Staffan Bensch

Lund University | LU · Department of Biology

Predicting coevolutionary outcomes is critical for understanding the consequences of climate change and biodiversity loss. The geographic mosaic theory of coevolution posits that coevolutionary interactions vary among populations, facilitating the persistence of such interactions over evolutionary timescales. Characterisation of geographic mosaics...

Many species are currently experiencing range shifts in response to climate change. The colonisation of new regions can have serious genetic consequences for the shifting population, given that repeated founder events and strong genetic drift are expected to erode genetic variation along the expansion axis. Such a loss of genetic diversity could re...

Haemoproteus bobricklefsi sp. nov. (Haemosporida, Haemoproteidae) was found in the dunnock Prunella modularis and represents the first blood parasite described in accentor birds of the Prunellidae. The description is based on the morphology of blood stages and includes information about a barcoding segment of the mitochondrial cytochrome b gene (li...

Background Small songbirds respond and adapt to various geographical barriers during their annual migration. Global flyways reveal the diverse migration strategies in response to different geographical barriers, among which are high-elevation plateaus. However, few studies have been focused on the largest and highest plateau in the world, the Qingh...

Animal migration is a fascinating phenomenon that has puzzled mankind since the time of ancient Greece. It is a process widespread across a varied range of taxa and it shines especially in birds which, because of their mobility, display an amazing diversity of routes and strategies. With the advances in tracking devices and improvements of sequenci...

Avian haemosporidian parasites are useful model organisms to study the ecology and evolution of parasite-host interactions due to their global distribution and extensive biodiversity. Detection of these parasites has evolved from microscopic examination to PCR-based methods, with the mitochondrial cytochrome b gene serving as barcoding region. Howe...

Differences in migratory behavior have been suggested to drive speciation, but the genetics underlying this process remain unknown. Identification and study of migratory divides can help us understand how differential migration can lead to reproductive isolation. Here, we genotyped Willow Warblers Phylloscopus trochilus , from the Åland islands, lo...

Background Small songbirds respond and adapt to various geographical barriers during their annual migration. Global flyways reveal the various migration strategies in response to different geographical barriers, among which are high-elevation plateaus. However, few studies have been focused on the largest and highest plateau in the world, the Qingh...

Stable isotope analysis provides valuable insights into the ecology of long-distance migratory birds during periods spent away from a specific study site. In a previous study, Swedish great reed warblers (Acrocephalus arundinaceus) infected with haemosporidian parasites differed in feather isotope ratios compared to non-infected birds, suggesting t...

Migratoriness in birds is evolutionary labile, with many examples of increasing or decreasing migration distances on the timescale of modern ornithology. In contrast, shifts of migration to more nearby wintering grounds seem to be a slow process. We examine the history of how Palearctic migratory landbirds have expanded their wintering ranges to in...

It is a long-standing view that the main mechanism maintaining narrow migratory divides in passerines is the selection against intermediate and suboptimal migratory direction, but empirical proof of this is still lacking. We present novel results from a willow warbler migratory divide in central Sweden from where birds take the typical SW and SE as...

Background Mitochondrial respiration plays a central role in the survival of many eukaryotes, including apicomplexan parasites. A 479-bp fragment from the mitochondrial cytochrome b gene is widely used as a barcode to identify genetic lineages of avian malaria parasites Plasmodium and related haemosporidians. Here we looked for evidence of selectio...

Background Migratory birds play an important part in the spread of parasites, with more or less impact on resident birds. Previous studies focus on the prevalence of parasites, but changes in infection intensity over time have rarely been studied. As infection intensity can be quantified by qPCR, we measured infection intensity during different sea...

Aim The increasing spread of vector‐borne diseases has resulted in severe health concerns for humans, domestic animals and wildlife, with changes in land use and the introduction of invasive species being among the main possible causes for this increase. We explored several ecological drivers potentially affecting the local prevalence and richness...

Parasite dispersal and host-switching may be better understood by knowing when they occurred. We estimated when the ancestor of a parasite of great reed warblers ( Acrocephalus arundinaceus ) dispersed to the Seychelles and began infecting the endemic Seychelles warbler ( A. sechellensis ). We used mitochondrial genomes and published molecular dive...

Structural rearrangements have been shown to be important in local adaptation and speciation, but have been difficult to reliably identify and characterize in non-model species. Here we combine long reads, linked reads and optical mapping to characterize three divergent chromosome regions in the willow warbler Phylloscopus trochilus, of which two a...

Migratory routes and remote wintering quarters in birds are often species and even population specific. It has been known for decades that songbirds mainly migrate solitarily, and that the migration direction is genetically controlled. Yet, the underlying genetic mechanisms remain unknown. To investigate the genetic basis of migration direction, we...

Background Hosts are often simultaneously infected with several parasite species. These co-infections can lead to within-host interactions of parasites, including mutualism and competition, which may affect both virulence and transmission. Birds are frequently co-infected with different haemosporidian parasites, but very little is known about if an...

Background Sequencing parasite genomes in the presence of host DNA is challenging. Sequence capture can overcome this problem by using RNA probes that hybridize with the parasite DNA and then are removed from solution, thus isolating the parasite DNA for efficient sequencing. Methods Here we describe a set of sequence capture probes designed to ta...

The Bluethroat subspecies Cyanecula svecica svecia occurs throughout the northern Palearctic. European C. s. svecia has been shown to follow a route west of the Himalaya to winter quarters in southern Asia. Out of 30 Bluethroats tagged with geolocators in eastern Siberia, we obtained one track suggesting that it migrated east of the Himalaya to a w...

Relative telomere length (RTL), an indicator of senescence, has been shown to be heritable but can also be affected by environmental factors, such as parental effects. Investigating heritability as well as parental effects and rearing environment can help us to understand the factors affecting offspring telomeres. Moreover, how phenotypic parental...

The genetic basis of bird migration has been the focus of several studies. Two willow warbler subspecies (Phylloscopus trochilus trochilus and Phylloscopus trochilus acredula) follow different migratory routes to wintering grounds in Africa. Their breeding populations overlap in contact areas or “migratory divides” located in central Scandinavia an...

The seasonal timing of moult in migratory birds is an adaptation to cope with time constraints in the annual cycle. Kiat and Izhaki analysed moult patterns in Palaearctic passerines and rejected the proposition that seasonally divided moult is an endogenously controlled strategy. Instead, they advocated the view that it occurs due to a flexible and...

Long-distance migration requires adaptations in a suite of behavioral, physiological and morphological traits. However, almost nothing is known about the genetic basis of these adaptations. The willow warbler Phylloscopus trochilus occurs in Europe with a northern and southern subspecies that show drastically different migration routes and winterin...

Telomere erosion has been proposed to be tightly associated with senescence, environmental stressors and life history trade‐offs. How telomere dynamics vary across life stages and especially in relation to (heritable) phenotypic traits is still unclear. The tawny owl Strix aluco display a highly heritable melanin‐based colour polymorphism, a grey a...

How blood parasite infections influence the migration of hosts remains a lively debated issue as past studies found negative, positive, or no response to infections. This particularly applies to small birds, for which monitoring of detailed migration behavior over a whole annual cycle has been technically unachievable so far. Here, we investigate h...

During the last two decades, molecular methods to study mitochondrial DNA sequence variation have become an important part in the studies of avian haemosporidians. Up until recently, these methods have primarily been used for identification of the parasites and for tentative phylogenetic reconstructions, allowing researchers not trained in traditio...

Parasite species evolve by switching to new hosts, cospeciating with their current hosts, or speciating on their current hosts. Vector transmitted parasites are expected to speciate by host switching, but confirming this hypothesis has proved challenging. Parasite DNA can be difficult to sequence, thus well resolved parasite phylogenies that are ne...

Background: The behaviour of blood-sucking arthropods is a crucial determinant of blood protozoan distribution and hence of host-parasite coevolution, but it is very challenging to study in the wild. The molecular identification of parasite lineages in vectors can be a useful key to understand the behaviour and transmission patterns realised by th...

Avian haemosporidians (Haemosporida) represent a globally distributed, species-rich multiparasite-multihost host-parasite system. Each year, many of these parasite lineages are carried between temperate and tropical regions by migratory birds. While several factors can limit the transmission of avian haemosporidians to new areas, recent studies hav...

Infection patterns of avian haemosporidians result from the evolution of their associations with hosts, and can be shaped by multiple biotic factors. However, at the level of parasite species, few studies have investigated the details of the temporal dynamics of infection patterns in wild bird communities. We hereby studied a wild bird community in...

Backgrounds: Geographic regions, where two closely related taxa with different migration routes come into contact, are known as migratory divides. Hybrids originating from migratory divides are hypothesized to migrate intermediately relative to the parental populations. Few studies have tested this hypothesis in wild birds, and only in hybrids tha...

Many hypotheses attempt to explain parasite‐host associations, but rarely are they examined together in a single community. For hosts, key traits are the proportion of infected individuals (prevalence) and the diversity of parasites infecting them. A key parasite trait is host specificity, ranging from specialists infecting one or a few closely rel...

Background The behaviour of blood-sucking arthropods is a crucial determinant of blood protozoan distribution and hence host-parasite coevolution but is very challenging to study in the wild. The molecular identification of parasite lineages in vectors can be a useful key to understand the behaviour and transmission patterns realised by these vecto...

Background The behaviour of blood-sucking arthropods is a crucial determinant of blood protozoan distribution and hence of host-parasite coevolution, but it is very challenging to study in the wild. The molecular identification of parasite lineages in vectors can be a useful key to understand the behaviour and transmission patterns realised by thes...

Background: The behaviour of blood-sucking arthropods is a crucial determinant of blood protozoan distribution and hence of host-parasite coevolution, but it is very challenging to study in the wild. The molecular identification of parasite lineages in vectors can be a useful key to understand the behaviour and transmission patterns realised by the...

Background: Geographic regions, where two close-related taxa with different migration routes come into contact, are known as migratory divides. Hybrids originating from migratory divides are hypothesized to migrate intermediately relative to the parental populations. However, to date and due to technical limitations, few studies testing this hypoth...

Evolutionary genomics has recently entered a new era in the study of host-pathogen interactions. A variety of novel genomic techniques has transformed the identification, detection and classification of both hosts and pathogens, allowing a greater resolution that helps decipher their underlying dynamics and provides novel insights into their enviro...

Migration can influence host–parasite dynamics in animals by increasing exposure to parasites, by reducing the energy available for immune defense, or by culling of infected individuals. These mechanisms have been demonstrated in several comparative analyses; however, few studies have investigated whether conspecific variation in migration distance...

Avian haemosporidian parasites are particularly diverse and widespread. To date, more than 3,000 distinct cytochrome b lineages have been recorded, of which some present extremely wide geographical distributions, even including multiple continents. Whether these isolates represent one or several cryptic species remains unknown. Here we carried out...

If the absence of the appropriate vector(s) is the limiting condition the infection will be successfully developed in experimental birds. P. relictum (GRW4 cyt.b lineage) in House Sparrow (Passer domesticus) P. ashfordi (GRW2 cyt.b lineage) in House Sparrow, Tree Sparrow (Passer montanus) and Song Thrush (Turdus philomelus)

Evolutionary genomics has recently entered a new era in the study of host-pathogen interactions. A variety of novel genomic techniques has transformed to the identification, detection and classification of both hosts and pathogens, allowing a greater resolution that helps decipher their underlying dynamics and provides novel insights into their env...

Evolutionary genomics has recently entered a new era in the study of host-pathogen interactions. A variety of novel genomic techniques has transformed to the identification, detection and classification of both hosts and pathogens, allowing a greater resolution that helps decipher their underlying dynamics and provides novel insights into their env...

Long-distance migrants with transcontinental breeding ranges are of particular interest for the study of local adaptation and geographic differentiation in birds. We compared phenotypes and genotypes between Far East Siberian Willow Warblers Phylloscopus trochilus yakutensis Ticehurst, 1935 with the European subspecies P. t. trochilus Linnaeus, 175...

Accurate detection and identification are essential components for epidemiological, ecological, and evolutionary surveys of avian haemosporidian parasites. Microscopy has been used for more than 100 years to detect and identify these parasites; however, this technique requires considerable training and high-level expertise. Several PCR methods with...

Members of the order Haemosporida are protist parasites that infect mammals, reptiles and birds. This group includes the causal agents of malaria, Plasmodium parasites, the genera Leucocytozoon and Fallisia, as well as the species rich genus Haemoproteus with its two subgenera Haemoproteus and Parahaemoproteus. Some species of Haemoproteus cause se...

Avian malaria and related haemosporidians (Plasmodium, [Para]Haemoproteus, and Leucocytoozoon) represent an exciting multi‐host, multi‐parasite system in ecology and evolution. Global research in this field accelerated after 1) the publication in 2000 of PCR protocols to sequence a haemosporidian mitochondrial (mtDNA) barcode, and 2) the developmen...

Aim The role of migratory birds in the spread of parasites is poorly known, in part because migratory strategies and behaviours potentially affecting transmission are not easy to study. We investigated the dynamics of infection by blood parasites through the annual cycle of a long‐distance Nearctic–Neotropical migratory songbird to examine the role...

Human‐induced climate change is expected to cause major biotic changes in species distributions and thereby including escalation of novel host‐parasite associations. Closely related host species that come into secondary contact are especially likely to exchange parasites and pathogens. Both the Enemy Release Hypothesis (where invading hosts escape...

Background: High-latitude bird migration has evolved after the last glaciation, in less than 10,000-15,000 years. Migrating songbirds rely on an endogenous migratory program, encoding timing, fueling, and routes, but it is still unknown which compass mechanism they use on migration. We used geolocators to track the migration of willow warblers (Ph...

Throughout the world, vulture populations have suffered precipitous declines in recent decades. The Cinereous Vulture Aegypius monachus, a highly philopatric scavenger distributed across southern Europe and the central Asian plateau is threatened in many parts of its range. Turkey holds the second largest population of this species in the Western P...

Avian malaria and related haemosporidians (Plasmodium, [Para]Haemoproteus, and Leucocytoozoon) represent an exciting multi-host, multi-parasite system in ecology and evolution. Global research in this field accelerated after 1) the publication in 2000 of PCR protocols to sequence a haemosporidian mitochondrial (mtDNA) barcode, and 2) the developmen...

Parasites that can infect multiple host species are considered to be host generalists with low host specificity. However, whether generalist parasites are better adapted to a subset of their host species remains unknown. To elucidate this possibility, we compared the variation in prevalence and infection intensity among host species of three genera...

Parasites can vary in the number of host species they infect, a trait known as "host specificity". Here we quantify phylogenetic signal-the tendency for closely related species to resemble each other more than distantly related species-in host specificity of avian haemosporidian parasites (genera Plasmodium, Haemoproteus and Leucocytozoon) using da...

Genomic sequencing of avian haemosporidian parasites (Haemosporida) has been challenging due to excessive contamination from host DNA. In this study, we developed a cost-effective protocol to obtain parasite sequences from naturally infected birds, based on targeted sequence capture and next generation sequencing. With the genomic data of Haemoprot...

The biogeographic histories of parasites and pathogens are infrequently compared with those of free-living species, including their hosts. Documenting the frequency with which parasites and pathogens disperse across geographic regions contributes to understanding not only their evolution, but also the likelihood that they may become emerging infect...

Lack of reliable information on the status of species often leads managers to exclusively rely on experiential knowledge, opinions or perceptions, usually derived from personnel associated with natural resource management agencies. Yet, the accuracy of these sources of information remains largely untested. We approached this challenge, which is par...

Supplementary information for "Counting bears in the Iranian Caucasus: Remarkable mismatch between scientifically sound population estimates and perceptions" by E. M. Moqanaki, J. Jiménez, S. Bensch and J. V. López-Bao

The light-dependent magnetic compass of birds provides orientation information about the spatial alignment of the geomagnetic field. It is proposed to be located in the avian retina, and be mediated by a light-induced, biochemical radical-pair mechanism involving cryptochromes as putative receptor molecules. At the same time, cryptochromes are know...

Heterogeneous patterns of genomic differentiation are commonly documented between closely related populations and there is considerable interest in identifying factors that contribute to their formation. These factors could include genomic features (e.g., areas of low recombination) that promote processes like linked selection (positive or purifyin...

Table S1. Datasets included in the present study. Table S2. Coverage of consensus genomes. Table S3. Repeatability measured using outlier status and overlap values. Table S4. Data used to measure the speciation continuum.

Table S5. Results from GLMs examining the relationship between repeatability and predictor variables related to genomic features.

Vector and host communities, as well as habitat characteristics, may have important but different impacts on the prevalence, richness and evenness of vector‐borne parasites. We investigated the relative importance of (1) the mosquito community composition, (2) the vertebrate community composition and (3) landscape characteristics on the prevalence,...

Parasites often have reduced genomes as their own genes become redundant when utilizing their host as a source of metabolites, thus losing their own de novo production of metabolites. Primate malaria parasites can synthesize vitamin B 1 (thiamine) de novo but rodent malaria and other genome-sequenced apicomplexans cannot, as the three essential gen...

Parasite-host and insect-plant research have divergent traditions despite the fact that most phytophagous insects live parasitically on their host plants. In parasitology it is a traditional assumption that parasites are typically highly specialized; cospeciation between parasites and hosts is a frequently expressed default expectation. Insect-plan...

Heterogeneous patterns of genomic differentiation are commonly documented between closely related populations and there is considerable interest in identifying factors that contribute to their formation. These factors could include genomic features (e.g., areas of low recombination) that promote processes like linked selection (positive or purifyin...

Urban environments pose novel challenges, as well as opportunities, for urban-dwelling wildlife. Although differences have been reported in several phenotypic traits (e.g. morphology, physiology and behaviour) between urban and rural populations, it is poorly understood whether this affects individual fitness. Telomere dynamics are posited as one p...

Over 50 avian Plasmodium species have been described. However, PCR-based information shows much broader diversity of genetic lineages in these parasites. This discrepancy indicates insufficient knowledge about taxonomic diversity and boundaries of a single species in avian Plasmodium species. In recent taxonomy, most of genetically closely related...

Abstract Parasites are expected to exert long-term costs on host fecundity and longevity. Understanding the consequences of heritable polymorphic variation in disease defence in wild populations is essential in order to predict evolutionary responses to changes in disease risk. Telomeres have been found to shorten faster in malaria-diseased indivi...

It is well established that differences in migratory behavior between populations of songbirds have a genetic basis but the actual genes underlying these traits remains largely unknown. In an attempt to identify such candidate genes we de novo assembled the genome of the willow warbler Phylloscopus trochilus, and used whole-genome resequencing and...