Ancestral Host Research Articles

ABCB transporters in a leaf beetle respond to sequestered plant toxins.

Phytophagous insects can tolerate and detoxify toxic compounds present in their host plants and have evolved intricate adaptations to this end. Some insects even sequester the toxins for their defence. This necessitates specific mechanisms, especially carrier proteins that regulate uptake and transport to specific storage sites or protect sensitive tissues from noxious compounds. We identified three ATP-binding cassette subfamily B (ABCB) transporters from the transcriptome of the cardenolide-sequestering leaf beetle Chrysochus auratus and analysed their functional role in the sequestration process. These were heterologously expressed and tested for their ability to interact with various potential substrates: verapamil (standard ABCB substrate), the cardenolides digoxin (commonly used), cymarin (present in the species's host plant) and calotropin (present in the ancestral host plants). Verapamil stimulated all three ABCBs and each was activated by at least one cardenolide, however, they differed as to which they were activated by. While the expression of the most versatile transporter fits with a protective role in the blood-brain barrier, the one specific for cymarin shows an extreme abundance in the elytra, coinciding with the location of the defensive glands. Our data thus suggest a key role of ABCBs in the transport network needed for cardenolide sequestration.

Host Plant Choice Determined by Reproductive Interference between Closely Related Butterflies

AbstractA number of empirical studies have concluded that reproductive interference (RI) contributes to parapatric species distributions or sexual exclusion. However, the possibility that divergent host plant use in phytophagous insects is due to sexual exclusion has seldom been considered. Here, we present evidence that RI is responsible for different host plant use by two Pierid butterfly species, Pieris napi and Pieris melete. When a novel host species was introduced about 50 years ago, two Pierid butterfly species at first used both the ancestral host species and the novel one. Subsequently, P. napi shifted to use only the novel host, while P. melete shifted to specialize on the ancestral host. To explain these patterns, we investigated whether the two host species differ in suitability for larval growth and survival. Additionally, we tested whether RI occurred between the two butterfly species using large outdoor field cages. Courtship of females by conspecific and heterospecific males reduces the number of eggs laid by approximately half. However, RI is asymmetric and would generate selection on P. melete females to evolve to avoid the more suitable host species preferred by P. napi. Thus, our study suggests that sexual exclusion can explain the shift in host plant use by these two butterfly species.

Phylogenomics of the lepidopteran endoparasitoid wasp subfamily Rogadinae (Hymenoptera: Braconidae) and related subfamilies

AbstractRogadinae are a cosmopolitan, species‐rich braconid wasp subfamily whose species are endoparasitoids that attack larvae of a number of lepidopteran families. Members of this subfamily are characterized by pupating within the mummified host larval skin. The subfamily contains six tribes whose relationships have only been partially clarified: Aleiodini, Betylobraconini, Clinocentrini, Rogadini, Stiropiini and Yeliconini. The limits and composition of the closely related subfamilies to the Rogadinae, Hormiinae and Lysiterminae, also remain unclear. Here, we generated ultraconserved element data to reconstruct an almost fully resolved phylogeny for the members of Rogadinae and related subfamilies. Based on our best estimate of phylogeny, we confirm the monophyly of Rogadinae including Betylobraconini, synonymize Xenolobus Fahringer and Bequartia Cameron within the species‐rich genus Aleiodes Wesmael (syn.n.) based on DNA, and synonymize Promesocentrus van Achterberg with Pilichremylus Belokobylskij (syn.n.) based on morphology. We also consistently recovered Hormiinae and Lysiterminae as not reciprocally monophyletic, and thus propose to unite their members under Hormiinae. The ancestral host preference for Rogadinae was probably attacking concealed lepidopteran larvae, with the occurrence of at least two main subsequent transitions to attack both concealed and exposed hosts, one within Rogadini and a second within Aleiodini. We highlight the importance of natural history collections as a source for conducting genomic‐based studies using techniques that allow to obtain a substantial amount of data from considerably old preserved insect specimens.

The Mayan Tropical Rainforest: An Uncharted Reservoir of Tritrophic Host-Fruit Fly-Parasitoid Interactions

Simple SummaryTropical rainforest can provide various ecological services to adjacent agricultural environments, including maintaining and amplifying the numbers of beneficial insects. However, forest fragmentation and the selective cutting of indigenous trees used by native species of fruit flies and their parasitoids, degrades the potential of forests to provide ecological services to agriculture. Over a two–year period, we surveyed natural areas of the Mayan rainforest in Quintana Roo, Mexico. We found 11 species of native fruit flies belonging to the genus Anastrepha associated with 25 species of fruits belonging to ten plant families. We report the first records of 10 host plant species of the genus Anastrepha. We also report a new undescribed species of Anastrepha. The interaction between fruit flies and their parasitoids with host plants depends on fruit availability, which is crucial for the survival of each of these species. Our findings indicate that the areas of the Mayan rainforest surveyed represent a highly important reservoir for the diversity of native parasitoids spatially and temporally that are practically absent in fruits of cultivated plants. Conserving the landscape of the Mayan rainforest is important not only for species conservation, but also for the maintenance of fruit fly host plants of biological control agents in orchard agroecosystems in southeastern Mexico.Over a two–year period, we surveyed natural areas of the Mayan rainforest in Quintana Roo, Mexico. We found 11 species of Anastrepha Schiner (Diptera: Tephritidae) infesting 25 species of fruits belonging to ten plant families. We report the first records of 10 host plant species of the genus Anastrepha, which include the first report of a plant family (Putranjivaceae) serving as host of Anastrepha fraterculus (Wiedemann) infesting Drypetes lateriflora (Sw.) Krug and Urb. (Putranjivaceae). Pouteria reticulata (Engl.) Eyma (Sapotaceae) was found, for the first time, to be infested by Anastrepha serpentina (Wiedemann) and by a new undescribed species of Anastrepha. We also report Casimiroa microcarpa Lundell (Rutaceae) as a possible ancestral host for the Mexican fruit fly, Anastrepha ludens (Loew), in Central America. The family Sapotaceae was the best-represented host group with three fruit fly species recovered: A. serpentina, an economically-important species, found in eight host plants, and A. hamata and A. sp. (new species). We recorded six species of koinobiont parasitoids: Doryctobracon areolatus Szepligeti, Utetes (Bracanastrepha) anastrephae Viereck, Opius hirtus Fisher, and Doryctobracon zeteki Musebeck, (all Braconidae), and Aganaspis pelleranoi (Brethés) and Odontosema anastrephae Borgmeier, (both Figitidae). All these parasitoid species represent at least a new report for their host plants. Of the whole parasitoid community, D. areolatus was the most important parasitoid species with 52.7% of presence in 12 host plant species, parasitizing six fruit fly species. The interaction between fruit flies and their parasitoids with host plants depends on fruit availability, which is crucial for the survival of each of these species. Conserving the landscape of the Mayan rainforest is important not only for species conservation, but also for the maintenance of fruit fly host plants in orchard agroecosystems in southeastern Mexico.

Differential Tropism of SARS-CoV and SARS-CoV-2 in Bat Cells.

Severe acute respiratory syndrome coronavirus 2 did not replicate efficiently in 13 bat cell lines, whereas severe acute respiratory syndrome coronavirus replicated efficiently in kidney cells of its ancestral host, the Rhinolophus sinicus bat, suggesting different evolutionary origins. Structural modeling showed that RBD/RsACE2 binding may contribute to the differential cellular tropism.

Viral CpG Deficiency Provides No Evidence That Dogs Were Intermediate Hosts for SARS-CoV-2.

Due to the scope and impact of the COVID-19 pandemic there exists a strong desire to understand where the SARS-CoV-2 virus came from and how it jumped species boundaries to humans. Molecular evolutionary analyses can trace viral origins by establishing relatedness and divergence times of viruses and identifying past selective pressures. However, we must uphold rigorous standards of inference and interpretation on this topic because of the ramifications of being wrong. Here, we dispute the conclusions of Xia (2020. Extreme genomic CpG deficiency in SARS-CoV-2 and evasion of host antiviral defense. Mol Biol Evol. doi:10.1093/molbev/masa095) that dogs are a likely intermediate host of a SARS-CoV-2 ancestor. We highlight major flaws in Xia’s inference process and his analysis of CpG deficiencies, and conclude that there is no direct evidence for the role of dogs as intermediate hosts. Bats and pangolins currently have the greatest support as ancestral hosts of SARS-CoV-2, with the strong caveat that sampling of wildlife species for coronaviruses has been limited.

El SARS-CoV-2 y otros virus emergentes y su relación con la inocuidad en la cadena alimentaria

Influenza pandemics are related to the viral flow of wild and migratory birds, passing to pigs and poultry (intermediate hosts), which would end up infecting humans. Process management through Good Farming Practices for animal production and Animal Welfare should be important preventive measures to avoid this contagion. In coronaviruses and Ebola there are ancestral hosts (such as bats) and intermediate hosts (such as Civettictis civetta or Paradoxurus hermaphroditus for SARS-CoV-1, dromedaries for MERS-CoV, and snakes and Manis javanica for SARS-CoV-2). In its natural state the inter-specie jump would take much longer to occur; however, the existence of wet markets, where animals are traded, would have allowed viruses to find a new host in humans. In addition to pneumonia, the invasion of SARS-CoV-2 produces temporary discomfort, such as diarrhea, due to the abundance of the ACE-2 receptor in both lung cells and enterocytes, where its replication would increase the probability of transmission by the fecal-oral route. This risk could be minimized by improving Good Manufacturing Practice (GMP) procedures in the food industry. Future research should clarify the relationship between viruses and their hosts, as well as the effect of climate change and proteins availability for human consumption, on these pandemics.

Molecular phylogenetic analyses reveal a deep dichotomy in the conifer-inhabiting genus Trisetacus (Eriophyoidea: Nalepellidae), with the two lineages differing in their female genital morphology and host associations.

We analyzed the phylogenetic relationships of the genus Trisetacus using two genes [cytochrome c oxidase subunit I (COI) and D1-D2 region of 28S rDNA (D1-D2 28S)], a representive taxon sampling (nearly 40% of known diversity), and a large set of close and distant outgroups. Our analyses suggest the presence of a dichotomy between Trisetacus associated with Cupressaceae and Pinaceae. The following smaller molecular clades were found: Pin-1 (bud mites, twig sheath mites, bark gall mites, and endoparasitic mites from pinaceans), Pin-2 (needle sheath mites from pines), Pin-2a (putative Nearctic group of needle sheath mites), Pin-2b (putative Palearctic group of needle sheath mites), Cup-1 and 2 (bud, cone, seed mites and mites living under bark scales from cupressaceans). The monophyly of the recently proposed subgenus Brevithecus nested within clade Cup-2 was confirmed. Ancestral character reconstruction analyses recovered: (1) Pinaceae as the ancestral hosts of Nalepellidae and Trisetacus, (2) repetitive reductions of the spermathecal tube independently occurred in two lineages of Trisetacus from Cupressaceae, and (3) several mite habitats on host (galls, cones, twig sheaths, seeds, inside leaves, and under scales) are evolutionarily derived states, whereas living in buds or needle sheaths are ancestral states for Trisetacus clades Cup and Pin. Using confocal microscopy, we identified six basic types of the female internal genitalia of Trisetacus based on shapes of the spermatheca and spermathecal tube. These genitalic types are strongly correlated with lineages recovered by molecular phylogenetic analyses, suggesting that the female genital morphology is both evolutionarily conserved and is a factor influencing macroevolutionary patterns in this group of mites.

Host Range Expansion of an Endemic Insect Herbivore is Associated With High Nitrogen and Low Fibre Content in Exotic Pasture Plants.

Endemic moth species of the genus Wiseana spp. (Hepialidae) have become serious pests of introduced pasture plants in New Zealand. The original native host plants of these moths have not been confirmed. This study investigated the performance (survival, development time, weight gain) of three Wiseana species on seven putative host plants: five native and two exotic species. The aim was to identify native hosts for the three Wiseana species and to compare their performance on native plants and exotic pasture plants. The chemical composition of the seven putative host plants was investigated to compare native and exotic plant chemistries, and to test for associations between plant characteristics and performance of selected Wiseana species. Carbon, nitrogen, silica and fibre contents were measured for each plant species; primary metabolite composition was determined by gas chromatography-mass spectrometry. For the three moth species, increased survival and weight gain were significantly associated with high nitrogen and low fibre contents in one exotic host plant, white clover (Trifolium repens), although one species, W. umbraculata, did not complete development to adult on any of the plants tested, including clover. Two exotic plants (T. repens, Lolium perenne × Lolium multiflorum), and two native plants (Aciphylla squarrosa and Festuca actae) supported W. copularis development to the adult stage, but only one exotic (T. repens) and one native (F. actae) species supported complete development of W. cervinata. Exotic and native plant species had distinct metabolite profiles, but there was no significant association between metabolite composition and Wiseana performance. We conclude that W. copularis and W. cervinata, but not W. umbraculata, have expanded their host range, because of their ability to use both native and new hosts. No evidence was found for a host shift, i.e., a loss of performance on the ancestral host compared with the new host.

Evolutionary Modification of Pereopods in Phronimid Amphipods (Crustacea: Amphipoda: Hyperiidea: Phronimidae) Reflects Host Differences.

Phronimid amphipods are oceanic crustaceans associated with gelatinous zooplankters. Their host organisms belong mainly to two taxonomic groups: tunicates (salps or pyrosomes; subphylum Tunicata) and siphonophores (Cnidaria). After these amphipods devour the inner tissues of their hosts, they display the unique behavior of modifying their hosts into hollow barrel-shaped shelters, which are then utilized as neonatal nurseries by the females. Although previous studies have revealed the host specificity of these amphipods, it has not been inferred which types of hosts ancestral phronimids could have originally used. Moreover, morphological changes associated with host switching have not yet been studied. To deduce the evolutionary patterns of host switching, we investigated the phylogenetic relationships of phronimid species by using two genes: (1) cytochrome c oxidase subunit I (COI) and (2) 18S ribosomal RNA (18S). In addition, a morphometric analysis was conducted in order to better understand the morphological relationships between phronimids and their host organisms. Our phylogenetic analysis suggests that the ancestral host animals of phronimids could have been tunicates and that the host organisms have independently switched from tunicates to siphonophores at least twice in the family Phronimidae. Our morphometric analysis revealed that phronimids using siphonophores as hosts have a relatively shorter pereopod 5 compared to those using tunicates. The shortening of pereopod 5 seems to be an adaptation to the narrower internal space of siphonophore barrels compared to those of tunicates.

Host Plant and Antibiotic Effects on Scent Bouquet Composition of Anastrepha ludens and Anastrepha obliqua Calling Males, Two Polyphagous Tephritid Pests.

In insects, the quality of sex pheromones plays a critical role in mating success and can be determined by the ability of larvae/adults to accrue chemical precursors. We tested the host-quality-effect hypothesis by analyzing the chemical composition of scent bouquets emitted by calling males of two polyphagous tephritid species (Anastrepha ludens and A. obliqua) that originated from 13 fruit species representing diverse plant families. In A. ludens, we worked with an ancestral host (Rutaceae), nine exotic ones (Rutaceae, Anacardiaceae, Rosaceae, Solanaceae, Lythraceae), and two species never attacked in nature but that represent candidates for host-range expansion (Solanaceae, Myrtaceae). In A. obliqua, we tested an ancestral, a native, and an exotic host (Anacardiaceae), one occasional (Myrtaceae), and one fruit never attacked in nature (Solanaceae). We identified a core scent bouquet and significant variation in the bouquet’s composition depending on the fruit the larvae developed in. We also tested the possible microbial role on the scent bouquet by treating adults with antibiotics, finding a significant effect on quantity but not composition. We dwell on plasticity to partially explain our results and discuss the influence hosts could have on male competitiveness driven by variations in scent bouquet composition and how this could impact insect sterile technique programs.

Feeding, Survival, and Fecundity of Adult Emerald Ash Borer (Coleoptera: Buprestidae) on Foliage of Two Novel Hosts and Implications for Host Range Expansion.

Insect herbivores are more likely to successfully use a novel host if the plant is closely related to the ancestral host and the insect is polyphagous. Emerald ash borer (EAB), Agrilus planipennis (Fairmaire) (Coleoptera: Buprestidae), is a specialist wood borer of ash (Fraxinus spp., Lamiales: Oleaceae) trees and one of the most destructive forest pests in North American forests. Recent studies have found that larvae can develop in stems of two ash relatives; white fringetree (Chionanthus virginicus (L.) [Laminales: Oleaceae]) and cultivated olive (Olea europaea (L.) [Laminales: Oleaceae]). For EAB adults, the ability to consume, successfully mate, and lay viable eggs on foliage of these hosts is unknown. Thus, we conducted two no-choice assays with adult EAB on foliage of white fringetree and olive paired with positive controls of susceptible ash. Larval performance was also examined in a reciprocal study with cut stems of white fringetree and green ash (Fraxinus pennsylvanica Marshall) to determine whether adult diet impacted the success of progeny. Longevity, consumption rates, and fecundity of adults were similar on white fringetree and ash foliage. In contrast, adults consuming olive died quickly, consumed more over time, and females laid far fewer eggs compared to those on ash. Adult diet did not impact larval success, but larvae in white fringetree stems grew slower. These results indicate that white fringetree is a suitable host for EAB to complete its lifecycle, although larvae perform more poorly on this host than in susceptible ash species. In contrast, the more distantly related olive appears to be a poor host for adult EAB, although some viable eggs were produced by females.

Host preference and sorus location correlate with parasite phylogeny in the smut fungal genus Microbotryum (Basidiomycota, Microbotryales)

The smut fungal genus Microbotryum (Microbotryales, Pucciniomycotina) contains species that parasitize plants from many different lineages of euasterids, with host specificity of individual parasite species in general being exceptionally high. Additionally, it has been shown that the location of spore production in some species is related to spore dispersal. In this phylogenetic study based on ITS and LSU rDNA data of 57 Microbotryum spp., host spectra and sorus location are mapped on the phylogeny of Microbotryum species in order to understand the macroevolutionary patterns of these two traits. We find that monophyletic parasite clades correspond well with monophyletic host clades and also that monophyletic parasite groups in general produce their spores in the same plant organ. Ancestral state reconstruction inferred the most probable ancestral trait for sorus location being leaves and the most probable ancestral host family for the genus Microbotryum as being the Polygonaceae. According to molecular analyses, a newly sequenced specimen of Ustilago ducellieri, a seed parasite on Arenaria leptoclados, previously treated as synonym of Microbotryum duriaeanum, belongs to a lineage distinct from specimens of M. duriaeanum. A new combination, Microbotryum ducellieri, is accordingly proposed. Taxonomic implications of the presented analyses for the genera Bauhinus and Haradaea are briefly discussed.

Bayesian Inference of Ancestral Host-Parasite Interactions under a Phylogenetic Model of Host Repertoire Evolution.

Intimate ecological interactions, such as those between parasites and their hosts, may persist over long time spans, coupling the evolutionary histories of the lineages involved. Most methods that reconstruct the coevolutionary history of such interactions make the simplifying assumption that parasites have a single host. Many methods also focus on congruence between host and parasite phylogenies, using cospeciation as the null model. However, there is an increasing body of evidence suggesting that the host ranges of parasites are more complex: that host ranges often include more than one host and evolve via gains and losses of hosts rather than through cospeciation alone. Here, we develop a Bayesian approach for inferring coevolutionary history based on a model accommodating these complexities. Specifically, a parasite is assumed to have a host repertoire, which includes both potential hosts and one or more actual hosts. Over time, potential hosts can be added or lost, and potential hosts can develop into actual hosts or vice versa. Thus, host colonization is modeled as a two-step process that may potentially be influenced by host relatedness. We first explore the statistical behavior of our model by simulating evolution of host–parasite interactions under a range of parameter values. We then use our approach, implemented in the program RevBayes, to infer the coevolutionary history between 34 Nymphalini butterfly species and 25 angiosperm families. Our analysis suggests that host relatedness among angiosperm families influences how easily Nymphalini lineages gain new hosts. [Ancestral hosts; coevolution; herbivorous insects; probabilistic modeling.]

Phylodynamics of sunflower chlorotic mottle virus, an emerging pathosystem

Distribution and epidemiological patterns of sunflower chlorotic mottle virus (SCMoV) in sunflower (Helianthus annuus L.) growing areas in Argentina were studied from 2006 to 2017. The virus was detected exclusively in the Pampas region (Entre Ríos, Santa Fe, Córdoba, La Pampa and Buenos Aires provinces). Phylodynamic analyses performed using the coat protein gene of SCMoV isolates from sunflower and weeds dated the most recent common ancestor (MRCA) back to 1887 (HPD95% = 1572–1971), which coincides with the dates of sunflower introduction in Argentina. The MRCA was located in the south of Buenos Aires province and was associated with sunflower host (posterior probability for the ancestral host, ppah = 0.98). The Bayesian phylodynamic analyses revealed the dispersal patterns of SCMoV, suggesting a link between natural host diversity, crop displacement by human activities and virus spread.

Varroa mite evolution: a neglected aspect of worldwide bee collapses?

While ectoparasitic Varroa mites cause minimal damage to their co-evolved ancestral host, the eastern honey bee (Apis cerana), they devastate their novel host, the western honey bee (Apis mellifera). Over several decades, the host switch caused worldwide population collapses, threatening global food security. Varroa management strategies have focused on breeding bees for tolerance. But, can Varroa overcome these counter-adaptations in a classic coevolutionary arms race? Despite increasing evidence for Varroa genetic diversity and evolvability, this eventuality has largely been neglected. We therefore suggest a more holistic paradigm for studying this host-parasite interaction, one in which 'Varroa-tolerant' bee traits should be viewed as a shared phenotype resulting from Varroa and honey bee interaction.

Molecular genotyping and epidemiology of equine piroplasmids in South Africa

Recently reported substantial genetic diversity within Theileria equi 18S rRNA gene sequences has led to the identification of five genotypes A, B, C, D, and E, complicating molecular and serological diagnosis. In addition, T. haneyi has lately been reported as a species closely related to the T. equi 18S rRNA genotype C (Knowles et al., 2018). Theileria spp. of this group have a monophyletic origin and are therefore referred to as Equus group to distinguish them from the remaining Theileria lineages (Jalovecka et al., 2019). In this study, we report on the development of genotype-specific quantitative real-time PCR assays capable of detecting and distinguishing between each parasite genotype. Alignment of complete 18S rRNA sequences available on GenBank allowed for the design of a single primer pair and five TaqMan minor groove binder (MGB™) probes specific for each genotype (A–E). The assays, evaluated as qPCR simplex and two qPCR multiplex formats (Multiplex EP–ABC and Multiplex EP–DE), were shown to be both efficient and specific in the detection of T. equi genotypes. The developed qPCR assays were used to study (i) the intra-specific diversity of parasite genotypes within horse and zebra, (ii) the inter-specific differences in parasite genotype diversity in horses as compared to zebra, and (iii) the geographic distribution of T. equi 18S rRNA genotypes in South Africa. In addition, (iv) the presence of T. haneyi in South Africa was evaluated. An assessment of 342 equine field samples comprising 149 field horses, 55 racehorses, and 138 wild zebra confirmed the previously reported presence of T. equi 18S rRNA genotypes A, B, C, and D, and absence of genotype E in South African equids. Theileria equi genotypes A, B, C, and D, were detected in zebra, whereas only genotypes A, C and D, could be identified in field horses, and only genotypes A and C in racehorses. Genotypes B and D were the dominant genotypes identified in zebra in South Africa, while horses were predominantly infected with T. equi genotypes A and C. The greater diversity of T. equi genotypes in zebra suggests that it is an ancestral host for this piroplasmid lineage. Importantly, evidence is presented that each identified T. equi genotype segregates independently in each of the three studied equid populations reinforcing the notion that they represent individual separate entities corresponding to species. Preliminary investigations of the relationship between T. equi genotype C infections and Theileria haneyi, suggest that in addition to the five currently known T. equi genotypes, South African equids are also infected with T. haneyi.

Extensive host-switching of avian feather lice following the Cretaceous-Paleogene mass extinction event

Nearly all lineages of birds host parasitic feather lice. Based on recent phylogenomic studies, the three major lineages of modern birds diverged from each other before the Cretaceous-Paleogene (K-Pg) mass extinction event. In contrast, studies of the phylogeny of feather lice on birds, indicate that these parasites diversified largely after this event. However, these studies were unable to reconstruct the ancestral avian host lineage for feather lice. Here we use genome sequences of a broad diversity of lice to reconstruct a phylogeny based on 1,075 genes. By comparing this louse evolutionary tree to the avian host tree, we show that feather lice began diversifying on the common ancestor of waterfowl and landfowl, then radiated onto other avian lineages by extensive host-switching. Dating analyses and cophylogenetic comparisons revealed that two of three lineages of birds that diverged before the K-Pg boundary acquired their feather lice after this event via host-switching.

Biogeography and early diversification of Tapinotaspidini oil-bees support presence of Paleocene savannas in South America

Worldwide distributed tropical savannas were established only in the Miocene, with climatic cooling and rise of C4 grasses. However, there is evidence for an earlier presence of savanna-like vegetation in southern parts of South America. Here we investigated the biogeographic history of a clade of solitary bees which have endemic groups in areas covered by savannas and other types of open vegetation as well as forested areas. We hypothesized that these bees originated in savanna-like biomes and that shifts to forested areas and floral host shifts increased species diversification along their evolutionary history. We reconstructed a comprehensive phylogeny for Tapinotaspidini bees based majorly on original DNA sequences. We then used macroevolutionary tools to estimate ancestral range area and reconstructed ancestral habitat (open versus forested) and host plant association to analyze the effects of shifts in vegetation type and flower hosts on their diversification. Tapinotaspidini bees originated in the Paleocene and in a savanna-type, Cerrado-like, which is reinforced by reconstruction of open vegetation as the most probable ancestral area, thus bringing additional evidence to a much earlier origin of this vegetation type in South America. Shifts to forested areas occurred at least three times in a period of 30 Ma and were responsible for slight increases in diversification rates. Malpighiaceae is the ancestral floral host; host broadening occurred only in the Miocene and at least in three occasions. Host shifts, i.e. from Malpighiaceae to other oil families, occurred in the Eocene and Miocene. Both host broadening and host shifts did not significantly alter diversification rates, however exploitation of other oil sources were important in occupying new habitats. The link between biomes and host plant shifts and changes in diversification rate brings us additional insights into the evolution of bees and associated flora in South America.

Molecular phylogeny and host‐plant use (Lamiaceae) of theThymogethespollen beetles (Coleoptera)

AbstractThe 24 members of the Euro‐Asiatic genusThymogethesare highly specialized pollen beetles associated as larvae with flowers of Lamiaceae Nepetoideae. All members of the genus were analysed in within the framework of an integrative taxonomy approach, which was aimed to reconstruct the phylogenetic relationships, as well as the possible pattern of evolution of their larval‐host‐plant association. Evidence from multiple molecular markers [COI; 16S; H3], combined with an estimation of divergence times using an average rate of 0.0177 substitutions/site/My among branches, placed the origin of the genus at a minimum of 9–10 Mya. This date of origin approximates the known evolution of the host plants in Euro‐Mediterranean areas. Evidence from combined molecular and cladistic morphological analyses resulted in suitable agreement with the previously established morphology‐based systematics of the genus, although members of theexilisspecies‐group were split into three clades. The only disagreement between results of this new combined phylogeny and previous classification is in the exclusion of “Thymogethes”grenieri. This species is herein positioned outside the genus, based on molecular evidence. Our analysis depicts severalThymogethesspecies differentiating in the last few Mys, specifically those included in theT. lugubrisspecies‐group. Combined evidence from DNA, morphology and ancestral state parsimony reconstruction of larval‐host‐plant associations suggests that subtribe Menthinae likely represents the ancestral host plants, with a series of independent host shifts during the radiation of the clade, in association first with Menthinae and subsequently with Lavandulinae and Nepetinae. Steno‐oligophagy is the most frequent (86%) condition, while strictly monophagous species are less numerous (14%).