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Attraction of Phyllopertha Horticola (Coleoptera, Scarabaeidae) Males to Fragrance Components of Ophrys Flowers (Orchidaceae, Section Fuciflorae)

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Attraction of Phyllopertha Horticola (Coleoptera, Scarabaeidae) Males to Fragrance Components of Ophrys Flowers (Orchidaceae, Section Fuciflorae) Attraction of Phyllopertha horticola (Coleoptera, Scarabaeidae) males to fragrance components of Ophrys flowers (Orchidaceae, section Fuciflorae) ANNA.KARIN BORG-KARLSON Borg-Karlson, A.-K.: Attraction of Phy llopertha horticola (Coleoptera, Scarabaeidae) males io fragrance components of Ophrys flowers (Orchidaceae .section Fuciflorae). ITriidgrirdsbor- rens hanar (Phyllopertha horticola) attraheras till komponenter i blomdoften frintre Ophrys arter (Orchidaceae . sektion Fuciflorae).1 - Ent. Tidskr. t09: 105-109. Umei, Sweden 1989. ISSN 001-l-886x. The North-European scarabaeid beetle Phyllopertha horicola has been obscrvcd to visit and exhibit pseudocopulatory behaviour on the nectarless flowers of C)pftrys species (section Fuci /orae). To evaluate the role of chemical signals in the beetles' attraction response io the flow- ers and to determine the active compounds involved, males from a swarming Population on Oland (Sweden) were tested for their attraction to different odour samples: hexane exlracts of Ophrys flower labella, hexane extracts of male and female beetles, and different blends of compounds identified in Opftrys flowcr fragrances. Male beetles were most strongly attracted to a blend of short-chained aliphatic alcohols present in extracts of O. hoktcericea labella. A.-K. Borg-Karkon, Ecological Research Sration of IJppsala Ilniversity, Olands Skogshy 6280, 5-386 00 Fiirjestaden, Sweden and Departmen of Orgqnic Chemistri', The Royal Institute oI 'feclnology. S-100 41, Stockhobn, Sweden. lntroduclion The scarabaeid bcetle Phyllopertha horticola (L.) arc attracted/excited primarily by odour, or has repeatedly been observed to visit flowers of whethcr visual and tactile cues are also necessary Ophrys L., sectioll Fuciflorae Rchb.f. (Kullen- to elicit the complete sequenced behavioural re- berg 1973, Warncke & Kullenberg 1984. Borg- sponses leading to pollination by pseudocopula- Karlson 1987). The often white or cerise sepals of tion. Oplrrys flowers are similar in coloration to th€ The purpose of this study was to experimentally beetles' food plants Cisrr.r (Cistaceae) and Crata- examine, using behavioural tests, the role ofolfac- egus (Rosaceae). It has been suggested that these tory stimuli in the pollination relationship betwe- colours may act as visual cues encouraging both en the beetle Phyllopertha horticola and the spe- male and female beetles to visit and forage on cies Ophrys holocericea, O. scolopax Cav. and O. tN Ophr,r,s flowcrs and thcre bv incidentally pollinate t ent h re d i nif e r a illd. them (Ferlan in lit. to Kullenberg. Kullenbcrg 1961). However, recent observations (Forster, Materials and methods Paulus, in lit. to Kullenberg 7976,7982,1983) re- vealed that P. horticola malesbehave as ifsexually The study took place on Oland, one of the large is- stimulated when visiting O. holocericea (Btrm. lands in the Baltic Sea closc to the mainland of fil.) Greuter l: O. fuciflora (Cranz) Moenckl southcrn Sweden. Odour attraction tcsts were flowers. Other scarabaeid beetles, such as Tropi- performed from end May to mid Junc 1983-85 on nota squalida Scop. and Oxythyrea funesta Poda, a field population of beetles inhabiting the border also visit and pollinate various Opftrys species in of a grass-lined gravel road surrounded by hazel- the section Fuciflorae (Ktsllenberg 1961). bushes (Corylus avellana L.). It has not been established whether the beetles To test the beetles attraction to different 106 Anns- Karitt l)org-Korlson tanoic acid 2. nonanoic acid 20. citral4, geraniol 1, citronellol 4. geranylacetone 4. The compositions of the floral blends were bascd on GC-MS (gas-chromatography mass spectrometry) analyses madc earlicr (Kullenberg ct al. 1984, Borg-Karlson 1987). The lest samples were applicd to dummies- picces of black velvet (5 x l0 mm) placed on insect h i.+o 66p. pins and covercd with green tullc. Abovc the tulle was attached an odourlcss piece of velvet to in- crease the visual and tactile stimuli of the dummy. Samples were tested one bv one in random order, for 5 minutes each, and in +10 repllcates. Re- sponses of beetles, within 0.2 m from the dummv, Fig. l. The odour-guidcd rcsponses of Pryllopertha hor- were noted. Results were evaluated statistically ricola male beetles are divided into four behaviour tvpcs using the chi-square test for 2 x 2 contingency ta- based on increasing degrees of attraction/excitation: ( l) blcs. For each test sample, the sum of behaviour p : uninlluenced passage, (2) q.i. : quick inspection, types (h.i. (3) h. i. : hovering inspection and w.i. = walking inspec- 3 and 4 [hovering ), walking-inspection tion, (1) h.i.-a.cop. : hovering inspection followed by (w.i.), and alighting rcsponscs on the dummy "copulation attempt" or w,i. q,cop. : walking inspec- (h.i.-a.cop., w.i.-a.cop.)], i.c. thc strongly at- tion folkrwed bv "attempted copulation", The insects tracted beetles (Fig. l), was dividcd by the total are regar(lcd as strr',nglv attracted in the behaviour types 3 and 4. number of responses (N). Portions of thc be- havioural sequences were filmed using a Paillard Det dottinducerade beteendet som trddgirdsborrehanar 16 utfdr vid attrappen Ar uppdelat i fyra olika typer med sti- Bolex mm film camera. gandc attraktivitctsgrad: (l) 2 : passage, (2) S.i. = Voucher beetle specimens are deposited at the snabb inspektion, (.3) n.i. = sidsvingande eller r,r,.i. = Ecological Research Station of Uppsala Univer- krvpandc inspcktittn. (1) h.i.--a.cop. eller w.i.-a.cop. = sity on Oland. Taxonomy and nomenclature of sidsvlngandc cllcr krypande inspektion som slutar med fiirsitk till kopulation pe artrappen. Insekterna iir starkt Ophrys is based on Nelson (1962), sce also Borg- attraherade om de reagerar med beteendetyp 3 cllcr 4. Karlson ( 1987) and rcfcrcnces therein. Results odours, eight different samples were presented to heetles within the confines of the swarm. Extracts In the mornings, male heetles started to walk were studied of Ophrys scolopax and O. lenth- whcn ambient temperaturc rcachcd 13"C (at 1.5 m rediniferu labella, and of female and inale /'ftyl- above ground) and to swarm at lzl-15"C. Swarm- lopertha horticola bodies. Fresh Oplrrys labella ing usually lasted for about three hours. Female were extracted in redistilled hexane (5 labclla/ml beetles were never seen swarming, bui were fre- hexane, Merck) for three days at 4'C and l-2 days quently observed feeding on leaves and grasses in old P. hortitolu bcctles were extracted in redistil- the vicinity. led pe ntanc (2 whole bodies/ml pentane, Merck) Only males were attracted to the dummies. The for 24 hours at 20'C. bchavioural rcsponses of malc bectles. both flying The following three authentic blends of com- and walking, to the dummies werc classified into 4 pounds identified in scents of Opftrys flowers were typcs as follows, listed in ordcr of increasing at- studied (volumes in pl; each blend diluted in 1.9 traction/excitation (Fig. 1). m.l hexane): Alcohol blend: l.hexanol 2, cis-3- 1. p passage: hexcn-l-ol 2. 1-octanol 10, l-hexadecanol l0; = uninfluenced by-passing without de- creasing flight speed or changing flight direction. Acid blend: pcntanoic acid 2, hexanoic acid 4, 2. q.i. = quick inspection: flight direction changed to- hcptanoic acid 10, octanoic acid 2, nonanoic acid wards dummy, sometimcs rcsulting in touch. 20; Ophrys tenthredinifera extruct blendi l-hexa- 3. r.i. : hovering inspection: mcandcring flight towards the odour source. hovering in front of it, sometimes fly- nol 2, ci.r-3-hexen- l -ol 2, 1-octanol 10, l-hexade- ing around it, always facing tho dummy. w.i. = walking canol 10, 6-methyl-5-hepten-2-one 6. pentanoic inspection: walking on ground (x in vcgetation towards acid 2, hexanoic acid 4. heptanoic acid l(). oc- clumml but without touch. Attractiort oJ Phyllopertha nales to Ophrys flowers 107 Tab. I . Respon sc s r tf P htlloperrha hortico,/a males towards each of the eight samples tested; f( 3 + 4) = the proportion of attracted beetles with behaviour iype 3 and 4 (e.g., h. i., w.i., h.i.-a.cop., w.i.-a.cop.) in relation to all beetles (within 0.2 m of the test dummy) for a given sample; f(4) = the proportion of attracted beetles with behaviour type ,[l N = thc total number of beetles observed for a given test sample; n = the number of replicates for each sample. Trddgirdsborrens betqcndcsvar gentemot 8 testamnen: f(3+4) : proportionen attraherade hanar med beteendetyP -j eller 4, jiimfort med alla hanar inom 0.2 m fran rest-attrappen; f(4) = proportionen attraherade hanar med beteende tr"p,+: N = totala antalet observerade skalbaggar; n = antal testtillfiillen frir varje iimne. Sample f(3+4) (4) - Alcohol blend 0.36* 0.19 64 10 O p hrys sc o lo pax labella extract 0.25- 0.10 40 8 Phyllopertha horticold nrale body extract 0.14 0.09 58 l0 P, horticolalemale body cxtract 0.13.* 0.06 70 8 O. tenthredinifera extract blcnd 0.12 0.10 50 l0 O. tenthre dinifero labclla extract 0.08 0.00 25 4 Acid blend 0.06 0.02 54 9 Hexane blank 0.05 0.02 56 10 ' p<lr Ill5) 'iSnrlicant!.l difl.r.nr lronr tllt hlanl lX:. '- \ignrfrcanlh difterenr tnnn rhc Allohrn bbnd (X:. p<0.05) 4. h .i.--a. cop. - hovering inspection followed by descent Discussion and walking on dummy.'fhe beetle performs move- ments that appear to represent a copulation attempt, Rcsults of the present odour-attraction Iests indi- tt.i.-a.cop. : walking inspection followed by contact cate that P. horticolq malcs are attracted to vol- with the durnmv and seeminglv attempted copulation. atile compounds prescnt in flower fragrances of Extracts of O. scolopax labella, which are domi the studied Op&rys species. White and cerise col- naled by the alcohols l-hexanol and 3-hexen-1-ol our cues were excluded in the experiments and the (Borg-Karlson 1987), proved to be highly attrac- beetles seemed to be only slightly, if at all, af- tive to P.
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