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Chilades Pandava Damage Among 85 Cycas Species in a Common

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Chilades Pandava Damage Among 85 Cycas Species in a Common HORTSCIENCE 47(12):1832–1836. 2012. Materials and Methods Three evaluators conducted this survey to Chilades pandava Damage among quantify the extent of damage to representa- tive field-planted Cycas species at Nong 85 Cycas Species in a Common Nooch Tropical Botanical Garden in Thai- land. The survey included 85 species growing Garden Setting in homogeneous abiotic and insect pressure conditions. Two arthropods exhibit chronic Thomas E. Marler1 pest pressures on the Cycas plants in this Western Pacific Tropical Research Center, College of Natural and Applied garden, Chilades pandava and Aulacaspis yasumatsui Takagi. The butterfly can feed Sciences, University of Guam, UOG Station, Mangilao, Guam 96923 on several tissue types, but for every sub- Anders J. Lindstrom¨ strate, it requires expanding tissue for ovipo- sitioning and caterpillar food. The various Nong Nooch Tropical Botanical Garden, 34/1 Sukhumvit Hgw, Najomtien, groups of Cycas species produce vegetative Sattahip, Chonburi 20250, Thailand flushes at varied times of the year, and some exotic Cycas taxa produce leaves throughout L. Irene Terry the year. These characteristics provide the Western Pacific Tropical Research Center, College of Natural and Applied resident butterfly population with a sustained Sciences, University of Guam, UOG Station, Mangilao, Guam 96923 supply of caterpillar food. Therefore, the pest pressures are relentless and constant through- Additional index words. Cycadaceae, leaf traits, plant–arthropod interactions, plant defense, out the year so that no matter when a Cycas volatiles has a flush of leaves, it is subjected to adult Abstract. The extent of Chilades pandava Horsfield herbivory among 85 Cycadaceae butterfly ovipositioning. A. yasumatsui, the species was determined by three evaluators in a common garden setting in Thailand to second resident Cycas pest in our garden, is identify patterns that may improve horticultural and conservation management an armored scale that infests any organ that practices. The significant differences in herbivory damage from this invasive lepidop- presents live exposed surfaces. Unidentified teran pest ranged 8.7-fold among the species. Phylogenetic sections of this monogeneric biological control organisms keep the wide- cycad family did not correspond to the relative differences among the species, and spread A. yasumatsui infestations from irrupt- country of nativity was also not informative for this purpose. We suggest the Cycas L. ing. Confusing the plant damage imposed by species that share native habitat with this butterfly or the closely related Theclinesthes these two arthropods is not possible; therefore, onycha Hewitson are among the least damaged taxa when they are comingled with other identifying butterfly damage is unambiguous. Cycas species in a common landscape. Grouping the most damaged Cycas species Individual plants were ranked for butterfly together in a managed landscape may reduce costs associated with plant protection. The damage in Mar. 2012. We assigned 0 for inclusion of non-native Cycas plants in gardens nearby native Cycas habitats carries the a plant exhibiting no visible damage and 10 potential of disrupting the delicate specialist relationship that native butterfly popula- for a plant with butterfly damage on every tions have with host Cycas species. leaf. A total of 673 plants in 85 species, representing all five Cycas sections, were evaluated. A Poisson regression model was Invasive arthropod herbivores comprise Nong Nooch Tropical Botanical Garden in fitted on log-transformed damage evaluation one of the greatest threats to cycad conserva- Thailand as part of our efforts to conserve this scores using SAS GLIMMIX procedure in tion both in situ and ex situ. The lepidopteran endangered cycad. In this setting where the Version 9.3 (SAS Institute Inc., Cary, NC). A Chilades pandava (Plains Cupid Butterfly, butterfly is native, herbivory is highly hetero- Type III test of fixed main effects was used Fig. 1B) is a specialist insect native to southern geneous among comingled Cycas species. to determine significance of evaluators and Asia that invaded Guam in recent years (Moore Some species like Cycas revoluta Thunb. species. The damage differences among the et al., 2005), and it is among the invasive (Fig. 1C) and C. micronesica (Fig. 1E) exhibit species were determined using pairwise com- species that are collectively threatening the extensive, unsightly damage, whereas other parison of least squares means on log scale. endemic Cycas micronesica K.D. Hill (Marler species exhibit minimal damage (Figs. 1D and and Lawrence, 2012). This insect depends on a 1F). Management decisions for this situation Results and Discussion sustained close relationship with native Cycas would benefit from understanding how the within its indigenous range. It causes ex- butterfly relates to its native host Cycas The mean damage score among species tensive damage to Cycas populations within species in natural habitats and how it relates was significant based on Type III test (P # habitats in its invasive range like Guam. These as a pest to novel Cycas host species in hor- 0.0001) after removing the evaluator differ- factors engender a horticultural conflict be- ticultural settings. ences. The log-transformed mean damage tween the need to protect Cycas species from Most of the 107 described Cycas species scores were back-transformed and presented damage within garden settings and within the (Osborne et al., 2012) are being conserved in asaLSMEANstable(Table1).Butterfly butterfly’s invasive range and the arguable the ex situ cycad germplasm collections in damage of the 85 Cycas speciesinthiscommon need to conserve its dependence on Cycas Nong Nooch Tropical Botanical Garden. Pes- garden setting exhibited an overall mean of plants in situ. ticide protection of susceptible Cycas taxa 2.093, and mean damage ranged 8.7-fold An extensive ex situ germplasm collection resulting from this pest represents one of the among the species (0.5947 to 5.1502). The of Guam’s C. micronesica was established at greatest costs for success of these conservation damage scores for these species partitioned into efforts. We used this germplasm collection 18 overlapping groups. Clearly, whether Cycas growing in a common garden setting to de- plants succumb to or resist damage by chronic termine relative butterfly herbivory at the butterfly pressure is governed by differences Received for publication 22 Aug. 2012. Accepted species level. Our objective was to then mine expressed at the species level. for publication 10 Oct. 2012. Support provided by U.S. Forest Service Project these data for linkages between resistance to Phylogeny. Cycas micronesica, C. petraea No. 10-DG-11059702-095. damage and group traits that may reduce A. Lindstr. & K.D. Hill, C. revoluta, We thank George Fernandez for statistical analyses. uncertainty in predictions of and expose C. seemannii A. Braun, C. tansachana K.D. 1To whom reprint requests should be addressed; mechanisms that explain disparity in butterfly Hill & S.L. Yang, and C. thouarsii R.Br. ex e-mail [email protected]. damage. Gaudich. emerged as the most damaged 1832 HORTSCIENCE VOL. 47(12) DECEMBER 2012 Fig. 1. Relationship of Cycas species with the specialist herbivore Chilades pandava.(A) Extent of herbivory damage for 85 Cycas species sorted into the five Cycadaceae sections, mean ± SD.(B) Female butterfly ovipositioning eggs on Cycas revoluta leaf. (C) C. revoluta plant showing typical extensive butterfly damage to leaflets. Damage score 9. (D) C. pachypoda plant with undamaged leaves. Damage score 0. (E) Production beds of the susceptible C. micronesica showing plant mortality and heterogeneous growth. (F) Production beds of the resistant C. chamaoensis showing homogeneous growth. species and C. aculeata K.D. Hill & T.H. variation within each of these sections. Asior- 1983). The mean for the eight species from Nguyeˆn, C. circinalis L., C. condaoensis K.D. ientales exhibited a mean score that was this subsection in our survey was 3.1281, and Hill & S.L. Yang, C. pachypoda K.D. Hill, greater than the overall survey mean, con- half of the Cycas species ranked as most C. pruinosa Maconochie, C. taitungensis C.F. tained C. taitungensis in the least damaged damaged (Table 1) were from this small Shen, K.D. Hill, C.H. Tsou & C.J. Chen, C. group, and C. revoluta in the most damaged subsection. An interesting example that un- tuckeri K.D. Hill, and C. wadei Merrill group. These major Cycas sections do not derscores these observations is the Philippine exhibited the least amount of damage among adequately inform our understanding of what island of Culion, where the strict endemic C. the 85 species (Table 1). Exploring character- underlies the differences in butterfly damage wadei (Wadeanae section without floating istics that explain these two extreme groups among the 85 species that we observed. seeds) and the widespread C. edentata de may establish a foundation for understanding Our results do not broadly correspond with Laub. (Rumphiae subsection within Cycas the controlling mechanisms of the disparity in phylogeny. However, the non-significant lower section with floating seeds) live on the main damage within the genus. Two obvious group damage scores in this garden survey for the island and its barrier islands (Lindstr¨om et al., traits to explore for this purpose are phyloge- species in the Wadeanae section indicate that 2008). Damage to C. edentata was three times netic sections and native biogeographic range. direct bioassays
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