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Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tnah20 Christmas (Anoplognathus spp., Coleoptera: ) mistake peppercorn trees for eucalypts Martin J. Steinbauer & Wolfgang J. Wanjura Published online: 06 Dec 2010.

To cite this article: Martin J. Steinbauer & Wolfgang J. Wanjura (2002) Christmas beetles (Anoplognathus spp., Coleoptera: Scarabaeidae) mistake peppercorn trees for eucalypts, Journal of Natural History, 36:1, 119-125, DOI: 10.1080/00222930010022917 To link to this article: http://dx.doi.org/10.1080/00222930010022917

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Christmas beetles (Anoplognathus spp., Coleoptera: Scarabaeidae) mistake peppercorn trees for eucalypts

MARTIN J. STEINBAUER² ³ and WOLFGANG J. WANJURA³ ² Co-operative Research Centre for Sustainable Production Forestry ³ CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia

(Accepted 20 July 2000)

Two of Christmas (Anoplognathus montanus Macleay and A. pallidic- ollis Blanchard (Coleoptera: Scarabaeidae)) are recorded feeding upon an exotic plant species (Schinus molle) belonging to a plant family (Anacardiaceae) diŒerent from their preferred hosts (species of (Myrtaceae)). The presence of similar monoterpenes in both S. molle and Eucalyptus might explain why Christmas beetles exhibit aberrant feeding on the former species. Preferential selection by the beetles for S. molle over neighbouring potential eucalypt hosts could be related to the absence in S. molle of 1,8-cineole, a monoterpene found in high concentrations in eucalypts that are resistant to Christmas beetles, combined with the presence of attractant monoterpenes.

Keywords: Interfamily host-shift, novel hosts, host-speci® city, terpenoids, repellents versus attractants, south-east Australia.

Introduction The peppercorn tree, Schinus molle, is an exotic species from South America that has been widely planted throughout Australia (Backer and Bakhuizen van den Brink, 1965). The history of the introduction of S. molle into Australia is sketchy. Some of the earliest records concerning the species derive from Western Australia during the 1830± 1840s, while the species was ® rst recorded from South Australia in Downloaded by [University of Florida] at 07:41 14 November 2014 1858 (G. Keighery, personal communication). Hence, S. molle would appear to have been in Australia for approximately 150 years. The berries and leaves of S. molle contain terpenes that exhibit insecticidal, fungicidal and allelopathic properties (Morton, 1978; Bernhard et al., 1983; Carr et al., 1985; Dikshit et al., 1986; Guardiola et al., 1990; Steinbauer, 1995; Wimalaratne et al., 1996; Gebre-Amlak and Azerefegne, 1999). S. molle continues to be planted throughout parts of Australia as an ornamental and shade tree by virtue of its drought tolerance, ease of propagation and longevity (Lawrence, 1984). Beetles belonging to the Anoplognathus Leach are endemic to Australia and are typically active during the austral summer, hence, their common name

Journal of Natural History ISSN 0022-2933 print/ISSN 1464-5262 online Ñ 2002 Taylor & Francis Ltd http://www.tandf.co.uk/journals DOI: 10.1080/00222930010022917 120 M. J. Steinbauer and W. J. Wanjura

`Christmas beetles’ (Carne et al., 1974). The adults are considered host-speci® c for species of eucalypt (Carne, 1957, 1958). Edwards et al. (1993) reported that diŒeren- tial resistance of some species of eucalypt to Christmas beetle herbivory was correl- ated with the percentage of several individual terpene components, including 1,8-cineole. Similar correlations between reduced herbivory by Christmas and leaf beetles (Chrysomelidae) and elevated 1,8-cineole content have been suggested by Edwards et al. (1990), Li (1993) and Stone and Bacon (1994). This note details observations of aberrant feeding behaviour by two species of Christmas beetle which we consider could be explained by the chemical basis of host selection in species of Anoplognathus.

Results and observations In surveys at a number of locations in southern New South Wales during the summer of 1991/92, adult Anoplognathus were regularly seen feeding upon the leaves of S. molle (table 1; ® gure 1a). The numbers of beetles that were observed, together with the disparate locations from which the phenomenon was recorded, suggested that such behaviour was not uncommon. Moreover, beetles were observed feeding upon S. molle even when susceptible species of Eucalyptus were nearby and despite the apparent di culty beetles had grasping the small lea¯ ets of S. molle (® gure 1b). These observations suggest that beetles were not forced to feed upon S. molle through overpopulation. The composition of monoterpenes in S. molle exhibited much similarity to that recorded from susceptible eucalypts (table 2, compare Lawrence (1984) versus Boland et al. (1991)) particularly with respect to a-phellandrene. A notable diŒerence between the two plant genera is the lower total oil content of S. molle compared with many eucalypts. Another important diŒerence between S. molle and some species of eucalypt that are found in its vicinity is the absence of 1,8-cineole in the monoterpene pro® le of S. molle (table 2).

Discussion Although S. molle has become a common exotic in Australia, there are no records in the primary literature concerning the ± plant associations that have developed following introduction. In Florida a related species, S. terebinthifolius Raddi, has acquired a phytophagous insect fauna comprising 46 species since its introduction some 100 years earlier (Cassani, 1986; Cassani et al., 1989). The number of phyto-

Downloaded by [University of Florida] at 07:41 14 November 2014 phagous associated with S. terebinthifolius in its native Brazil has not been

Table 1. Anoplognathus spp. collected in 85-cm diameter frass traps beneath Schinus molle trees.

Date Location No. trees A. montanus A. pallidicollis

18 December 1991 Baldry cemetery 1 20 males, 7 females ± (32ß 52¾ S 148ß 30¾ E) 15 January 1992 Coolac tearooms 1 42 males, 14 females 2 females (34ß 56¾ S 148ß 10¾ E) 1 February 1992 `Rock® eld’, Boorowa 1 22 males, 7 females 1 male (34ß 27¾ S 148ß 44¾ E) 8 February 1992 Baldry cemetery 2 49 males, 17 females ± (32ß 52¾ S 148ß 30¾ E) Christmas beetles mistake peppercorn trees for eucalypts 121 Downloaded by [University of Florida] at 07:41 14 November 2014

Fig. 1. (a) Anoplognathus spp. on Schinus molle at `Rock® eld’ via Boorowa, NSW, January 1992. (b) Anoplognathus montanus feeding on Schinus molle at `Rock® eld’ via Boorowa, NSW, January 1992. 122 M. J. Steinbauer and W. J. Wanjura , ; d t e e c l e i y r a t o r a t s l 4 3 8 8 8 h s x . . . . . a c e a 0 1 0 1 0 t a o e e e r T n e e n p o w r s s e t e n v e l o l i a a t o e t a e o L 3 2 2 1 c t n . . . . i . o 0 3 7 3 0 l f d C 6 7 2 o e - o r 8 i e , w u t g 1 q a t e m r n o e l r i c f . t r s e n e d u n u p t e e t 3 2 p . s . s C y m 3 a 1 0 0 0 e l ß . y v a 0 s r d c r C 2 e a u o t h s Õ h E i t l e t f u r a o s a e e e s d w e n n e e i e h e s r t c r p e o e r d f t v e p s 9 1 4 n o a t . . . s e a 7 0 0 1 e 1 l l n l r d L 3 a o e e n i . u h s a w y d s P r i i e - s v o l e i l b t m l i r d o r p e n r m I p e m s a T . e s u ) l h n 3 e t e a i t 9 n g h i h 9 e c a t p i 1 n i 4 7 7 4 l S a . . . . ( o o w 7 5 0 7 1 C . n F l i m ) i a n . 3 s L a y 9 e t i r l e 9 n o a 1 e t s r ( p t a d r . s r r l e u o a t a e b o A w t n a n e l d e e r o E s h e d t d m h 6 7 6 n r . . . n t i f a n a 7 0 1 0 8 l i o l o 2 1 2 h w d t e t s e d i h n n n E i P w r o l - i u t ) t a t i m u 9 e s o o r o r 5 f r s p n a . o ( n t m f e s y h o e k e e g h c C i a e n i p t r e 0 e 0 5 4 9 t . a . . . . n n w Downloaded by [University of Florida] at 07:41 14 November 2014 r a . 1 3 8 2 7 i e r o g 2 1 r P h o o t - e s u t d l a t p e a o b r e a i f l a k l m m f T e o d a r m e m n h l . a c o r n E s f s o g a d ) d a g d n e e b l e t a ) g s b c t c i n ) e i a a t n i l t e c l b s a p l s i ) t o e u l a n b t s i l i c c i o t n s s o p s c s e a p e u e t a . e r s n d s ( ( g c e e i E w l s a s p a a t a r e u e r r r e r s n e o g o o ( ( s t f e a d d c i o ² a a l n r a o o i n e i i t c c o e l l l s i i l l l o a p e F n n d e e i o e l o o ² m D c c c c m m m a e r t a . . . . . p l o o S S E E E E p f t Christmas beetles mistake peppercorn trees for eucalypts 123

fully documented; however, some 200 species have been reported from this plant (Medal et al., 1999). Given the present understanding of host selection in Anoplognathus species, we suggest that these beetles may recognize S. molle as a potential host. This recognition could be mediated through the presence of a subset of particular components or the overall terpenoid composition of the plant. The beetles may be particularly attracted to S. molle in preference to nearby susceptible eucalypts by virtue of the absence of 1,8-cineole and similarities in levels of a-phellandrene. This suggestion requires validation, however, Morrow (1977) did suggest that eucalypt-eating insects might ® nd many endemic myrtaceous species acceptable hosts by virtue of their similar terpene and polyphenol compositions. Moreover, we have based this suggestion only on the monoterpenes found in the oils of these plants. Schinus molle and eucalypts possess complex pro® les of compounds belonging to many other classes. The eŒect of these compounds on insects is not known. There is evidence of similar phenomena in other insect± plant systems. For example, Furth and Young (1988) reported that species of Blepharida Chevrolat (Chrysomelidae), which are normally host-speci® c for species of Rhus (Anacardiaceae), exhibit a feeding preference for the novel host S. terebinthifolius. This preference was considered to have arisen due to similarities in ¯ avonoid com- position between Rhus and S. terebinthifolius. In this instance similarities in ¯ avonoid composition are thought to be the result of the recent common ancestry of the plants (Furth and Young, 1988). Some of the insects known to feed on S. tereb- inthifolius in Florida also exhibit a preference for citrus (Medal et al., 1999). Citrus contains monoterpenes common to some found in S. terebinthifolius. A species of native Australian geometrid (Chlenias Guene e sp.) caused considerable defoliation of plantations of Pinus radiata (Madden and Bashford, 1977a, 1977b). Likewise, P. radiata contains monoterpenes common to some found in Eucalyptus. If plant species with comparable secondary compounds, whether by virtue of related lineage or not, are assessed similarly by phytophagous insects, then in Australia we might predict that S. molle should acquire a suite of otherwise eucalypt-speci® c phytophages. There is some evidence that supports this suggestion. For example, larvae of the endemic emperor gum ( eucalypti (Scott), : ) have often been collected on S. molle (Froggatt, 1907; Tillyard, 1926; Common, 1990).

Downloaded by [University of Florida] at 07:41 14 November 2014 Acknowledgements The occurrence of Anoplognathus species on S. molle was ® rst brought to the attention of Dr Penny Edwards in March 1991 by W. Semple (then NSW Soil Conservation Service, Orange). Greg Keighery (Department of Conservation and Land Management, Perth) provided information concerning the occurrence of S. molle in Australia. We wish to thank Drs Saul Cunningham (CSIRO Entomology, Canberra), Penny Edwards (CSIRO Entomology, Brisbane), Rob Floyd (CSIRO Entomology, Canberra) and Mamoru Matsuki (presently CRC for Sustainable Production Forestry, Hobart) for their comments on this note.

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