Special Feature: Diversity of insect-plant interactions in the eastern Andes of Ecuador

James S. Miller1,2,a and Lee Dyer1,b

1 Department of Biology, University of Nevada, Reno, NV 89557
2 American Museum of Natural History, Division of Invertebrate Zoology, New York, NY

Correspondence: ajmiller@amnh.org, bldyer@tulane.edu

Published: 2 June 2009

ISSN: 1536-2442 | Volume 9, Number 26

Cite this paper as:
Miller JS, Dyer L. 2009. Special Feature: Diversity of insect-plant interactions in the eastern Andes of Ecuador. Journal of Insect Science 9:26, available online: insectscience.org/9.26


Introduction
Table of Contents
References

Over half the described organisms in the world are involved in plant-insect-parasitoid interactions (Hawkins and Sheehan 1994), and these provide the basis for our understanding of fundamental issues in ecology and evolutionary biology (e.g., Hairston et al. 1960; Ehrlich and Raven 1964; Janzen 1973; Lawton and McNeill 1979; Price et al. 1980; Karban and Myers 1989; Gentry and Dyer 2002). In fact, relationships between parasitoids, herbivorous insects, and their host plants are among the most productive systems for understanding multi-trophic interactions (e.g., Abrahamson and Weiss 1997; English-Loeb et al. 1993; Hochberg and Ives 2000; Jervis and Kidd 1996; Quicke 1997; Singer and Stireman 2003; Whitfield 1998, 2003). Documenting these ubiquitous systems has lagged far behind more traditional biotic surveys and inventories. The resulting lack of basic natural history data has led to unsubstantiated paradigms, as well to numerous disagreements in ecology and evolutionary biology. For example, it has been assumed commonly that herbivorous insects are highly host-specific (reviewed by Futuyma and Moreno 1988; Jaenike 1990; Irschick et al. 2005), especially in the tropics. More recently, however, this paradigm has been challenged by the results of an extensive survey of herbivore-plant interactions (Novotny et al. 2006), sparking a vigorous debate (Dyer et al. 2007; Stork 2007). Collecting and inventory methods that rely on rearing wild-caught specimens preserve much of the ecological, environmental and behavioral context of specimens, while also providing high quality taxonomic data.

Caterpillars and Parasitoids of the Eastern Andes in Ecuador is an ongoing, long-term rearing project dedicated to the inventory and dissemination of information on lepidopteran larvae, their host plants and their parasitoids. It provides important natural history data from a uniquely diverse and highly threatened ecosystem. This survey, officially begun in 2001 with funds provided by Earthwatch Institute, has continued with funding from a variety of sources, including National Geographic and the National Science Foundation. It is part of a coordinated trans-American effort taking place at sites in Ecuador, Costa Rica, Arizona and Louisiana (Dyer et al. 2007; Stireman et al. 2005; Gentry and Dyer 2002; www.caterpillars.org). The primary goals of the project are:

  1. To survey and inventory a diverse community of Lepidoptera and their associated parasitoid Hymenoptera and Diptera, thus providing specimens for future morphological and molecular systematic research.

  2. To collect baseline natural history information documenting caterpillar-host plant-parasitoid relationships, larval development rates, and other life cycle information for described and undescribed species. These data can be used to test ecological and evolutionary hypotheses.

  3. To disseminate this information via a searchable database, publicly accessible through the internet and available worldwide.

Ecuador is at the edge of the biological frontier when it comes to our understanding of insect-plant biodiversity and the taxonomic composition of tropical biological communities. On the slopes of the equatorial Andes, considered by many to be home to the highest levels of species endemism in the world (e.g., Brehm et al. 2005), most plant and animal groups remain poorly studied (Suarez and Ulloa 1993; Kessler 2000; Cresswell et al. 1999). Due to rapid rates of habitat destruction and growing human populations, these montane habitats are under immediate danger of deforestation (Suarez and Ulloa 1993). Areas of relatively low relief, more suitable for cattle grazing, are under particular threat.

Yanayacu Biological Station (YBS) is located at 2200 meters in the Quijos Valley, Napo Province, in the Andes of northeastern Ecuador (00°35.9’S, 77°53.4’W). The intermediate elevation of the station (situated in montane wet forest sensu Holdridge et al. 1971) provides easy access to a unique diversity of habitats, ranging from paramo (3800 m) to lowland rain forest (800 m and below). Much of the 2000 hectares encompassed by the YBS comprises flat, pristine cloud forest, some of the last remaining habitat of this type found anywhere in Ecuador.

This series of articles showcases basic natural history information, as well as complex tri-trophic interactions, in a poorly known fauna (Neotropical caterpillars and host plants, as well as their parasitic Hymenoptera and Diptera). Integrated information of this sort, novel even for well-studied faunas, is crucial for those wanting to test hypotheses involving tropical taxa. In this special feature, adult Lepidoptera and parasitoids are associated with their immature stages, providing new taxonomic characters for the families Geometridae, Nymphalidae, Hesperiidae, Tachinidae and Braconidae, groups for which the systematics and biodiversity are poorly known (e.g., Scoble et al. 1995; Penz 1999; Dolphin and Quicke 2001; Whitfield et al. 2001; Penz and DeVries 2002; Greeney and Jones 2003). The information generated by this project has also allowed examination of diversity patterns at larger geographical scales (see Dyer et al. 2007; Connahs et al. this issue) and across habitat types, because the sampling protocol includes Andean habitats broadly distributed along a north-south axis and across an elevational gradient.

These papers are a first attempt to summarize some of the taxonomic and ecological results from the rearing inventory in Ecuador. The overall collecting effort encompasses a huge variety of caterpillar and host plant taxa, but certain focal groups were particularly well sampled. These include the host plant genera Piper (Piperaceae) and Chusquea (Poaceae), lepidopteran larvae in the genus Eois (Geometridae), as well as caterpillars in the butterfly families Hesperiidae and Nymphalidae. Future features will provide broad syntheses treating additional taxonomic groups, such as the Arctiidae, Saturniidae, Sphingidae, Noctuidae and Limacodidae, as the data accumulate and species identifications become further refined.

Introduction
Table of Contents
References

Table of Contents

Characterizing the cauline domatia of two newly discovered Ecuadorian ant plants in Piper: an example of convergent evolution
Eric J. Tepe, Walter A. Kelley, Genoveva Rodriguez-Castañeda, and Lee A. Dyer 		HTML PDF
Geographic variation in host-specificity and parasitoid pressure of an herbivore (Geometridae) associated with the tropical genus Piper (Piperaceae)
Heidi Connahs, Genoveva Rodríguez-Castañeda, Toni Walters, Thomas Walla, and Lee Dyer 		HTML PDF
A key to New World Distatrix Mason (Hymenoptera: Braconidae), with descriptions of six new reared Neotropical species
Christopher C. Grinter, James B. Whitfield, Heidi Connahs, Lee A. Dyer, Winifred Hallwachs, and Daniel H. Janzen 		HTML PDF
The early stages and natural history of Antirrhea adoptiva porphyrosticta (Watkins, 1928) in eastern Ecuador (Lepidoptera: Nymphalidae: Morphinae)
Harold F. Greeney, Philip J. DeVries, Carla M. Penz, Rafael B. Granizo, Heidi Connahs, John O. Stireman, Thomas R. Walla, and Lee A. Dyer 		HTML PDF
Reared microgastrine wasps (Hymenoptera: Braconidae) from Yanayacu Biological Station and environs (Napo Province, Ecuador): Diversity and host specialization
James B. Whitfield, Josephine J. Rodriguez, and Paul K. Masonick 		HTML PDF
The life history and shelter building behavior of Vettius coryna coryna Hewitson, 1866 in eastern Ecuador (Lepidoptera: Hesperiidae: Hesperiinae)
Harold F. Greeney and Andrew D. Warren 		HTML PDF
The immature stages and shelter building behavior of Falga jeconia ombra Evans, 1955 in eastern Ecuador (Lepidoptera: Hesperiidae: Hesperiinae)
Harold F. Greeney and Andrew D. Warren 		HTML PDF
A new species of solitary Meteorus (Hymenoptera: Braconidae) reared from caterpillars of toxic butterflies (Lepidoptera: Nymphalidae) in Ecuador
Scott R. Shaw and Guinevere Z. Jones 		HTML PDF
The immature stages and natural history of Veladyris pardalis (Salvin, 1869) in eastern Ecuador (Lepidoptera: Nymphalidae: Ithomiinae)
Harold F. Greeney, Ryan I. Hill, Marco Ramiro Hualingua-L, Wilmer Rosendo Simbaña, and Grant Gentry 		HTML PDF
A new species of Andesipolis Whitfield & Choi from the eastern Andes of Ecuador with notes on biology and classification (Hymenoptera: Braconidae: Rhysipolinae)
Andrew C. Townsend and Scott R. Shaw 		HTML PDF
Nine new species of Aleiodes Wesmael reared at Yanayacu Biological Station (Hymenoptera: Braconidae: Rogadinae) in eastern Ecuador
Andrew C. Townsend and Scott R. Shaw 		HTML PDF
The early stages of Pedaliodes poesia (Hewitson, 1862) in eastern Ecuador (Lepidoptera: Satyrinae: Pronophilina)
Harold F. Greeney, Tomasz W. Pyrcz, Marco Ramiro Hualingua-L, Philip J. DeVries, and Lee A. Dyer 		HTML PDF
Species richness and host associations of Lepidoptera- attacking Tachinidae in the northeast Ecuadorian Andes
John O. Stireman, Harold F. Greeney, and Lee A. Dyer 		HTML PDF
Introduction
Table of Contents
References

References

  1. Abrahamson WG, Weiss AE. 1997. Evolutionary Ecology across Three Trophic Levels: Goldenrods, Gallmakers, and Natural Enemies. Princeton University Press.  Abrahamson WG, Weiss AE. 1997. Evolutionary Ecology across Three Trophic Levels: Goldenrods, Gallmakers, and Natural Enemies. Princeton University Press.
  2. Brehm G, Pitkin LM, Hilt N, Fiedler K. 2005. Montane Andean rain forests are a global diversity hotspot of geometrid moths. Journal of Biogeography 32: 1621-1627. Brehm G, Pitkin LM, Hilt N, Fiedler K. 2005. Montane Andean rain forests are a global diversity hotspot of geometrid moths. Journal of Biogeography 32: 1621-1627.
  3. Cresswell W, Hughes M, Mellanby R, Bright S, Catry P, Chaves J, Freile J, Gabela A, Martineau H, Macleod R, Mcphie F, Anderson N, Holt S, Barabas S, Chapel C, Sanchez T. 1999. Densities and habitat preferences of Andean cloud-forest birds in pristine and degraded habitats in north-eastern Ecuador. Bird Conservation International 9: 129-145. Cresswell W, Hughes M, Mellanby R, Bright S, Catry P, Chaves J, Freile J, Gabela A, Martineau H, Macleod R, Mcphie F, Anderson N, Holt S, Barabas S, Chapel C, Sanchez T. 1999. Densities and habitat preferences of Andean cloud-forest birds in pristine and degraded habitats in north-eastern Ecuador. Bird Conservation International 9: 129-145.
  4. Dolphin K, Quicke DLJ. 2001. Estimating the global species richness of an incompletely described taxon: an example using parasitoid wasps (Hymenoptera: Braconidae). Biological Journal of the Linnean Society 73: 279-286. Dolphin K, Quicke DLJ. 2001. Estimating the global species richness of an incompletely described taxon: an example using parasitoid wasps (Hymenoptera: Braconidae). Biological Journal of the Linnean Society 73: 279-286.
  5. Dyer LA, Singer MS, Lill JT, Stireman JO, Gentry GL, Marquis RJ, Ricklefs RE, Greeney HF, Wagner DL, Morais HC, Diniz IR, Kursar TA, Coley PD. 2007. Host specificity of Lepidoptera in tropical and temperate forests. Nature 448: 696-699. Dyer LA, Singer MS, Lill JT, Stireman JO, Gentry GL, Marquis RJ, Ricklefs RE, Greeney HF, Wagner DL, Morais HC, Diniz IR, Kursar TA, Coley PD. 2007. Host specificity of Lepidoptera in tropical and temperate forests. Nature 448: 696-699.
  6. Ehrlich PR, Raven PH. 1964. Butterflies and plants: a study in coevolution. Evolution 18: 568-608. Ehrlich PR, Raven PH. 1964. Butterflies and plants: a study in coevolution. Evolution 18: 568-608.
  7. English-Loeb GM, Brody AK, Karban R. 1993. Host-Plant-Mediated interactions between a generalist folivore and its tachinid parasitoid. Journal of Animal Ecology 62: 465-471. English-Loeb GM, Brody AK, Karban R. 1993. Host-Plant-Mediated interactions between a generalist folivore and its tachinid parasitoid. Journal of Animal Ecology 62: 465-471.
  8. Futuyma DJ, Moreno B. 1988. The evolution of ecological specialization. Annual Review of Ecology and Systematics 19: 207-233. Futuyma DJ, Moreno B. 1988. The evolution of ecological specialization. Annual Review of Ecology and Systematics 19: 207-233.
  9. Gentry GL, Dyer LA. 2002. On the conditional nature of neotropical caterpillar defenses against their natural enemies. Ecology 83: 3108-3119. Gentry GL, Dyer LA. 2002. On the conditional nature of neotropical caterpillar defenses against their natural enemies. Ecology 83: 3108-3119.
  10. Greeney HF, Jones MT. 2003. Shelter building in the Hesperiidae: a classification scheme for larval shelters. Journal of Research on the Lepidoptera 37: 27-36. Greeney HF, Jones MT. 2003. Shelter building in the Hesperiidae: a classification scheme for larval shelters. Journal of Research on the Lepidoptera 37: 27-36.
  11. Hairston NG, Smith FE, Slobodkin LB. 1960. Community structure, population control, and competition. American Naturalist 94: 421-424. Hairston NG, Smith FE, Slobodkin LB. 1960. Community structure, population control, and competition. American Naturalist 94: 421-424.
  12. Hawkins BA, Sheehan W. 1994. Parasitoid Community Ecology. Oxford University Press.  Hawkins BA, Sheehan W. 1994. Parasitoid Community Ecology. Oxford University Press.
  13. Hochberg ME, Ives AR. 2000. Parasitoid Population Biology. Princeton University Press.  Hochberg ME, Ives AR. 2000. Parasitoid Population Biology. Princeton University Press.
  14. Holdridge LR, Grenke WC, Hatheway WH, Liang T, Tosi JA. 1971. Forest Environments in Tropical Life Zones. Pergamon Press.  Holdridge LR, Grenke WC, Hatheway WH, Liang T, Tosi JA. 1971. Forest Environments in Tropical Life Zones. Pergamon Press.
  15. Irschick D, Dyer L, Sherry TW. 2005. Phylogenetic methodologies for studying specialization. Oikos 110: 404-408. Irschick D, Dyer L, Sherry TW. 2005. Phylogenetic methodologies for studying specialization. Oikos 110: 404-408.
  16. Jaenike J. 1990. Host specialization in phytophagous insects. Annual Review of Ecology and Systematics 21: 243-273. Jaenike J. 1990. Host specialization in phytophagous insects. Annual Review of Ecology and Systematics 21: 243-273.
  17. Janzen DH. 1973. Comments on host-specificity of tropical herbivores and its relevance to species richness. In: Heywood VH, editor. Taxonomy and Ecology, pp. 201-211. Academic Press.  Janzen DH In: Heywood VH, editor. 1973. Comments on host-specificity of tropical herbivores and its relevance to species richness. Taxonomy and Ecology201-211. Academic Press.
  18. Jervis MA, Kidd NAC. 1996. Insect Natural Enemies: Practical Approaches to their Study and Evaluation. Chapman and Hall.  Jervis MA, Kidd NAC. 1996. Insect Natural Enemies: Practical Approaches to their Study and Evaluation. Chapman and Hall.
  19. Karban R, Myers JH. 1989. Induced plant responses to herbivory. Annual Review of Ecology and Systematics331-348. Karban R, Myers JH. 1989. Induced plant responses to herbivory. Annual Review of Ecology and Systematics331-348.
  20. Kessler M. 2000. Elevational gradients in species richness and endemism of selected plant groups in the central Bolivian Andes. Plant Ecology 149: 181-193. Kessler M. 2000. Elevational gradients in species richness and endemism of selected plant groups in the central Bolivian Andes. Plant Ecology 149: 181-193.
  21. Lawton JH, McNeill S. 1979. Between the devil and the deep blue sea: on the problem of being a herbivore. Symposium of the British Ecological Society 20: 223-244. Lawton JH, McNeill S. 1979. Between the devil and the deep blue sea: on the problem of being a herbivore. Symposium of the British Ecological Society 20: 223-244.
  22. Novotny V, Drozd P, Miller SE, Kulfan M, Janda M, Basset Y, Weiblen GD. 2006. Why are there so many species of herbivorous insects in tropical rainforests? Science 313: 1115-1118. Novotny V, Drozd P, Miller SE, Kulfan M, Janda M, Basset Y, Weiblen GD. 2006. Why are there so many species of herbivorous insects in tropical rainforests? Science 313: 1115-1118.
  23. Penz CM. 1999. Higher level phylogeny for the passion-vine butterflies (Nymphalidae, Heliconiinae) based on early stage and adult morphology. Zoological Journal of the Linnean Society 127: 277-344. Penz CM. 1999. Higher level phylogeny for the passion-vine butterflies (Nymphalidae, Heliconiinae) based on early stage and adult morphology. Zoological Journal of the Linnean Society 127: 277-344.
  24. Penz CM, DeVries PJ. 2002. Phylogenetic analysis of Morpho butterflies (Nymphalidae, Morphinae): Implications for classification and natural history. American Museum Novitates 3374: 1-33. Penz CM, DeVries PJ. 2002. Phylogenetic analysis of Morpho butterflies (Nymphalidae, Morphinae): Implications for classification and natural history. American Museum Novitates 3374: 1-33.
  25. Price PW, Bouton EE, Gross P, McPheron BA, Thompson JN, Weiss AE. 1980. Interactions among three trophic levels: influence of plants on interactions between insect herbivores and natural enemies. Annual Review of Ecology and Systematics 11: 41-65. Price PW, Bouton EE, Gross P, McPheron BA, Thompson JN, Weiss AE. 1980. Interactions among three trophic levels: influence of plants on interactions between insect herbivores and natural enemies. Annual Review of Ecology and Systematics 11: 41-65.
  26. Quicke DLJ. 1997. Parasitic Wasps. Chapman & Hall.  Quicke DLJ. 1997. Parasitic Wasps. Chapman & Hall.
  27. Scoble MJ, Gaston KJ, Crook A. 1995. Using taxonomic data to estimate species richness in Geometridae. Journal of the Lepidopterists’ Society 49: 136-147. Scoble MJ, Gaston KJ, Crook A. 1995. Using taxonomic data to estimate species richness in Geometridae. Journal of the Lepidopterists’ Society 49: 136-147.
  28. Singer MS, Stireman JO. 2003. Does anti-parasitoid defense explain host-plant selection by a polyphagous caterpillar? Oikos 100: 554-562. Singer MS, Stireman JO. 2003. Does anti-parasitoid defense explain host-plant selection by a polyphagous caterpillar? Oikos 100: 554-562.
  29. Stireman JO, Dyer LA, Janzen DH, Singer MS, Li JT, Marquis RJ, Ricklefs RE, Gentry GL, Hallwachs W, Coley PD, Barone JA, Greeney HF, Connahs H, Barbosa P, Morais HC, Diniz IR. 2005. Climatic unpredictability and parasitism of caterpillars: Implications of global warming. Proceedings of the National Academy of Sciences of the United States of America 102: 17384-17387. Stireman JO, Dyer LA, Janzen DH, Singer MS, Li JT, Marquis RJ, Ricklefs RE, Gentry GL, Hallwachs W, Coley PD, Barone JA, Greeney HF, Connahs H, Barbosa P, Morais HC, Diniz IR. 2005. Climatic unpredictability and parasitism of caterpillars: Implications of global warming. Proceedings of the National Academy of Sciences of the United States of America 102: 17384-17387.
  30. Stork NE. 2007. Biodiversity - World of insects. Nature 448: 657-658. Stork NE. 2007. Biodiversity - World of insects. Nature 448: 657-658.
  31. Suarez L, Ulloa R. 1993. La Diversidad Biologica del Ecuador. In: Mena PA, Suarez L, editors. La Investigacion para la Conservacion de la Diversidad Bioliogica en al Ecuador, pp. 13-27. EcoCiencia, Ecuador.  Suarez L, Ulloa R. 1993. La Diversidad Biologica del Ecuador. In: Mena PA, Suarez L, editors. La Investigacion para la Conservacion de la Diversidad Bioliogica en al Ecuador, pp. 13-27. EcoCiencia, Ecuador.
  32. Whitfield JB. 2003. Phylogenetic insights into the evolution of parasitism in Hymenoptera. In: Littlewood TJ, editor. The Evolution of Parasitism - A Phylogenetic Approach. Advances in Parasitology 54: , pp. 69-100. Whitfield JB In: Littlewood TJ, editor. 2003. Phylogenetic insights into the evolution of parasitism in Hymenoptera. The Evolution of Parasitism - A Phylogenetic Approach. Advances in Parasitology 54: 69-100.
  33. Whitfield JB. 1998. Phylogeny and evolution of host-parasitoid interactions in hymenoptera. Annual Review of Entomology 43: 129-151. Whitfield JB. 1998. Phylogeny and evolution of host-parasitoid interactions in hymenoptera. Annual Review of Entomology 43: 129-151.
  34. Whitfield JB, Cameron SA, Ramirez SR, Roesch K, Messinger S, Taylor M. 2001. Review of the Apanteles species (Hymenoptera: Braconidae) attacking Lepidoptera in Bombus (Fervidobombus) (Hymenoptera: Apidae) colonies in the New World, with description of a new species from tropical South America. Annals of the Entomological Society of America 94: 851-857. Whitfield JB, Cameron SA, Ramirez SR, Roesch K, Messinger S, Taylor M. 2001. Review of the Apanteles species (Hymenoptera: Braconidae) attacking Lepidoptera in Bombus (Fervidobombus) (Hymenoptera: Apidae) colonies in the New World, with description of a new species from tropical South America. Annals of the Entomological Society of America 94: 851-857.