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Plants and herbivorous insects have dominated terrestrial ecosystems for over 300 million years. Uniquely in the fossil record, foliage with well-preserved insect damage offers abundant and diverse information both about producers and about ecological and sometimes taxonomic groups of consumers. These data are ideally suited to investigate food web response to environmental perturbations, and they represent an invaluable deep-time complement to neoecological studies of global change. Correlations between feeding diversity and temperature, between herbivory and leaf traits that are modulated by climate, and between insect diversity and plant diversity can all be investigated in deep time. To illustrate, I emphasize recent work on the time interval from the latest Cretaceous through the middle Eocene (67-47 million years ago (Ma)), including two significant events that affected life: the end-Cretaceous mass extinction (65.5 Ma) and its ensuing recovery; and globally warming temperatures across the Paleocene-Eocene boundary (55.8 Ma). Climatic effects predicted from neoecology generally hold true in these deep-time settings. Rising temperature is associated with increased herbivory in multiple studies, a result with major predictive importance for current global warming. Diverse floras are usually associated with diverse insect damage; however, recovery from the end-Cretaceous extinction reveals uncorrelated plant and insect diversity as food webs rebuilt chaotically from a drastically simplified state. Calibration studies from living forests are needed to improve interpretation of the fossil data.

Vascular-plant hosts, their arthropod herbivores, and associated functional feeding groups are distributed spatiotemporally into four major herbivore expansions during the past 420 m.y. They are: (1) a Late Silurian to Late Devonian (60 m.y.) phase of myriapod and apterygote, hexapod (perhaps pterygote) herbivores on several clades of primitive vascular-plant hosts and a prototaxalean fungus; (2) a Late Mississippian to end-Permian (85 m.y.) phase of mites and apterygote and basal pterygote herbivores on pteridophyte and basal gymnospermous plant hosts; (3) a Middle Triassic to Recent (245 m.y.) phase of mites, orthopteroids (in the broadest sense) and hemipteroid and basal holometabolan herbivores on pteridophyte and gymnospermous plant hosts; and (4) a mid Early Cretaceous to Recent (115 m.y.) phase of modern-aspect orthopteroids and derived hemipteroid and holometabolous herbivores on angiospermous plant hosts. These host-plant and herbivore associations are mediated by seven functional feeding groups: a) external foliage feeding, b) piercing-and-sucking, c) boring (Phase 1 origins); d) galling, e) seed predation, f) nonfeeding oviposition (Phase 2 origins); and leaf mining (early Phase 3 origin). Within about 20 m.y. of each herbivore expansion, there is a biota that expresses the nearly full spectrum of later plant-arthropod associations. These four associational phases may be linked to the paleoclimatologic variables of greenhouse/icehouse cycles and atmospheric O2 and CO2 levels by uncertain causes, although some relationship probably is present. The 7 functional feeding groups persist through most of the sampled interval but harbor host-plants and arthropod herbivores that are spatiotemporally ephemeral. Poor understanding of associations in Phases 1 to 3 is attributed to disproportionate focus on the angiosperm and holometabolan insect associations of Phase 4.

In this study, we investigated whether patterns of gene expression in larvae feeding on different plants can explain important aspects of the evolution of insect–plant associations, such as phylogenetic conservatism of host use and re-colonization of ancestral hosts that have been lost from the host repertoire. To this end, we performed a phylogenetically informed study comparing the transcriptomes of 4 nymphalid butterfly species in Polygonia and the closely related genus Nymphalis. Larvae were reared on Urtica dioica, Salix spp., and Ribes spp. Plant-specific gene expression was found to be similar across butterfly species, even in the case of host plants that are no longer used by two of the butterfly species. These results suggest that plant-specific transcriptomes can be robust over evolutionary time. We propose that adaptations to particular larval food plants can profitably be understood as an evolved set of modules of co-expressed genes, promoting conservatism in host use and facilitating re-colonization. Moreover, we speculate that the degree of overlap between plant-specific transcriptomes may correlate with the strength of trade-offs between plants as resources and hence to the probability of colonizing hosts and complete host shifts.

Using molecular phylogenetic data and methods we inferred divergence times and diversification patterns for the weevil subfamily Ceutorhynchinae in the context of host-plant associations and global climate over evolutionary time. We detected four major diversification shifts that correlate with both host shifts and major climate events. Ceutorhynchinae experienced an increase in diversification rate at ~53 Ma, during the Early Eocene Climate Optimum, coincident with a host shift to Lamiaceae. A second major diversification phase occurred at the end of the Eocene (~34 Ma). This contrasts with the overall deterioration in climate equability at the Eocene-Oligocene boundary, but tracks the diversification of important host plant clades in temperate (higher) latitudes, leading to increased diversification rates in the weevil clades infesting temperate hosts. A third major phase of diversification is correlated with the rising temperatures of the Late Oligocene Warming Event (~26.5 Ma); diversification rates then declined shortly after the Middle Miocene Climate Transition (~14.9 Ma). Our results indicate that biotic and abiotic factors together explain the evolution of Ceutorhynchinae better than each of these drivers viewed in isolation.

We studied host plant abundance, host use, and oviposition preference in metapopulations of the butterfly Melitaea cinxia within an area of 3500 km2in 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 lanceolata occurs in practically all meadows otherwise suitable for the butterfly, whereas the distribution of V. spicata is largely restricted to the northwestern part of the study area. Based on observations of 6500 prediapause larval groups during 1993-1996, we document spatial variation in host plant use in relation to their abundance (electivity). The fraction of larval groups found on V. spicata increased disproportionally with the relative cover of V. spicata in the habitat patches. Additionally, the probability of Veronica use in a population increased with increasing number of larval groups found on Veronica in the surrounding populations but decreased with increasing use of Plantago in the neighborhood. This regional effect on host use at the scale of migrating butterflies could be caused either by spatial variation in the insect (in either preference or performance) or by spatial variation in plants (in resistance to attack by the butterflies). To study the first possibility, we conducted oviposition preference experiments using butterflies from five metapopulations located 2-55 km from each other and characterized by differences in host plant availability and host use. We found clear genetic differences in oviposition preference between the five metapopulations consistent with the observed host use patterns in the field. We conclude that the spatial host use patterns of M. cinxia in the study area are driven both by direct effects of local host abundance and by indirect effects mediated through metapopulation-level adaptation to the regionally more abundant host plant.

• Insect fluid-feeding on fossil vascular plants is an inconspicuous and underappreciated mode of herbivory that can provide novel data on the evolution of deep-time ecological associations and indicate the host-plant preferences of ancient insect herbivores. Previous fossil studies have documented piercing-and-sucking herbivory but often are unable to identify culprit insect taxa. • One line of evidence are punctures and scale-insect impression marks made by piercing-and-sucking insects that occasionally provide clues to the systematic identities and relationships of particular insect herbivores. • We report here the earliest occurrences of piercing and sucking on early angiosperms as evidenced by scale insect covers, impression marks, punctures and body fossils – notably a mealybug – from the Lower Cretaceous Rose Creek Flora of the Dakota Formation (c. 103 Ma), in southeastern Nebraska, USA. The mealybug, two other scale insect taxa, and several distinctive damage types on laurel leaves and seed-plant stems at Rose Creek document a diverse guild of piercing-and-sucking insects on early angiosperms. • The discovery of an Early Cretaceous female mealybug indicates an early herbivorous association with a laurel host. These data provide direct evidence for co-associations and possible coevolution of scale insects and their plant hosts during early angiosperm diversification.

1. The desert locust is a migratory pest whose population development in remote areas must be monitored to prevent outbreaks, upsurges and plagues. Monitoring would be very much facilitated if the area of search could be restricted to sites of likely population increase. 2. The spatial distribution of solitarious desert locusts on the Red Sea coastal plain of Sudan was determined over 3 years from November to March. Additional observations were made on habitat factors, such as plant community, soil texture, soil moisture and land use. 3. Locust densities varied according to the amount and distribution of rainfall and longevity of the annual green vegetation, with virtually no locusts being observed in the driest season. 4. Samples on a grid of 120 sites within a 120-km stretch of coastal plain showed that locusts were prevalent only in the millet-Heliotropium plant community, which is found at sites with a fine sandy soil texture and comparatively high and long-lasting soil moisture in wadi deltas. These sites constitute less than 5% of the area of this part of the plain. 5. An accessory study showed association between desert locust distribution and millet cropping in an area where no Heliotropium was found. Other samples confirmed the association between solitarious desert locust and millet agriculture. 6. Synthesis and applications. The results indicate that surveys for early detection and control of desert locust on the Red Sea coast of Sudan can focus on millet cropping areas. The results suggest that the efficiency of monitoring migratory pest outbreaks in remote areas could be enhanced by using associations between plant communities and herbivorous insects to predict risk areas and target survey efforts.

Part of the exoskeleton of some wood-inhabiting insects is modified to form a mycangium, which is a specialized organ used to convey fungal spores or yeasts to their offspring. Although most stag beetles (Coleoptera: Lucanidae) are known to have female-specific mycangia and associated yeast symbionts, the evolutionary origin of the mycangium in this group remains unresolved. Here, we report the presence of a mycangium and associated yeast symbionts in the European horned stag beetle Sinodendron cylindricum (L.), which belongs to an ancestral clade of the Lucanidae. The mycangium of S. cylindricum is shown to be female-specific and have the same developmental origin as that of other stag beetles. A total of five yeast strains were isolated from adult mycangia and larval gut of S. cylindricum. Of these, we suggest that SICYAM1 is an undescribed yeast with taxonomic novelty, and have identified SICYLG3 as the xylose-fermenting yeast Scheffersomyces insectosa using nuclear ribosomal RNA and ITS sequences. The remaining three yeast strains, SICYAM2, SICYLG1, and SICYLG2, were assigned to the genus Sugiyamaella. Yeast density in the adult mycangium was lower than that of the more evolutionarily advanced stag beetles, the European Lucanus cervus (L.) and Dorcus parallelipipedus (L.), which were also examined in this study. No living yeasts were isolated from the adult guts. However, a third instar larva of S. cylindricum harbored 104–106 living yeasts in each gut region, which suggests that gut yeasts play an important role in these wood-feeding larvae.

An engraving made by a scolytid bark beetle, assigned to the genus Dendroctonus of the tribe Tomicini, has been identified on a mummified, middle Eocene (45 Ma) specimen of Larix altoborealis wood from the Canadian High Arctic. Larix altoborealis is the earliest known species of Larix, a distinctive lineage of pinaceous conifers that is taxonomically identifiable by the middle Eocene and achieved a broad continental distribution in northern North America and Eurasia during the late Cenozoic. Dendroctonus currently consists of three highly host-specific lineages that have pinaceous hosts: a basal monospecific clade on Pinoideae (Pinus) and two sister clades that consist of a speciose clade associated exclusively with Pinoideae and six species that breed overwhelmingly in Piceoideae (Picea) and Laricoideae (Pseudotsuga and Larix). The middle Eocene engraving in L. altoborealis represents an early member of Dendroctonus that is ancestral to other congeneric species that colonized a short-bracted species of Larix. This fossil occurrence, buttressed by recent data on the phylogeny of Pinaceae subfamilies and Dendroctonus species, indicates that there was phylogenetically congruent colonization by these bark-beetle lineages of a Pinoideae + (Piceoideae + Laricoideae) host-plant sequence. Based on all available evidence, an hypothesis of a geochronologically early invasion during the Early Cretaceous is supported over an alternative view of late Cenozoic cladogenesis by bark beetles onto the Pinaceae. These data also suggest that host-plant chemistry may be an effective species barrier to colonization by some bark-beetle taxa over geologically long time scales.

The yellow-necked dry-wood termite Kalotermes flavicollis (Fabricius, 1793) (Blattodea: Kalotermitidae) is an important pest that infests wood in Europe. An increase in attacks by K. flavicollis has been seen in buildings and in churches in Palermo (Italy), with attacks on both structural elements and artefacts. Future climate changes are expected to lead to increasing temperatures, which will probably affect the pest status of this species, which is difficult to control. In this context, it is important to identify potential natural antagonists of K. flavicollis. During a survey of the K. flavicollis population, several dead individuals with evident fungal efflorescence were found. Therefore, a study aimed at the isolation and identification of these microorganisms was conducted. Fungal colonies isolated from mycelial structures grown on insects were identified based on morphological characteristics and DNA profiling. Three different species were identified: Aspergillus nomius, A. subramanianii, and A. tamarii. This is the first time that fungi have been recorded in association with the yellow-necked dry-wood termite and, in addition, this study reports the first association of A. subramanianii with Isoptera.