Michael C. Singer

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• BA Oxford 1967; PhD Stanford 1971
• Student at Theoretical and Experimental Ecology Station

Abstract As plant species expand their upper limits of distribution under current warming, some retain both traditional climate space and biotic environment while others encounter novel conditions. The latter is the case for Rhododendron campanulatum, a woody shrub that grows both above and below treeline at our study site in the Eastern Himalayas...

Despite growing evidence that "connectedness" of humans with nature creates multiple benefits for both humans and nature, these benefits are not fully considered by health and conservation policymakers. Studies are scattered across scientific disciplines including health, education, psychology and biology, making it difficult to get a complete over...

The Working Group II contribution to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change (IPCC) provides a comprehensive assessment of the scientific literature relevant to climate change impacts, adaptation and vulnerability. The report recognizes the interactions of climate, ecosystems and biodiversity, and human societie...

Insects have been key players in assessments of biodiversity impacts of anthropogenically-driven environmental change, including the evolutionary and ecological impacts of climate change. Populations of Edith's Checkerspot Butterfly (Euphydryas editha) adapt rapidly to diverse environmental conditions, with numerous high-impact studies documenting...

Evolutionary change impacts the rate at which insect pests, pollinators, or disease vectors expand or contract their geographic ranges. Although evolutionary changes, and their ecological feedbacks, strongly affect these risks and associated ecological and economic consequences, they are often underappreciated in management efforts. Greater rigor a...

Studies in birds and trees show climatic stresses distributed across species' ranges, not only at range limits. Here, new analyses from the butterfly Euphydryas editha reveal mechanisms generating these stresses: geographic mosaics of natural selection, acting on tradeoffs between climate adaptation and fitness traits, cause some range-central popu...

Scientists often need to know whether pairs of entities tend to occur together or independently. Standard approaches to this issue use co-occurrence indices such as Jaccard, Sørensen-Dice, and Simpson. We show that these indices are sensitive to the prevalences of the entities they describe and that this invalidates their interpretability. We propo...

Internationally agreed sustainability goals are being missed. Here, we conduct global meta-analyses to assess how the extent to which humans see themselves as part of nature-known as human-nature connectedness (HNC)-can be used as a leverage point to reach sustainability. A meta-analysis of 147 correlational studies shows that individuals with high...

Internationally-agreed sustainability goals are being missed. Here, we document a potential reason for this failure and show how the extent to which humans see themselves as part of nature – known as human-nature connectedness (HNC) – can be used as a leverage point for increasing public engagement towards sustainability targets. We conduct the fir...

Internationally-agreed sustainability goals are being missed. Here, we show how the extent to which humans see themselves as part of nature –known as human-nature connectedness (HNC) –can be used as a leverage point for reaching sustainability. We conduct the first global meta-analysis of the HNC literature. Meta-analysis of 147 correlational studi...

As species' poleward range limits expand under climate change, generalists are expected to be better colonists than specialists, extending their ranges faster. This effect of specialization on range shifts has been shown, but so has the reverse cause‐effect: in a global meta‐analysis of butterfly diets it was range expansions themselves that caused...

This review was solicited as an autobiography. The “problems” in my title have two meanings. First, they were professional difficulties caused by my decision to study oviposition preferences of butterflies that were not susceptible to traditional preference-testing designs. Until I provided video, my claim that the butterflies duplicate natural pos...

This is a reponse to Kharouba and Wolkovich's (2020) review of consumer-resource phenological synchrony. They provide a valuable review and cogent advocacy for future work. However, they misunderstand and misinterpret examples from plant-insect interactions. Their detailed case study involves phenological synchrony/ asynchrony between spring hatchi...

Dynamics of herbivorous insect diet breadth are important in generation of novel pests, biological control of weeds and as indicators of global change impacts. But what forces and events drive these dynamics? Here we present evidence for a novel scenario: that specialization increases in persistent populations, but that, at the species level, this...

Alpine treelines are expected to shift upward due to recent climate change. However, interpretation of changes in montane systems has been problematic because effects of climate change are frequently confounded with those of land use changes. The eastern Himalaya, particularly Langtang National Park, Central Nepal, has been relatively undisturbed f...

We illustrate an evolutionary host shift driven by increased fitness on a novel host, despite maladaptation to it in six separate host‐adaptive traits. Here, local adaptation is defined as possession of traits that provide advantage in specific environmental contexts; thus individuals can have higher fitness in benign environments to which they are...

1. This study provides evidence that a heliophilic butterfly, the Glanville fritillary (Melitaea cinxia) has adapted differently to environmental variation across latitudes and elevations. 2. In cool air, basking M. cinxia orient themselves perpendicular to the sun's rays to gain heat and take off. During flight, solar heating is reduced because or...

Global transport of organisms by humans provides novel resources to wild species, which often respond maladaptively. Native herbivorous insects have been killed feeding on toxic exotic plants, which acted as 'ecological traps'1-4. We document a novel 'eco-evolutionary trap' stemming from the opposite effect; that is, high fitness on an exotic resou...

Gregarious, social caterpillars have stimulated research because group size may affect survival, growth rate, thermoregulation, and interactions with other species, yet group size is often variable both within and among populations. We used a combination of observations and experiments to study the importance of group size for egg and larval surviv...

Clumped distributions of herbivorous insect eggs often result from independent assessments of individual plants by different ovipositing females. Here we ask whether, in addition, plants might be rendered more or less attractive to ovipositing Melitaea cinxia butterflies by presence of conspecific eggs and/or by prior larval attack. Both eggs and l...

Studies of heat shock response show a correlation with local climate, although this is more often across altitudinal than latitudinal gradients. In the present study, differences in constitutive but not inducible components of heat shock response are detected among populations of the Glanville fritillary butterfly Melitaea cinxia L. that exist at t...

The work of the butterfly field biologist typically involves travels with humans and larvae interspersed in the same vehicle, spontaneous interrogations from passing strangers or park rangers, and attempts to perform meaningful research in collaboration with disparate personalities at remote sites with no facilities. In the course of this work, I b...

Relationships between biased dispersal and local adaptation are currently debated. Here, I show how prior work on wild butterflies casts a novel light on this topic. “Preference” is defined as the set of likelihoods of accepting particular resources after encountering them. So defined, butterfly oviposition preferences are heritable habitat adaptat...

The butterfly Euphydryas editha is known to be vulnerable to climate events that exacerbate natural phenological asynchrony between insect and hosts. In prior work, populations of E. editha have been more persistent at high latitudes and high elevations than in the south and at low elevations, consistent with response to observed warming climate. H...

The climate‐sensitive butterfly Euphydryas editha exhibited interpopulation variation in both phenology and egg placement, exposing individuals to diverse thermal environments. We measured ‘eggspace’ temperatures adjacent to natural egg clutches in populations distributed across a range of latitudes (36°8′–44°6′) and altitudes (213–3171 m). Eggs la...

Ecotypic variation among populations may become associated with widespread genomic differentiation, but theory predicts that this should happen only under particular conditions of gene flow, selection and population size. In closely related species, we might expect the strength of host-associated genomic differentiation (HAD) to be correlated with...

There is increasing pressure from policymakers for ecologists to generate more detailed ‘attribution’ analyses aimed at quantitatively estimating relative contributions of different driving forces, including anthropogenic climate change (ACC), to observed biological changes. Here, we argue that this approach is not productive for ecological studies...

Background Until recently the isolation of microsatellite markers from Lepidoptera has proved troublesome, expensive and time-consuming. Following on from a previous study of Edith's checkerspot butterfly, Euphydryas editha, we developed novel microsatellite markers for the vulnerable marsh fritillary butterfly, E. aurinia. Our goal was to optimize...

Map showing location of all 28 populations sampled in the present study. Populations 18 to 28 are those also included in a previous study using AFLP markers [28]. (TIF)

Sites across the UK and Catalonia region of Europe from which E. aurinia was sampled. (DOCX)

Spatial mosaics occur in both evolutionary and ecological properties of species' interactions. Studies of these patterns have facilitated description and prediction of evolutionary responses of interacting species to each other and to changing environments. We propose seeking complementary understanding of community assembly and dynamics by studyin...

Background/Question/Methods The study documents final outcomes of two independent anthropogenic host shifts by the North American butterfly Euphydryas editha. The first, at Schneider’s Meadow, Carson City, involved inclusion of an exotic, Plantago lanceolata, into the diet in addition to the traditional host, Collinsia parviflora. Field experiments...

The biological world is responding rapidly to a changing climate, but attempts to attribute individual impacts to rising greenhouse gases are ill-advised.

This thesis describes a relationship between maternal preference and offspring performance in a population of the butterfly Euphydryas editha that used two host plants, Pedicularis semibarbata and Collinsia torreyi from 1979 to 2001, but now no longer uses Collinsia. In the light of the known history of diet change in this butterfly population, it...

Changing climate can disrupt existing phenological relations between interacting species. We might expect the historical baseline for these effects to be precise synchrony between the season at which a consumer most requires food and the time when its resources are most available. When this is the case, change in any direction would be detrimental...

No sign of intrinsic incompatibilities in hybrids.Different-host hybrids showed no evidence of reduced fitness in terms of egg viability, early larval growth rate on a “neutral” host, adult female teneral weight, sex ratio, or fertility. Plots show means ± SEM for various cross types. Both same- and different-host hybrid families had mean egg viabi...

Early larval performance on Ctor (A) and Psem (B). Pure and same-host hybrid insects grew and survived at indistinguishable rates (compare C versus CC and P versus PP). Different-host hybrids grew slightly more slowly on Psem, although the effect was only significant when predators were excluded. Colored bars show LS means ± SEM. Numbers inside bar...

ANOVA tables from analyses of early larval performance on Psem . (0.08 MB PDF)

The modeled relationship between the discrimination phase (d-phase) and the probability of laying on a particular host. Positive and negative d-phases indicate a preference for Ctor and Psem, respectively. For a given d-phase, d (in days), the probability of laying on Ctor was modeled as (tan(4d)+1) / 2. It is biologically realistic that females wi...

ANOVA tables from analyses of early larval performance on Ctor . (0.08 MB PDF)

Adaptive surface shown in terms of absolute survival probability. This surface is the same as that shown in Figure 6 except that its height reflects expected offspring survival in absolute terms rather than expected offspring survival relative to that of the optimal clutch size on each respective host (see Methods). (1.80 MB PDF)

ANOVA tables from analyses of the effects of foraging height on mature Ctor . (0.09 MB PDF)

ANOVA tables from analyses of the effects of foraging height on Psem . (0.09 MB PDF)

ANOVA tables from analyses of the effects of oviposition behavior on offspring growth (A) and survival (B) on Ctor . (0.08 MB PDF)

ANOVA tables from analyses of the effects of oviposition site height on offspring development time on Psem . (0.08 MB PDF)

Oviposition site choice in a PC female butterfly. Females with one parent adapted to Psem and the other adapted to Ctor do not usually take the time to explore their host and instead lay their eggs at the point where they first contact an acceptable host plant. In this example, a PC female is placed at the top of Psem, immediately becomes intereste...

ANOVA tables from analyses of larval performance on mature Ctor . (0.08 MB PDF)

ANOVA tables from analyses of the effects of female clutch size on offspring growth (A) and survival (B) on Psem . (0.07 MB PDF)

Summary of six studies of EPI in insects that specialize on distinct host plants. (0.07 MB PDF)

Oviposition site choice in a PP female butterfly. Females whose parents were both adapted to Psem are very deliberate in choosing oviposition sites near the ground. In this typical example, a PP female is placed at the top of Psem and immediately dips her antennae towards the leaf, signaling her interest. Rather than laying right away, however, she...

Gene flow between populations that are adapting to distinct environments may be restricted if hybrids inherit maladaptive, intermediate phenotypes. This phenomenon, called extrinsic postzygotic isolation (EPI), is thought to play a critical role in the early stages of speciation. However, despite its intuitive appeal, we know surprisingly little ab...

Climate change alters phenological relations between interacting species. We might expect the historical baseline, or starting-point, for such effects to be precise synchrony between the season at which a consumer most requires food and the time when its resources are most available. We synthesize evidence that synchrony was not the historical cond...

Background/Question/Methods Changing climate can disrupt existing phenological relations between interacting species. We might expect the historical baseline for these effects to be precise synchrony between the season at which a consumer most requires food and the time when its resources are most available. When this is the case, change in any d...

Sample sizes and locations of the populations used for polymorphism screening. (0.10 MB DOC)

This file lists all the primers tested in the study, and the results of polymorphism testing based on a small sample of 8 individuals. Loci used for further analysis are highlighted in gray. (0.12 MB DOC)

The isolation of microsatellite markers remains laborious and expensive. For some taxa, such as Lepidoptera, development of microsatellite markers has been particularly difficult, as many markers appear to be located in repetitive DNA and have nearly identical flanking regions. We attempted to circumvent this problem by bioinformatic mining of micr...

Divergent adaptation to host plant species may be the major mechanism driving speciation and adaptive radiations in phytophagous insects. Host plants can differ intrinsically in a number of attributes, but the role of natural enemies in host plant specialization is often underappreciated. Here, we report behavioural divergence between the European...

When populations use different resources, they tend to diverge in traits that affect performance on those resources. The extent and complexity of divergence that is achieved will depend on gene flow, genetic constraints, and the character of divergent selection. We describe divergent host adaptation among Californian populations of the Melitaeine b...

The generic names Euptychia, Cissia and Argyreuplychia are reviewed in relation to the Cissia confusa species-group in Costa Rica and Trinidad, for which the generic name Cissia Doubleday is formally resurrected. Three new species of Cissia are described, C. confusa itself is resurrected from synonymy, three neotypes and three lectotypes are design...

1. Euphydryas editha (Boisduval) (Nymphalidae) butterflies exhibited no consistent pattern of change in egg weight over their lifetimes. This was the case for captive butterflies fed diets containing sugars and amino acids, for butterflies fed diets containing only sugar, and for those left completely at liberty in their natural habitats between ov...

1. The tendency of Euptychia libye butterflies to oviposit with greater likelihood on isolated plants than on members of clumps is evident in the field and can be duplicated under controlled conditions in the laboratory. 2. The causes and effects of this behaviour and the difficulties inherent in inferring preference from egg distributions in the f...

Because weevils are used as biocontrol agents against thistles, it is important to document and understand host shifts and the evolution of host-specificity in these insects. Furthermore, such host shifts are of fundamental interest to mechanisms of speciation. The mediterranean weevil Larinus cynarae normally parasitizes either one of two thistle...

It is difficult to imagine how warning colours evolve in unpalatable prey. Firstly, novel warningly coloured variants gain no protection from their colours, since predators have not previously encountered and learnt their colour patterns. This leads to a frequency-dependent disadvantage of a rare variant within a species. Secondly, novel warningly...

This chapter describes a small but unbiased sample of evolutionary changes: three evolutionary host shifts undertaken by a single butterfly species, Edith's checkerspot, Euphydryas editha. The first host shift is a straightforward and clearly anthropogenic event at Schneider's Meadow (Carson City, Nevada) triggered by the introduction of an exotic...

Abstract 1. Degree of host specialisation was a continuous variable in a population of Edith’s checkerspot butterfly (Euphydryas editha). A novel host, Collinsia torreyi, had been added to the diet in response to anthropogenic disturbance, and then abandoned prior to the current study. Butterflies either showed no preference or preferred their trad...

Natural selection acting on timing of metamorphosis can be sex-specific, resulting in differences in timing between males and females. Insects with discrete generations frequently show protandry: males usually mature before females. Both Euphydryas editha and E. aurinia butterflies followed this trend. The present study was motivated by the unusual...

Checkerspot butterflies have been used as an extraordinarily successful model system for more than four decades. This volume presents the first synthesis of the broad range of studies of that system as conducted in Ehrlich’s research group in Stanford, in Hanski’s research group in Helsinki and elsewhere. Ehrlich’s long - term research project on E...

Checkerspot butterflies have been used as an extraordinarily successful model system for more than four decades. This volume presents the first synthesis of the broad range of studies of that system as conducted in Ehrlich’s research group in Stanford, in Hanski’s research group in Helsinki and elsewhere. Ehrlich’s long - term research project on E...

Previous work on the Glanville fritillary butterfly (Melitaea cinxia) shows substantial inbreeding depression in both of our two study regions, Finland and southern France. The influence of inbreeding depression on population dynamics should depend on the strength of inbreeding avoidance. We conducted mate choice experiments to ascertain whether an...

Summary To predict evolution in plant-insect systems we can begin by defining potentially heritable traits of plants that describe how they interact with insects and potentially heritable traits of insects that describe how they interact with plants. Examples are "acceptability" as a plant trait and "preference" as an insect trait. Some practical a...

We investigate whether egg load (a surrogate for fecundity) drives host specificity in a herbivorous insect. In many insects, including our study organism (Edith's checkerspot butterfly), both egg load and tendency to accept low-ranked hosts increase during each search for an oviposition site. Effects on host acceptance of egg load and passage of t...

1. The butterfly Melitaea cinxia uses two host plant species in the Åland Islands of south-west Finland. Survey data show that host plant use is spatially variable and that the two species are not used (fed on by M. cinxia larvae) in proportion to their abundances. The pattern of host plant use by M. cinxia has been attributed in part to plant dist...

In experiments that investigate species' interactions, individuals are often chosen at random to represent their populations. However, this practice can generate misleading results when individuals of different species do not interact at random. We illustrate this effect by examining oviposition preferences of Euphydryas aurinia butterflies from th...

The effects of inbreeding on fitness and the maintenance of genetic load in metapopulations of the endangered Glanville fritillary butterfly (Melitaea cinxia) were examined in four laboratory experiments. In Finland M. cinxia occurs as a large metapopulation consisting of small local populations with fast turnover, whereas in southern France the sp...

Species living in highly fragmented landscapes often occur as metapopulations with frequent population turnover. Turnover rate is known to depend on ecological factors, such as population size and connectivity, but it may also be influenced by the phenotypic and genotypic composition of populations. The Glanville fritillary butterfly (Melitaea cinx...

Species living in highly fragmented landscapes often occur as metapopulations with frequent population turnover. Turnover rate is known to depend on ecological factors, such as population size and connectivity, but it may also be influenced by the phenotypic and genotypic composition of populations. The Glanville fritillary butterfly (Melitaea cinx...

We successfully trained adult Edith's checkerspot butterflies, Euphydryas editha, to modify their alighting preference for a novel flower and to reduce their handling time when searching for nectar in a flower that was difficult for them to use. Butterflies caught feeding from the difficult flower (manzanita, Arctostaphylos spp., Ericaceae) in the...

Laboratory and field studies on Melitaea cinxia in southwestern Finland showed a significant difference in the performance of experimental inbred and crossbred populations, which supports previous observational results on inbreeding-related extinctions in this species.

We studied host plant abundance, host use, and oviposition preference in metapopulations of the butterfly Melitaea cinxia within an area of 3500 km2 in the Aland islands, southwestern Finland. In the study area, M. cinxia has ;400 small local populations on dry meadows with the larval host plants, Plantago lanceolata and Veronica spicata. Plantago...

Ecologists and evolutionary biologists have a common interest in plant–insect interactions. Ecologists develop terminology describing patterns of association between plants and insects, while evolutionary biologists use the same words to denote potentially heritable traits of individuals. Use of the same terms to describe both traits of the interac...

Experiments designed to reveal variation among individual parasites in preference for different host species may generate misleading results. Apparent variation in the order of preference among host species can be generated solely from variation in the strength of discriminations made within host species. We illustrate this with a study of oviposit...

The population size of insects associated with any resource patch is determined by local birth and death in that patch and by migration into and out of the patch. When resource patches are small and close together, individuals move readily between patches, so very high emigration and immigration rates dominate patterns of local distribution (Kareiv...

Increasing emphasis is being placed on the large-scale and long-term dynamics of populations. A butterfly (Euphydryas editha) metapopulation that was naturally restricted to rocky outcrops in an area of coniferous forest suffered two major perturbations in 30 yr. First, humans clear-cut patches of forest in about 1967. The butterfly colonized the c...

We studied effects of spatiotemporally variable natural selection on local adaptation. A butterfly (Euphydras editha) metapopulation contained patches of two types: undisturbed outcrops and logged clearings (where the insects' traditional host had been killed). Butterflies retained their traditional host in the outcrops, but in the clearings they u...

A population of Edith's checkerspot butterfly,Euphydryas editha, laid eggs on two morphologically distinct host-plant species, one of which it had recently colonized. Search behaviour of two groups of females was compared in the field. ‘Naive’ females were experimentally deprived of both flight experience and host encounter, while ‘experienced’ fem...

A tree of mitochondrial DNA (mtDNA) haplotypes was constructed to estimate the number of evolutionary changes of host-plant preference needed to account for variation among 24 populations of the butterfly Euphydryas editha. Using 17 restriction endonucleases, 22 mtDNA haplotypes were found among 24 populations of this butterfly species. We allowed...

A tree of mitochondrial DNA (mtDNA) haplotypes was constructed to estimate the number of evolutionary changes of host-plant preference needed to account for variation among 24 populations of the butterfly Euphydryas editha. Using 17 restriction endonucleases, 22 mtDNA haplotypes were found among 24 populations of this butterfly species. We allowed...

The authors measured oviposition preferences and offspring performance (survival of young larvae) within Euphydryas editha. There was a significant association between speed of preference evolution and within-population diversity of preference: preference ranks were more diverse within populations where preference was evolving rapidly than within "...