Sigurður S. Snorrason

Distribution of male yellow dungflies around ovipasition sites: the effect of body size

Abstract. 1 Spatial and temporal variation in body size of yellow dungflies, Scatophaga stercoraria, gathering on and around cow droppings was studied in an Icelandic population in order to elucidate the effect of male and female size on male mating tactics. 2 Males copulating on droppings were on average larger than males copulating in the grass, but of similar size to males guarding ovipositing females. Males searching on droppings were smaller than males copulating or guarding females on droppings but larger than males copulating in the grass. No such differences were found in female size. 3 Resource‐holding power of males (RHP, i.e. male: female size ratio) differed between the three mating groups and was highest for males on the droppings. Size and RHP clearly affect the tactics of copulating males. Males with low RHP tend to copulate in the grass in spite of the cost of longer copulation duration. We argue that this is caused by risk of takeovers from large searching males. 4 There was no change in male size with the age of individual droppings. Contrary to what might be expected, large searching males are not predominantly found at fresh droppings when the probability of catching unpaired females is highest. We suggest instead that good prospects in taking females over from other males must make the strategy to search for females on older droppings profitable. 5 RHP did not change with age of dropping in the three mating groups. The size of ovipositing females increased with age of dropping, probably reflecting longer copulation and egg‐laying times of large females. 6 We found an overall positive relationship between sizes of male and female partners. This correlation was highly significant for copulating pairs in the grass. This is probably a consequence of males with low RHP copulating in the grass and fights in which larger males take over females from smaller males. A weaker, but significant, correlation was found amongst ovipositing pairs. This must be due to take‐over effects. No size correlation was found for pairs copulating on droppings.

The Relationship between Body and Scale Growth Proportions and Validation of Two Back-Calculation Methods Using Individually Tagged and Recaptured Wild Atlantic Salmon

Abstract Using individually tagged wild Atlantic salmon Salmo salar from two Icelandic rivers, we evaluated scale growth in relation to body growth and validated smolt length estimates derived from the Dahl–Lea and Fraser–Lee back-calculation models. Smolts were measured and individually tagged during their seaward migration, and scale samples were taken. The scale readers accuracy was tested, and the bias was found to be less than 5% on average. Spring freshwater growth measurements (i.e., riverine growth during the spring before seaward migration) were validated using scales from smolts and again from the same individuals as adults. Spring growth was largely underestimated (average = 31–34%) when back-calculated from adult scales. Isometry in body growth and scale growth depended on the fish growth rate. The Dahl–Lea back-calculation model overestimated smolt lengths in both populations (average = 0.76–1.43 cm). Use of the Fraser–Lee back-calculation model resulted in slightly more extensive overestima...

Validation of reference genes for expression studies during craniofacial development in arctic charr.

Arctic charr (Salvelinus alpinus) is a highly polymorphic species and in Lake Thingvallavatn, Iceland, four phenotypic morphs have evolved. These differences in morphology, especially in craniofacial structures are already apparent during embryonic development, indicating that genes important in the formation of the craniofacial features are expressed differentially between the morphs. In order to generate tools to examine these expression differences in Arctic charr, the aim of the present study was to identify reference genes for quantitative real-time PCR (qPCR). The specific aim was to select reference genes which are able to detect very small expression differences among different morphs. We selected twelve candidate reference genes from the literature, identified corresponding charr sequences using data derived from transcriptome sequencing (RNA-seq) and examined their expression using qPCR. Many of the candidate reference genes were found to be stably expressed, yet their quality-rank as reference genes varied considerably depending on the type of analysis used. In addition to commonly used software for reference gene validation, we used classical statistics to evaluate expression profiles avoiding a bias for reference genes with similar expression patterns (co-regulation). Based on these analyses we chose three reference genes, ACTB, UB2L3 and IF5A1 for further evaluation. Their consistency was assessed in an expression study of three known craniofacially expressed genes, sparc (or osteonectin), matrix metalloprotease 2 (mmp2) and sox9 (sex-determining region Y box 9 protein) using qPCR in embryo heads derived from four charr groups at three developmental time points. The three reference genes were found to be very suitable for studying expression differences between the morphotypes, enabling robust detection of small relative expression changes during charr development. Further, the results showed that sparc and mmp2 are differentially expressed in embryos of different Arctic charr morphotypes.

Resource polymorphism and diversity of Arctic charr Salvelinus alpinus in a series of isolated lakes

Morphological, dietary and life-history variation in Arctic charr Salvelinus alpinus were characterized from three geographically proximate, but isolated lakes and one large lake into which they drain in south-western Alaska. Polymorphism was predicted to occur in the first three lakes because S. alpinus tend to become polymorphic in deep, isolated lakes with few co-occurring species. Only one morph was evident in the large lake and two of the three isolated lakes. In the third isolated lake, Lower Tazimina Lake, small and large morphs were found, the latter including two forms differing in growth rate. The small morph additionally differed from the two large forms by having more gill rakers and a deeper body than same-sized individuals of the large morph, consuming more limnetic and fewer benthic resources, having a greater gonado-somatic index and maturing at a smaller size. The two large forms consumed only slightly different foods (more terrestrial insects were consumed by the medium-growth form; more snails by the high-growth form). Trends in consumption of resources with body shape also differed between lakes. Variability in life history of S. alpinus in these Alaskan lakes was as broad as that found elsewhere. This variability is important for understanding lake ecosystems of remote regions where this species is commonly dominant.

Fine-scale parallel patterns in diversity of small benthic Arctic charr (Salvelinus alpinus) in relation to the ecology of lava/groundwater habitats

It is critical to study factors that are important for origin and maintenance of biological diversity. A comparative approach involving a large number of populations is particularly useful. We use this approach to study the relationship between ecological factors and phenotypic diversity in Icelandic Arctic charr (Salvelinus alpinus). Numerous populations of small benthic charr have evolved in lava springs in Iceland. These charr appear morphologically similar, but differ in important morphological features related to feeding. We found a clear relationship between diversity in morphology, diet, and ecological factors among populations. In particular, there were clear differences in morphology and diet between fish coming from habitats where the lava spring flowed on as a stream compared to habitats where the lava spring flowed into a pond. Our study shows that ecological factors are important for the origin and maintenance of biological diversity. The relationship between phenotype and ecological factors are observed on a fine scale, when comparing numerous populations that are phenotypically similar. This strongly suggests that for understanding, managing, and conserving biological diversity important ecological variables have to be taken into the account.

Patterns of MiRNA Expression in Arctic Charr Development

Micro-RNAs (miRNAs) are now recognized as a major class of developmental regulators. Sequences of many miRNAs are highly conserved, yet they often exhibit temporal and spatial heterogeneity in expression among species and have been proposed as an important reservoir for adaptive evolution and divergence. With this in mind we studied miRNA expression during embryonic development of offspring from two contrasting morphs of the highly polymorphic salmonid Arctic charr (Salvelinus alpinus), a small benthic morph from Lake Thingvallavatn (SB) and an aquaculture stock (AC). These morphs differ extensively in morphology and adult body size. We established offspring groups of the two morphs and sampled at several time points during development. Four time points (3 embryonic and one just before first feeding) were selected for high-throughput small-RNA sequencing. We identified a total of 326 conserved and 427 novel miRNA candidates in Arctic charr, of which 51 conserved and 6 novel miRNA candidates were differentially expressed among developmental stages. Furthermore, 53 known and 19 novel miRNAs showed significantly different levels of expression in the two contrasting morphs. Hierarchical clustering of the 53 conserved miRNAs revealed that the expression differences are confined to the embryonic stages, where miRNAs such as sal-miR-130, 30, 451, 133, 26 and 199a were highly expressed in AC, whereas sal-miR-146, 183, 206 and 196a were highly expressed in SB embryos. The majority of these miRNAs have previously been found to be involved in key developmental processes in other species such as development of brain and sensory epithelia, skeletogenesis and myogenesis. Four of the novel miRNA candidates were only detected in either AC or SB. miRNA candidates identified in this study will be combined with available mRNA expression data to identify potential targets and involvement in developmental regulation.

Bones in motion: Ontogeny of craniofacial development in sympatric arctic charr morphs

Background: The impressive diversity in the feeding apparatus often seen among related fish species clearly reflects differences in feeding modes and habitat utilization. Such variation can also be found within species. One example of such intraspecific diversity is the Arctic charr in Lake Thingvallavatn, where four distinct morphs coexist: two limnetic, with evenly protruding jaws, and two benthic, with subterminal lower jaws. We used these recently evolved morphs to study the role of ontogenetic variation in shaping craniofacial diversity. Results: The segmental development of the pharyngeal arches and the order of events in craniofacial development is the same as has been described for teleosts, emphasizing the conserved nature of this process. However, our morphometric analyses reveal differences between morphs. Hatching is accompanied by increase in size and allometric shape changes in Arctic charr. Ontogenetic trajectories of craniofacial shape also differ significantly between morphs. Conclusions: The results point to hatching as a significant developmental event in Arctic charr and possibly other fishes. Also, the developmental origins of limnetic and benthic specializations in the craniofacial elements of Arctic charr may stem from events around hatching. This calls for investigations of the mechanisms and consequences of hatching, in the context of development and evolution. Developmental Dynamics 244:1168–1178, 2015.

Variability in the functional role of Arctic charr Salvelinus alpinus as it relates to lake ecosystem characteristics

This study investigated how dietary habits vary with lake characteristics in a fish species that exhibits extensive morphological and ecological variability, the Arctic charr Salvelinus alpinus. Iceland is a hotspot of geological activity, so its freshwater ecosystems vary greatly in physical and chemical attributes. Associations of dietary items within guts of charr were used to form prey categories that reflect habitat-specific feeding behavior. Six prey categories were defined and dominated by snails (Radix peregra), fish (Gasterosteus aculeatus), tadpole shrimp (Lepidurus arcticus), chironomid pupae, pea clam (Pisidium spp.), and the cladoceran Bosmina sp.. These reflected different combinations of feeding in littoral stone, offshore benthic, and limnetic habitats. Certain habitat-specific feeding strategies consistently occurred alongside each other within lakes. For example, zooplanktivory occurred in the same lakes as consumption from offshore habitats; piscivory occurred in the same lakes as consumption from littoral benthic habitats. Redundancy analyses (RDA) were used to investigate how lake environment was related to consumption of different prey categories. The RDA indicated that piscivory exhibited by Arctic charr was reduced where brown trout were abundant and lakes were shallow, greater zooplanktivory occurred at lower latitudes and under decreased nutrient but higher silicon dioxide concentrations, and benthic resource consumption was associated with shallower lakes and higher altitudes. This study showed that trends previously observed across fish species were supported at the intraspecific level, indicating that a single species with flexible dietary habits can fill functional roles expected of multiple species in more diverse food webs.

Habitat complexity affects how young of the year Atlantic cod Gadus morhua perceive predation threat from older conspecifics

The response of age 0+ year juvenile Atlantic cod Gadus morhua to the presence of age 1+ and age 3+ year conspecifics was measured with and without cover available. Juveniles reacted by aggregating more closely and maintaining distance from older conspecifics in an experimental setting without cover but only to age 3+ year conspecifics when cover was available. The results indicate that prior residence of older juveniles can affect age 0+ year juveniles during benthic settlement and highlights the conservation value of structurally complex nursery habitats.

Differential gene expression during early development in recently evolved and sympatric Arctic charr morphs

Phenotypic differences between closely related taxa or populations can arise through genetic variation or be environmentally induced, leading to altered transcription of genes during development. Comparative developmental studies of closely related species or variable populations within species can help to elucidate the molecular mechanisms related to evolutionary divergence and speciation. Studies of Arctic charr (Salvelinus alpinus) and related salmonids have revealed considerable phenotypic variation among populations and in Arctic charr many cases of extensive variation within lakes (resource polymorphism) have been recorded. One example is the four Arctic charr morphs in the ∼10,000 year old Lake Thingvallavatn, which differ in numerous morphological and life history traits. We set out to investigate the molecular and developmental roots of this polymorphism by studying gene expression in embryos of three of the morphs reared in a common garden set-up. We performed RNA-sequencing, de-novo transcriptome assembly and compared gene expression among morphs during an important timeframe in early development, i.e., preceding the formation of key trophic structures. Expectedly, developmental time was the predominant explanatory variable. As the data were affected by some form of RNA-degradation even though all samples passed quality control testing, an estimate of 3′-bias was the second most common explanatory variable. Importantly, morph, both as an independent variable and as interaction with developmental time, affected the expression of numerous transcripts. Transcripts with morph effect, separated the three morphs at the expression level, with the two benthic morphs being more similar. However, Gene Ontology analyses did not reveal clear functional enrichment of transcripts between groups. Verification via qPCR confirmed differential expression of several genes between the morphs, including regulatory genes such as AT-Rich Interaction Domain 4A (arid4a) and translin (tsn). The data are consistent with a scenario where genetic divergence has contributed to differential expression of multiple genes and systems during early development of these sympatric Arctic charr morphs.