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Figure 16 in Geographic variation in host selection in the spider wasps Entypus unifasciatus (Say) and Tachypompilus ferrugineus (Say) (Hymenoptera: Pompilidae), II

Figure 16. Combined geographic occurrence records of host species of Agelenidae, Ctenidae, Lycosidae, Pisauridae, Selenopidae, Sparassidae, Trechaleidae and Zoropsidae for Entypus unifasciatus and Tachypompilus ferrugineus based on 19,097 SCAN and GBIF collection records and online images. Northwestern Mexico is poorly sampled on this map. Note scarcity of records from the Pacific Northwest.

Figure 15 in Geographic variation in host selection in the spider wasps Entypus unifasciatus (Say) and Tachypompilus ferrugineus (Say) (Hymenoptera: Pompilidae), II

Figure 15. Geographic occurrence records of host species of Agelenidae, Ctenidae, Lycosidae, Pisauridae, Selenopidae, Sparassidae, Trechaleidae and Zoropsidae for Entypus unifasciatus and Tachypompilus ferrugineus based on 19,097 SCAN and GBIF collection records and online images. Northwestern Mexico is poorly sampled on this map. Note scarcity of records from the Pacific Northwest.

Figures 1–6. Entypus unifasciatus. 1 in Geographic variation in host selection in the spider wasps Entypus unifasciatus (Say) and Tachypompilus ferrugineus (Say) (Hymenoptera: Pompilidae), II

Figures 1–6. Entypus unifasciatus. 1) Entypus unifasciatus unifasciatus (Say), female, with immobilized Dolomedes tenebrosus Hentz (Pisauridae), adult female, Caitlin Dunbar Nature Center, Lichester, Howard County, MD. © Photographer unknown. 2) Entypus unifasciatus cressoni (Banks), female, attempting to re-sting immobilized Dolomedes tenebrosus, adult female, Lewisville, Denton County, TX. Photograph © Ruthanne Thomas. 3) Entypus unifasciatus cressoni, female, with immobilized Rabidosa rabida (Walckenaer) (Lycosidae), adult female, Cook's Slough Nature Park, Uvalde County, TX. Photograph © Tripp Davenport. 4) Entypus unifasciatus californicus (Townes), female, with immobilized Schizocosa mccooki (Montgomery) (Lycosidae), adult female, San Luis Obispo, San Luis Obispo County, CA. Photograph © Morgan Cantrell. 5) Entypus unifasciatus cressoni, female, with immobilized Ctenus sp. (Ctenidae), adult or subadult female, Tequila, Jalisco State, Mexico. Photograph © Chris Lloyd. 6) Entypus unifasciatus urichi (Banks), female, with immobilized Ancylometes bogotensis (Keyserling) (Ctenidae), subadult female or juvenile, San Antonio del Tequendama, Cundinamarca Department, Colombia. Photograph © German Leonel Sarmiento Cruz.

opencc-zeroMay 2022View details →

Data from: Speciation in Nearctic oak gall wasps is frequently correlated with changes in host plant, host organ, or both

Quantifying the frequency of shifts to new host plants within diverse clades of specialist herbivorous insects is critically important to understand whether and how host shifts contribute to the origin of species. Oak gall wasps (Hymenoptera: Cynipidae: Cynipini) comprise a tribe of ~1000 species of phytophagous insects that induce gall formation on various organs of trees in the family Fagacae —primarily the oaks (genus Quercus; ~435 sp). The association of oak gall wasps with oaks is ancient (~50 my), and most oak species are galled by one or more gall wasp species. Despite the diversity of both gall wasp species and their plant associations, previous phylogenetic work has not identified the strong signal of host plant shifting among oak gall wasps that has been found in other phytophagous insect systems. However, most emphasis has been on the Western Palearctic and not the Nearctic where both oaks and oak gall wasps are considerably more species rich. We collected 86 species of Nearctic oak gall wasps from 10 of the 14 major clades of Nearctic oaks and sequenced >1000 Ultra Conserved Elements (UCEs) and flanking sequences to infer wasp phylogenies. We assessed the relationships of Nearctic gall wasps to one another and, by leveraging previously published UCE data, to the Palearctic fauna. We then used phylogenies to infer historical patterns of shifts among host tree species and tree organs. Our results indicate that oak gall wasps have moved between the Palearctic and Nearctic at least four times, that some Palearctic wasp clades have their proximate origin in the Nearctic, and that gall wasps have shifted within and between oak tree sections, subsections, and organs considerably more often than previous data have suggested. Given that host shifts have been demonstrated to drive reproductive isolation between host-associated populations in other phytophagous insects, our analyses of Nearctic gall wasps suggest that host shifts are key drivers of speciation in this clade, especially in hotspots of oak diversity. Though formal assessment of this hypothesis requires further study, two putatively oligophagous gall wasp species in our dataset show signals of host-associated genetic differentiation unconfounded by geographic distance, suggestive of barriers to gene flow associated with the use of alternative host plants.

One to host them all: genomics of the diverse bacterial endosymbionts of the spider Oedothorax gibbosus

Bacterial endosymbionts of the groups Wolbachia, Cardinium and Rickettsiaceae are well-known for their diverse effects on their arthropod hosts, ranging from mutualistic relationships to reproductive phenotypes. Here, we analyzed a unique system in which the dwarf spider Oedothorax gibbosus is co-infected with up to five different endosymbionts affiliated with Wolbachia, ‘Candidatus Tisiphia’ (formerly Torix group Rickettsia), Cardinium, and Rhabdochlamydia. Using short-read genome sequencing data, we show that the endosymbionts are heterogeneously distributed among O. gibbosus populations and are frequently found co-infecting spider individuals. To study this intricate host-endosymbiont system on a genome resolved level, we used long-read sequencing to reconstruct closed genomes of the Wolbachia, ‘Ca. Tisiphia’ and Cardinium endosymbionts. We provide insights in the ecology and evolution of the endosymbionts and shed light on the interactions with their spider host. We detected high quantities of transposable elements in all endosymbiont genomes and provide evidence that ancestors of the Cardinium, ‘Ca. Tisiphia’ and Wolbachia endosymbionts have co-infected the same hosts in the past. Our findings contribute to broadening our knowledge about endosymbionts infecting one of the largest animal phyla on earth, and show the usefulness of transposable elements as an evolutionary “contact-tracing” tool.

opencc-zeroJun 2022View details →

Data from: Adaptive division of growth and development between hosts in helminths with two-host life cycles

Parasitic worms (helminths) with complex life cycles divide growth and development between successive hosts. Using data from 597 species of acanthocephalans, cestodes, and nematodes with two-host life cycles, we found that helminths with larger intermediate hosts were more likely to infect larger, endothermic definitive hosts, although some evolutionarily shifts in definitive host mass occurred without changes in intermediate host mass. Life-history theory predicts parasites to shift growth to hosts in which they can grow rapidly and/or safely. Accordingly, helminth species grew relatively less as larvae and more as adults if they infected smaller intermediate hosts and/or larger, endothermic definitive hosts. Growing larger than expected in one host, relative to host mass/endothermy, was not associated with growing less in the other host, implying a lack of cross-host tradeoffs. Rather, some helminth orders had both large larvae and large adults. Within these taxa, though, size at maturity in the definitive host was unaffected by changes to larval growth, as predicted by optimality models. Parasite life-history strategies were mostly (though not entirely) consistent with theoretical expectations, suggesting that helminths adaptively divide growth and development between the multiple hosts in their complex life cycles.

opencc-zeroJun 2022View details →

Data from: Plant host traits mediated by foliar fungal symbionts and secondary metabolites

Fungal symbionts living inside plant leaves ("endophytes") can vary from beneficial to parasitic, but the mechanisms by which the fungi affect the plant host phenotype remain poorly understood. Chemical interactions are likely the proximal mechanism of interaction between foliar endophytes and the plant, as individual fungal strains are often exploited for their diverse secondary metabolite production. Here, we go beyond single strains to examine commonalities in how 16 fungal endophytes shift plant phenotypic traits such as growth and physiology, and how those relate to plant metabolomics profiles. We inoculated individual fungi on switchgrass, Panicum virgatum L. This created a limited range of plant growth and physiology (2–370% of fungus-free controls on average), but effects of most fungi overlapped, indicating functional similarities in unstressed conditions. Overall plant metabolomics profiles included almost 2000 metabolites, which were broadly correlated with plant traits across all the fungal treatments. Terpenoid-rich samples were associated with larger, more physiologically active plants and phenolic-rich samples were associated with smaller, less active plants. Only 47 metabolites were enriched in plants inoculated with fungi relative to fungus-free controls, and of these, LASSO regression identified 12 metabolites that explained from 14–43% of plant trait variation. Fungal long-chain fatty acids and sterol precursors were positively associated with plant photosynthesis, conductance, and shoot biomass, but negatively associated with survival. The phytohormone gibberellin, in contrast, was negatively associated with plant physiology and biomass. These results can inform ongoing efforts to develop metabolites as crop management tools, either by direct application or via breeding, by identifying how associations with more beneficial components of the microbiome may be affected.

opencc-zeroJul 2022View details →

Timescale reverses the relationship between host density and infection risk

Host density shapes infection risk through two opposing phenomena. First, when infective stages are subdivided among multiple hosts, greater host densities decrease infection risk through "safety in numbers". Hosts, however, represent resources for parasites, and greater host availability also fuels parasite reproduction. Hence, host density increases infection risk through "density-dependent transmission". Theory proposes that these phenomena are not disparate outcomes but occur over different timescales. That is, higher host densities may reduce short-term infection risk, but because they support parasite reproduction, may increase long-term risk. We tested this theory in a zooplankton-disease system with laboratory experiments and field observations. Supporting theory, we found that negative density-risk relationships ('safety in numbers') sometimes emerged over short timescales, but these relationships reversed to 'density-dependent transmission' within two generations. By allowing parasite numerical responses to play out, time can shift the consequences of host density, from reduced immediate risk to amplified future risk.

Fig. 30 in Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants

Fig. 30 (continued on next page). Distribution of the lutko species group fritillaries in combination with the distribution of known and suspected species of host plants from the genus Phlomoides Blume. = Melitaea lutko Evans, 1932; = M. timandra timandra Coutsis & Oorschot, 2014; = M. timandra binaludica subsp. nov.; = M. shahvarica sp. nov.; = M. mimetica mimetica Higgins, 1940; = M. mimetica delerei Heidemann, 1954;? = unconfirmed findings of M. timandra; = M. timandra with an unclear subspecies status; = Phlomoides regeliana (Aitch. & Hemsl.) Adylov, Kamelin & Makhm.; = Phlomoides boissieriana (Regel) Adylov, Kamelin & Makhm.; = Phlomoides laciniata (L.) Kamelin & Makhm.; = Phlomoides labiosiformis (Popov) Adylov, Kamelin & Makhm.; = Phlomoides loasifolia (Benth.) Kamelin & Makhm.; = Phlomoides molucelloides (Bunge) Salmaki; = Phlomoides acaulis (Beck ex Rech.f.) Salmaki; = Phlomoides labiosa (Bunge) Adylov, Kamelin & Makhm. A. Pakistan, Chitral, Chaghbini CGNP, alt. 2700–3000 m. B. Pakistan, Khyber Pakhtunkhwa, Drosh. C. Pakistan, Khyber Pakhtunkhwa, Keon Nullah. D. Pakistan, Khyber Pakhtunkhwa, Malakand. E. Pakistan, Khyber Pakhtunkhwa, Birmoglasht. F. Turkmenistan, Badkhyz, Kepeli, alt. 700 m. G. Turkmenistan, Badkhyz, Kyzyl-Jar, alt. 700 m. H. Turkmenistan, Kushka, alt. 700 m. I. Turkmenistan, Murgab river, Sary-Yazy, alt. 300 m. J. Turkmenistan, 30 km E of BairamAli, Zahmet, alt. 240 m. K. Turkmenistan, Bairam-Ali, alt. 230 m. L. Turkmenistan, Kara-Kum desert, 30 km W of Mary, alt. 200 m.M. Turkmenistan, Dushak, alt. 250 m.N. Turkmenistan, Chaacha, alt. 400 m. O. Turkmenistan, Bakharden, alt. 200 m. P. Iran, Khorossan Razavi, Kuh-e-Binalud Mts, Qadamgah area, Gerina, alt. 2000 m. Q. Iran, Khorasan Razavi, Kuh-e-Binalud Mts, 15 km SW of Zoshk, alt. 2300– 2500 m. R. Iran, S Khorosan, 75 km N of Birjant, Sedeh, alt. 1500 m. S. Iran, S Khorosan, 35 km N of Birjant, alt. 1500 m. T. Afghanistan, Bamian, Band-e-Amir, Dzhudoi-Kvak Gorge, alt. 3200 m. U. Afghanistan, Bamian, Band-e-Amir, Hazarajat, alt. 3000–3200 m. V. Afghanistan, Bamian, Koh-iBaba Mts, Joshanak, alt. 2800 m. W. Afghanistan, Heart, Qala-i-Naw, Kashka pass. X. Iran, Semnan, Shahvar Mt., alt. 2200–2500 m. Y. Turkmenistan, Kara-Kala, Monjukly Ridge, 300–700 m. Z. Iran, Golestan, E Maraveh Tappeh, N Ghazan Ghayeh, Palizan Mts. A". Pakistan, Balochistan, Quetta, Urak, alt. 2500 m. B". Pakistan, Balochistan, Ziarat, alt. 2500 m. C". Pakistan, Balochistan, Khojak, alt. 1700 m. D". Pakistan, Balochistan, Zaghum, alt. 1600 m; E". Pakistan, Punjab, Gawar, alt. 500 m. F". Pakistan, Balochistan, Sheik Wazil, alt. 1600 m. G". Afghanistan, Bamian, Hushkak, alt. 2700–2800 m. H". Afghanistan, Bamian, Punjub Distr., 10 km NE of Varas, alt. 2400 m. I". Afghanistan, Ghor, 17 km E of Changcharan, 15 km S of Bandi-Ali, Gazak Mts, alt. 2400 m. J". Afghanistan, Ghor, Bayan Range, 15 km S of Changcharan, Kindival valley, alt. 2700 m. K". Afghanistan, Bamiyan, Kohi-Baba Mts, Panjao, alt. 3000 m. Afghanistan, Bamiyan, Koh-i-Baba Mts, Shah-tu-Kotal, alt. 4000 m. L". Afghanistan, Kapisa, Pandshir valley, alt. 2200–2800 m. M". Afghanistan, Kabul. N". Iran, Tehran, Elburz Ridge, Demavend Mt., Ask, alt. 1800 m. O". Iran, Semnan, Foulad Mohaleh, alt. 2200 m. P". Pakistan, Punjab, Murree.

Fig. 29 in Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants

Fig. 29.Differences in the structure of valva and aedeagus of the lutko species group.A. Melitaea shahvarica sp. nov. B, D, H. M. timandra binaludica subsp. nov. C–E. M. mimetica Higgins, 1940. F. M. lutko Evans, 1932. G. M. timandra timandra Coutsis &van Oorschot, 2014. A. Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mts, alt. 2200–2400 m. B. Iran, Rezavi Khorassan Prov., Kuh-e-Binalud Mts, Dorrud v. vicinity, alt. 2430 m. C. Afghanistan, Bamian Prov., Punjub Distr., 10 km NE of Varas v., alt. 2400 m. D. Afghanistan, Band-i-Amir, Hazarajat. E. Pakistan, Balochistan, Quetta, Urak, alt. 2400– 2700 m. F. Pakistan, Chitral, Gol National Park, alt. 2700 m. G. Turkmenistan, Sary-Yazy, alt. 700 m. H. Iran, Rezavi Khorassan Prov., Kuh-e-Binalud Mts, Dorrud v. vicinity, alt. 2430 m.

Fig. 28 in Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants

Fig. 28.Eggs and caterpillars of Melitaea shahvarica sp. nov. in nature and in the laboratory.A–B. Freshly laid eggs under a leaf of a host plant, May 2018, Iran, Shahvar Mt., alt. 2200 m. C–D. IV–V instar caterpillars on the leaves of the host plant Phlomoides molucelloides (Bunge) Salmaki, July 2019, Iran, Shahvar Mt., alt. 2500 m. E. I instar caterpillars in the laboratory, Moscow, May 2018. F. VI instar caterpillars during diapause, Moscow, October 2018.

Fig. 18. Male genitalia and harpe. A–C in Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants

Fig. 18. Male genitalia and harpe. A–C. Melitaea shahvarica sp. nov. D–E. M. lutko Evans, 1932. F–I. M. mimetica Higgins, 1940. A–C. Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mts, alt. 2200–2400 m. D–E. Pakistan, Chitral, Chaghbini, CGNP [Chitral Gol National Park], alt. 2700 m. F–G. Afghanistan, Bamian Prov., Punjub Distr., 10 km. NE Varas v., alt. 2400 m. H–I. Afghanistan, Bamian Prov., Panjub Distr., 10 km. NE Varas vil., alt. 2400 m.

Fig. 8 in Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants

Fig. 8. Distribution of Melitaea mimetica Higgins, 1940. For a description of the symbols with letters, see Fig. 30. = M. mimetica mimetica Higgins, 1940; = M. mimetica delerei Heidemann, 1954.

Fig. 14 in Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants

Fig. 14. Distribution of Melitaea timandra timandra Coutsis & van Oorschot, 2014, M. timandra binaludica subsp. nov. and M. shahvarica sp. nov. For a description of the points by letters, see Fig. 30. = Melitaea timandra timandra; = Melitaea timandra binaludica subsp. nov.; = Melitaea shahvarica sp. nov.;? = unconfirmed finds of Melitaea timandra; = Melitaea timandra with uncertain subspecies status.

Fig. 17 in Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants

Fig. 17. Male genitalia (without aedeagus) in dorsal projection. A–C. Melitaea timandra timandra Coutsis & van Oorschot, 2014. D–I. M. timandra binaludica subsp. nov. A–C. S Turkmenistan, SaryYazy, alt. 300 m. D –F. Iran, Rezavi Khorassan Prov., Kuh-e-Binalud Mts, Dorrud v. vicinity, alt. 2430 m. G. Iran, Horossan Prov., 35 km N of Birjant t. H. Afghanistan, Bamian Prov., Band-e-Amir, alt. 3200 m. I. Central Afghanistan, Bamian Prov., Band-e-Amir, Dzhudoi-Kvak Gorge, alt. 3200 m.

Fig. 24 in Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants

Fig. 24. First instar caterpillar of Melitaea shahvarica sp. nov. A. Head, bottom view. B. Head, front view. C–D. Head, lateral view. E–F. Caterpillar, lateral view.

Fig. 6 in Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants

Fig. 6. Adults of M. mimetica Higgins, 1940. A –H. UPS. I –P. UNS. A–B, I–J. ♂, Pakistan, Balochistan, Quetta, Urak, alt. 2400–2700 m. C, K. ♂, Afghanistan, Ghowr Prov., Bayan Range, 15 km S of Chagcharan, Kindival vall. D, L. ♂, Afghanistan, Bamian Prov., Punjub Distr., 10 km NE of Varas v., alt. 2400 m. E–F, M–N. ♀, Pakistan, Balochistan, Quetta, Urak, alt. 2400–2700 m. G, O. ♀, Central Afghanistan, Bamian Prov., 8 km S of Bamian, Koh-e-Baba Mts, Dara-e-Khushkak, alt. 2930 m. H, P. ♀, Afghanistan, Bamian Prov., Punjub Distr., 10 km NE of Varas v., alt. 2400 m. B, J photos by Dr Eckweiler).

Fig. 5 in Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants

Fig. 5. Distribution of M. lutko Evans, 1932. For a description of the dots with letters see Fig. 30.

Fig. 4 in Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants

Fig. 4. Female genitalia (without the anal papillae, copulatory bursa, and anterior and posterior apophyses). A–D. M. shahvarica sp. nov., Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mts, alt. 2200–2400 m. E. M. lutko Evans, 1932, Chitral Gol National Park, Chitral, Pakistan, alt. 2700 m. F–I. M. mimetica Higgins, 1940. F. Pakistan, Balochistan, Quetta, Urak, alt. 2400– 2700 m. G–H. Afghanistan, Bamian Prov., Punjub Distr., 10 km NE of Varas v., alt. 2400 m. I. Central Afghanistan, Ghor Prov., 17 km E of Changcharan, 15 km S of Bandi-Ali, Gazak Mts, alt. 2400 m.

Fig. 13 in Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants

Fig. 13. Biotopes and biology of Melitaea timandra timandra Coutsis & van Oorschot, 2014. A. Bumpycellular overgrown sands on the left bank of the Murgab River, alt. 300 m, April 1993, the habitat of M. timandra timandra. B. Phlomoides regeliana (Aitch. & Hemsl.) Adylov, Kamelin & Makhm. – host plant for M. timandra timandra. The female laying eggs is visible under the leaves of the plant. C. The female of M. timandra timandra is ovipositing on the soil under the host plant leaf. D. Egg clutch of M. timandra timandra on the basal leaf rosette of Ph. regeliana.