a shallowing upward trend from mid to inner ramp. The individuals lived and accumulated in oolitic shoals within the inner ramp, in a shallow, well‐lit, high‐energy setting above fair‐weather wave base. Substrate was oxygenated and loose. The nerineoids are shown to belong to one species of the genus Eunerinea, and through the functional morphology of the shells they are tentatively interpreted as infaunal or semi‐infaunal. It is suggested that the recorded monospecific nerineoid shell beds indicate that the palaeoenvironmental conditions may have been favourable for the development of abundant populations of these gastropods in the northern part of the Neuquén Basin during a short time interval in the Hauterivian–Barremian boundary. This could have been related to a brief warming episode, but other factors may have also been involved. □Argentina, Early Cretaceous, Gastropods, nerineoids, Neuquén Basin, shell beds, taphonomy.     

A pre‐Miocene Irano‐Turanian cradle: Origin and diversification of the species‐rich monocot genus Gagea (Liliaceae)

The Irano‐Turanian (IT) floristic region is considered an important center of origin for many taxa. However, there is a lack of studies dealing with typical IT genera that also occur in neighboring areas. The species‐rich monocot genus Gagea Salisb. shows a center of diversity in IT region and a distribution in adjacent regions, therefore representing a good study object to investigate spatial and temporal relationships among IT region and its neighboring areas (East Asia, Euro‐Siberia, Himalaya, and Mediterranean). We aimed at (a) testing the origin of the genus and of its major lineages in the IT region, (b) reconstructing divergence times, and (c) reconstructing colonization events. To address these problems, sequences of the ribosomal DNA internal transcribed spacer (ITS) region of 418 individuals and chloroplast intergenic spacers sequences (psbA‐trnH, trnL‐trnF) of 497 individuals, representing 116 species from all sections of the genus and nearly its entire distribution area were analyzed. Divergence times were estimated under a random molecular clock based on nrITS phylogeny, which was the most complete data set regarding the representation of species and distribution areas. Ancestral distribution ranges were estimated for the nrITS data set as well as for a combined data set, revealing that Gagea most likely originated in southwestern Asia. This genus first diversified there starting in the Early Miocene. In the Middle Miocene, Gagea migrated to the Mediterranean and to East Asia, while migration into Euro‐Siberia took place in the Late Miocene. During the Pleistocene, the Arctic was colonized and Gagea serotina, the most widespread species, reached North America. The Mediterranean basin was colonized multiple times from southwestern Asia or Euro‐Siberia. Most of the currently existing species originated during the last 3 Ma.     

Systematics, shell structure and affinities of the Palaeozoic Problematicum Cornulites

The genus Cornulites, with the type species C. serpularius Schlotheim, 1820, from the Silurian of Gotland, comprises annulated, conical or tubular calcite shells, often found attached to the hard parts of other organisms. No consensus has ever been reached over the zoological affinities of the taxon, and no examples of soft‐part preservation are known: detailed examination of shell structures and growth patterns provide the only means of assessing its systematic position. Using transverse and longitudinal thin sections of C. serpularius Vine, 1882, and C. cellulosus sp. nov. , from the Much Wenlock Limestone Formation of England, the shell structure of Cornulites is shown to be lamellar, but with conspicuous internal chambers (camerae) at the apical end of the shell and, particularly in C. cellulosus, numerous smaller vacuities (cellulae) between the lamellae in the apertural shell region. Growth of the shell was by the secretion of low‐magnesian calcite increments within one another, giving a cone‐in‐cone structure, with the prominent development of cellulae in C. cellulosus probably a constructional feature relating to an upright life position. By comparison of morphology and shell structure with other taxa, the zoological affinities of Cornulites are re‐examined; previously suggested affinities with annelids, foraminifers, molluscs and poriferans can be ruled out. Specific shell structures, most notably pseudopuncta similar to those of bryozoans and brachiopods, have led some recent workers to interpret cornulitids as lophophorates. However, it is shown that they can be interpreted alternatively as solitary, aseptate members of the stem‐Zoantharia (Cnidaria: Anthozoa). Four cornulitid species are recognized in the Much Wenlock Limestone Formation: C. cellulosus sp. nov. , C. gremialis sp. nov. , C. scalariformis and C. serpularius. In the absence of the type material, C. serpularius is here restricted to cornulitids closely resembling the specimens originally figured by Schlotheim. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 150 , 681–699.     

A comparison of indices and measured values of eggshell thickness of different shell regions using museum eggs of 230 European bird species

The thickness of avian eggshells is used to assess shell quality in wild and domestic species, as an indicator of environmental pollution and as an adaptive explanation for shell maculation. Both direct measurements and calculated eggshell thickness indices (ETI) are used in such research, yet this is the first study to quantify, across a large spectrum of bird families (and thus egg shapes), the correlation between measured thicknesses and ETI. Furthermore, few studies have quantified thickness variation across the entire length of the shell, although this variation may influence both gas transfer and embryonic development. We measured the thickness of 942 eggshells of 230 European bird species from the Class II material at the Natural History Museum, Tring, UK, both in the conventional manner, at the equator through the blowhole and, uniquely, after a single longitudinal, cut at its equator at the blunt and pointed ends. Over half of the samples revealed shell defects, cautioning against the indiscriminate use of museum specimens. Strong positive associations were found between species‐specific means of shell thickness with each other and also with ETI, especially those derived from Schönwetter's ‘Handbuch der Oologie’ method, validating the interspecific comparative use of ETI. Thickness measurements and ETI factors are provided for all 230 species. Eggshells were usually thinner at the blunt end (the location of the air sac) than at the equator, but of equal thickness in passerine eggs. This difference was greatest in species producing elongate eggs and suggests that there is a functional significance of shell thickness variation among species that requires further investigation.     

Two new species in the Chaetoceros socialis complex (Bacillariophyta): C. sporotruncatus and C. dichatoensis,and characterization of its relatives,C. radicans and C. cinctus

The diatom genus Chaetoceros is one of the most abundant and diverse phytoplankton in marine and brackish waters worldwide. Within this genus, Chaetoceros socialis has been cited as one of the most common species. However, recent studies from different geographic areas have shown the presence of pseudo‐cryptic diversity within the C. socialis complex. Members of this complex are characterized by curved chains (primary colonies) aggregating into globular clusters, where one of the four setae of each cell curves toward the center of the cluster and the other three orient outwards. New light and electron microscopy observations as well as molecular data on marine planktonic diatoms from the coastal waters off Chile revealed the presence of two new species, Chaetoceros sporotruncatus sp. nov. and C. dichatoensis. sp. nov. belonging to the C. socialis complex. The two new species are similar to other members of the complex (i.e., C. socialis and C. gelidus) in the primary and secondary structure of the colony, the orientation pattern of the setae, and the valve ultrastructure. The only morphological characters that can be used to differentiate the species of this complex are aspects related to resting spore morphology. The two newly described species are closely related to each other and form a sister clade to C. gelidus in molecular phylogenies. We also provide a phylogenetic status along with the morphological characterization of C. radicans and C. cintus, which are genetically related to the C. socialis complex.     

Uneven Distribution of Mating‐Type Alleles Among Togninia minima Isolates,One of the Causal Agents of Leaf Stripe Disease on Grapevines in Northwest Iran

Togninia minima is the main fungal species associated with grapevine leaf stripe disease worldwide. This species is mainly known from its asexual state in nature; nevertheless, a biallelic heterothallic mating strategy has been confirmed for this species based on in vitro crossing studies. There are no data available on the incidence of an active sexual cycle within the populations of this species in many grapevine‐producing countries as well as Iran. The possibility of a clandestine sexual cycle within the Iranian isolates of T. minima was evaluated by analysing the distribution and frequency of the mating‐type alleles on a microspatial and a macrogeographical scales. Towards this aim, a total of 90 T. minima isolates were recovered from grapevines with esca disease from the vineyards in north and north‐western Iran. A multiplex PCR method previously designed by authors was applied for simultaneous identification and determination of the mating‐type alleles in T. minima populations. The results on the screening of mating‐type alleles using multiplex PCR method revealed the mating‐type identity of 77 isolates as Mat1‐2 and 23 isolates as Mat1‐1. Our results showed that both Mat1‐1 and Mat1‐2 isolates are present in a single vineyard and even on single vines. The distribution of mating‐type alleles in the sampled area skewed from the 1 : 1 ratio (77 : 23); however, co‐occurrence of both mating types in a single vineyard and even on single vines is suggestive for the presence of an active sexual cycle for T. minima in north‐western Iran.     

Phylogenetic and mixed Yule‐coalescent analyses reveal cryptic lineages within two South American marine snails of the genus Crepipatella (Gastropoda: Calyptraeidae)

The presence of sibling species within the marine gastropod genus Crepipatella has complicated the taxonomy of members of the group. Since the establishment of the genus, 15 species have been described, but recent studies have indicated that there are only five valid species, two of which inhabit the coasts of Chile, namely C. dilatata and C. fecunda. The two species are morphologically indistinguishable as adults, but can be differentiated on the basis of their encapsulated developmental stages. The primary aim of this study was to reconstruct phylogeny within the genus, and to establish species limits of C. dilatata and C. fecunda, using mitochondrial DNA data. To this end, we used maximum parsimony, maximum likelihood, and Bayesian inference to reconstruct phylogenies using 589 bp of the cytochrome oxidase I (COI) gene. The mtDNA phylogenies were then used as input in a general mixed Yule‐coalescent (GMYC) analysis to estimate species boundaries. In addition, quarter likelihood mapping was used to test a posteriori the confidence of inner branch patterns in the phylogenetic tree. Both DNA tree‐based and GMYC methods provide support for five isolated lineages within this species complex. Our data also suggest that Late Pleistocene and Holocene fragmentation and subsequent range expansion events may have shaped contemporary genetic patterns of Crepipatella in South America.     

Evolutionary history of spiny‐tailed lizards (Agamidae: Uromastyx) from the Saharo‐Arabian region

The subfamily Uromastycinae within the Agamidae is comprised of 18 species: three within the genus Saara and 15 within Uromastyx. Uromastyx is distributed in the desert areas of North Africa and across the Arabian Peninsula towards Iran. The systematics of this genus has been previously revised, although incomplete taxonomic sampling or weakly supported topologies resulted in inconclusive relationships. Biogeographic assessments of Uromastycinae mostly agree on the direction of dispersal from Asia to Africa, although the timeframe of the cladogenesis events has never been fully explored. In this study, we analysed 129 Uromastyx specimens from across the entire distribution range of the genus. We included all but one of the recognized taxa of the genus and sequenced them for three mitochondrial and three nuclear markers. This enabled us to obtain a comprehensive multilocus time‐calibrated phylogeny of the genus, using the concatenated data and species trees. We also applied coalescent‐based species delimitation methods, phylogenetic network analyses and model‐testing approaches to biogeographic inferences. Our results revealed Uromastyx as a monophyletic genus comprised of five groups and 14 independently evolving lineages, corresponding to the 14 currently recognized species sampled. The onset of Uromastyx diversification is estimated to have occurred in south‐west Asia during the Middle Miocene with a later radiation in North Africa. During its Saharo‐Arabian colonization, Uromastyx underwent multiple vicariance and dispersal events, hypothesized to be derived from tectonic movements and habitat fragmentation due to the active continental separation of Arabia from Africa and the expansion and contraction of arid areas in the region.     

Diversity of Halimeda (Bryopsidales,Chlorophyta) in New Caledonia: a Combined Morphological and Molecular Study

Halimeda is a genus of calcified and segmented green macroalgae in the order Bryopsidales. In New Caledonia, the genus is abundant and represents an important part of the reef flora. Previous studies recorded 19 species that were identified using morphological criteria. The aim of this work was to reassess the diversity of the genus in New Caledonia using morpho‐anatomical examinations and molecular analyses of the plastid tufA and rbcL genes. Our results suggest the occurrence of 22 species. Three of these are reported for the first time from New Caledonia: Halimeda kanaloana, H. xishaensis, and an entity resembling H. stuposa. DNA analyses revealed that the species H. fragilis exhibits cryptic or pseudocryptic diversity in New Caledonia. We also show less conclusive evidence for cryptic species within H. taenicola     

Polygyny does not explain the superior competitive ability of dominant ant associates in the African ant‐plant,Acacia (Vachellia) drepanolobium

The Acacia drepanolobium (also known as Vachellia drepanolobium) ant‐plant symbiosis is considered a classic case of species coexistence, in which four species of tree‐defending ants compete for nesting space in a single host tree species. Coexistence in this system has been explained by trade‐offs in the ability of the ant associates to compete with each other for occupied trees versus the ability to colonize unoccupied trees. We seek to understand the proximal reasons for how and why the ant species vary in competitive or colonizing abilities, which are largely unknown. In this study, we use RADseq‐derived SNPs to identify relatedness of workers in colonies to test the hypothesis that competitively dominant ants reach large colony sizes due to polygyny, that is, the presence of multiple egg‐laying queens in a single colony. We find that variation in polygyny is not associated with competitive ability; in fact, the most dominant species, unexpectedly, showed little evidence of polygyny. We also use these markers to investigate variation in mating behavior among the ant species and find that different species vary in the number of males fathering the offspring of each colony. Finally, we show that the nature of polygyny varies between the two commonly polygynous species, Crematogaster mimosae and Tetraponera penzigi: in C. mimosae, queens in the same colony are often related, while this is not the case for T. penzigi. These results shed light on factors influencing the evolution of species coexistence in an ant‐plant mutualism, as well as demonstrating the effectiveness of RADseq‐derived SNPs for parentage analysis.     

Genome‐wide association analyses reveal polygenic genomic architecture underlying divergent shell morphology in Spanish Littorina saxatilis ecotypes

Gene flow between diverging populations experiencing dissimilar ecological conditions can theoretically constrain adaptive evolution. To minimize the effect of gene flow, alleles underlying traits essential for local adaptation are predicted to be located in linked genome regions with reduced recombination. Local reduction in gene flow caused by selection is expected to produce elevated divergence in these regions. The highly divergent crab‐adapted and wave‐adapted ecotypes of the marine snail Littorina saxatilis present a model system to test these predictions. We used genome‐wide association (GWA) analysis of geometric morphometric shell traits associated with microgeographic divergence between the two L. saxatilis ecotypes within three separate sampling sites. A total of 477 snails that had individual geometric morphometric data and individual genotypes at 4,066 single nucleotide polymorphisms (SNPs) were analyzed using GWA methods that corrected for population structure among the three sites. This approach allowed dissection of the genomic architecture of shell shape divergence between ecotypes across a wide geographic range, spanning two glacial lineages. GWA revealed 216 quantitative trait loci (QTL) with shell size or shape differences between ecotypes, with most loci explaining a small proportion of phenotypic variation. We found that QTL were evenly distributed across 17 linkage groups, and exhibited elevated interchromosomal linkage, suggesting a genome‐wide response to divergent selection on shell shape between the two ecotypes. Shell shape trait‐associated loci showed partial overlap with previously identified outlier loci under divergent selection between the two ecotypes, supporting the hypothesis of diversifying selection on these genomic regions. These results suggest that divergence in shell shape between the crab‐adapted and wave‐adapted ecotypes is produced predominantly by a polygenic genomic architecture with positive linkage disequilibrium among loci of small effect.