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Trophic Interactions Involving Herpetogramma Phaeopteralis (Lepidoptera: Pyralidae) and Passiflora Incarnata (Passifloraceae)

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Trophic Interactions Involving Herpetogramma Phaeopteralis (Lepidoptera: Pyralidae) and Passiflora Incarnata (Passifloraceae) 136 Florida Entomologist 91(1) March 2008 TROPHIC INTERACTIONS INVOLVING HERPETOGRAMMA PHAEOPTERALIS (LEPIDOPTERA: PYRALIDAE) AND PASSIFLORA INCARNATA (PASSIFLORACEAE) ANDREI SOURAKOV McGuire Center for Lepidoptera Research, Florida Museum of Natural History, University of Florida, Gainesville, FL 32611 E-mail: asourakov@flmnh.ufl.edu During several nights in Aug, 2007, I observed nidae), feeds on EFN of Neoraimondia arequipen- tropical sod webworm moths, Herpetogramma sis cactus (Vila & Eastwood 2006). Hesperopsis phaeopteralis Guenée (Lepidoptera: Pyralidae) graciliae (MacNeill) (Lepidoptera: Hesperiidae) swarming in large numbers around flowering pur- commutes back and forth between its host plant ple passion vines, Passiflora incarnata L. (Passi- and mesquite, Prosopis glandulosa (Fabaceae), floraceae) in suburban north central Florida. utilizing the EFN of the latter plant (Wesenborn Tropical sod webworms use a variety of Poaceae 1997). Pink bollworm moths, Pectinophora gossyp- species as larval host plants and probably were iella (Saunders) (Lepidoptera: Gelechiidae) are at- emerging from sod grass. Upon closer observation tracted to cotton flower volatiles that serve as with flash photography, I found that moths fed on chemical cues for the moths to find the EFN (Wie- the extrafloral nectaries (EFN) located on the senborn & Baker 1990). A variety of African lipte- bracteoles of the plants (Fig. 1). In this note, I am nine lycaenid butterflies (Lycaenidae) feed on reporting this observation with the understand- EFN of bamboos and vines (Callaghan 1992). ing that (like many observations in tropical biol- The importance of feeding on EFN was shown ogy) it might be based on a sporadic rather than in increased fertility in soybean looper, Pseudo- on a reoccurring interaction. I also discuss poten- plusia includens (Walker) (Lepidoptera: Noctu- tial ecological significance of this interaction idae). It was even suggested that cultivars of cot- while reviewing the pertinent literature. ton lacking EFN should be used to reduce her- Exploitation of EFN by a variety of insects has bivory and thus increase crop production (Beach been reported for a number of plants. For instance, et al. 1985). Oviposition by Heliothis puntigera Ipomoea carnea (Convolvulaceae) in Costa Rica Wallengren (Lepidoptera: Noctuidae) on cotton attracts insects of 70 different families of Co- also was correlated with EFN production (Adjei- leoptera, Hemiptera, Lepidoptera, Neuroptera, Maafo & Wilson 1983). Fecundity and longevity of Diptera, and Hymenoptera (Keeler 1978). A dry Grapholita molesta (Busck) (Lepidoptera: Tortri- area specialist in northwestern Peru, Strymon jac- cidae) doubled when moths were provided EFN of queline Nicolay & Robbins (Lepidoptera: Lycae- peaches (Atanassov & Shearer 2005). EFN has high nutritional value. It was shown to be better than sucrose for increasing flight abil- ity of Cotesia glomerata L. (Hymenoptera: Bra- conidae) (Wanner et al. 2006), and better than honey for increasing longevity and fecundity of Microplitis croceipes (Cresson) (Hymenoptera: Braconidae) (Roese et al. 2006). EFN-feeding had similar beneficial effect on Trichogramma minu- tum Riley (Hymenoptera: Trichogrammatidae) (Shearer & Atanassov 2004) and on lacewing lar- vae (Limburg & Rosenheim 2001). Attraction of such natural enemies to EFN helps suppress her- bivores. For instance, parasitism of Lepidoptera larvae is higher, while diversity of herbivore spe- cies is lower on the EFN-bearing plants of Solanum adherens Roem. and Shult. (Solanaceae) than on species of Solanum lacking nectaries (Gentry 2003). According to Rudgers (2004), EFN production can have detrimental properties for a plant, by the way of attracting female moths, which not only feed on EFN but also may oviposit on the Fig. 1. Tropical sod webworm moth, Herpetogramma plant. Ants were thought to be frequently exploi- phaeopteralis Guenée, feeding on extrafloral nectary of tive of EFN-producing plants (e.g., Wheeler 1913). Passiflora incarnata L. Ants attracted to extrafloral nectaries can deter Scientific Notes 137 pollinators such as bees from flowers (Ness 2006). access to ants is denied (pers. obs.). As it was, de- In the neotropics, several coexisting Passiflora spite of hundreds of eggs laid on the plants daily, species develop different degrees of attractiveness few survived past the first instar due to preda- to ants through differences in their EFN develop- tion. However, after the first cold temperatures in ment and productivity, which suggests that there Nov, the EFNs seemingly shut down production, are costs and benefits to EFN presence (Apple & little attendance by ants was observed, and the Feener 2001). Nevertheless, attendance of extra- survival of the caterpillars feeding on the vines floral nectaries by ants was shown to reduce her- increased dramatically, leading to complete defo- bivory in Passiflora (e.g., Labeyrie et al. 2001), liation of the vines. From the view point of benefit and specifically in P. incarnata (McLaine 1983). to the plants, such strategy is explicable because In a field experiment, young Heliconius (Lepi- the above-ground tissues of P. incarnate typically doptera: Nymphalidae) larvae disappeared from are killed by freezing temperatures in north cen- Passiflora plants with EFN at a rate double that tral Florida. of plants without the nectaries (Smiley 1985). By Extrafloral nectar feeding by H. phaeopteralis protecting plants against herbivores and nectar- reported here possibly falls into the category of stealing butterflies, ants positively influenced re- quadro-trophic interactions involving plants, but- productive success of Passiflora coccinea in Bra- terfly caterpillars, ants, and adult moths. The zil, where their presence increased fruit and seed fourth level of trophic interactions (moths “steal- production (Leal et al. 2006). ing” EFN at night) does not necessarily have sig- EFN production can be induced by herbivory. nificant influence on the tri-trophic interaction Feeding by larvae of Ceratomia catalpae (Boisdu- between plants, herbivores, and ants described by val) (Lepidoptera: Sphingiidae), increased sugars previous authors. Although in a study by Heil et in EFN of Catalpa bignonioides Walter (Bignoni- al. (2004) the number of EFN-feeding flies corre- aceae) and led to up to ten-fold increase in atten- lated negatively with the number of ants attend- dance by ants (Ness 2003). Similarly, feeding by ing the plants, the extent to which H. phaeoptera- Heliconius charithonius L. and Agraulis vanillae lis may have similar effect on ants attending P. in- L. (Lepidoptera: Nymphalidae) (which was very carnata is unclear. heavy on the plants in my observation) might be a I thank George T. Austin (FLMNH) for identi- factor that induced nectar production in EFN of fying the moths. Passiflora, and contributed to attracting the moths. The latter supposition, however, needs to SUMMARY be experimentally tested. I must note though, that if not for the EFN and the numerous ants at- Feeding on extrafloral nectar of Passiflora in- tracted to them during the day, H. charithonius carnata by tropical sod webworm moths, Herpeto- and A. vanillae larvae would have completely de- gramma phaeopteralis, in Florida is reported and voured the plants, as happens to the plants when ecological implications of this observation are dis- cussed. REFERENCES CITED ADJEI-MAAFO, I. K., AND L. T. WILSON. 1983. Association of cotton nectar production with Heliothis punctigera (Lepidoptera: Noctuidae) oviposition. Environ. Ento- mol. 12: 1166-1170. APPLE, J. L., AND D. H. FEENER, JR. 2001. Ant visitation of extrafloral nectaries of Passiflora: the effects of nectary attributes and ant behavior on patterns in facultative ant-plant mutualisms. Oecologia 127: 409-416. ATANASSOV, A., AND P. W. SHEARER. 2005. Peach extra- floral nectar impacts life span and reproduction of adult Grapholita molesta (Busck) (Lepidoptera: Tor- tricidae). J. Agric. and Urban Entomol. 22: 41-47. BEACH, R. M., J. W. TODD, AND S. H. BAKER. 1985. Nec- taried and nectariless cotton cultivars as nectar sources for the adult soybean looper. J. Entomol. Sci. 20: 233-236. CALLAGHAN, C. J. 1992. Biology of epiphyll feeding but- terflies in a Nigerian cola forest (Lycaenidae: Lipten- inae). J. Lepidopterists’ Soc. 46: 203-214. Fig. 2. Florida carpenter ants, Camponotus florida- GENTRY, G. 2003. Multiple parasitoid visitors to the extra- nus (Buckley), feeding on extrafloral nectary of Passi- floral nectaries of Solanum adherens. Is S. adherens an flora incarnata L. insectary plant? Basic and Appl. Ecol. 4: 405-411. 138 Florida Entomologist 91(1) March 2008 HEIL, M., A. HILPERT, R. KRUEGER, AND K. E. LINSEN- tum) as a food source for parasitic wasps. Functional MAIR. 2004. Competition among visitors to extraflo- Ecol. 20: 67-74. ral nectaries as a source of ecological costs of an RUDGERS, J. A. 2004. Extrafloral nectar as a resource indirect defense. J. Trop. Ecol. 20: 201-208. mediating multispecies interactions. Ecology 85: KEELER, K. H. 1978. Insects feeding at extrafloral nec- 1495-1502. taries of Ipomoea carnea (Convolvulaceae). Entomol. SHEARER, P. W., AND A. ATANASSOV. 2004. Impact of News 89: 163-168. peach extrafloral nectar on key biological charac- LABEYRIE, E., L. PASCAL, J. DELABIE, J. ORIVEL, A. DE- teristics of Trichogramma minutum (Hymenoptera: JEAN, AND M. HOSSAERT-MCKEY. 2001. Protection of Trichogrammatidae). J. Econ. Entomol. 97: 789- Passiflora glandulosa (Passifloraceae) against her- 792. bivory: impact of ants exploiting extrafloral necta-
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