Skip to main content
SpringerLink
Log in

For sex and more: attraction of the tephritid parasitoid Psyttalia concolor (Hymenoptera: Braconidae) to male sex pheromone of the olive fruit fly, Bactrocera oleae

  • Original Paper
  • Published:
Journal of Pest Science Aims and scope Submit manuscript

Abstract

The braconid parasitoid Psyttalia concolor foraging for larvae of Bactrocera oleae (Diptera: Tephritidae) uses olfactory cues from the larval microhabitat. However, it could rely on the sex pheromones of adults of its host, since B. oleae mating leks usually occur on olive plants and a high number of infested fruits can be easily located in their close proximity. We quantified the behavioural responses of P. concolor towards different doses of the two major sex pheromone components of B. oleae, the female-borne 1,7-dioxaspiro[5.5]undecane (DSU) and the male-borne (Z)-9-tricosene (ZT). Results showed that P. concolor females and males were attracted only to the highest amounts of ZT (10 B. oleae male equivalents), regardless of the mating status. Wasps remained longer on the surfaces treated with high ZT amounts over the control. P. concolor females could exploit the ZT produced by lekking olive fruit fly males as a foraging kairomone. This could allow them to rapidly track B. oleae-infested olive plants, thus raising their chances to locate host larvae during their lifespan. On the other hand, the perception of ZT by P. concolor males could raise their chances to locate receptive females emerging from parasitised B. oleae pupae or also ovipositing in olive fruits. This study increases the knowledge of the reproductive biology of P. concolor, highlighting a case of chemical espionage in which a sex pheromone component from adult stages of the host is exploited both as a foraging kairomone by female parasitoids and as a sexual kairomone by male wasps.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Log in via an institution

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Institutional subscriptions

Fig. 1

Similar content being viewed by others

References

  • Aldrich JR, Kochansky JP, Abrams CB (1984) Attractant for beneficial insect and its parasitoids: pheromone of the predatory spined soldier bug, Podisus maculiventris (Hemiptera: Pentatomidae). Environ Entomol 13:1031–1306

    CAS  Google Scholar 

  • Allen GR (1995) The calling behaviour and spatial distribution of male bushcrickets (Sciarasaga quadrata) and their relationship to parasitism by acoustically orienting tachinid flies. Ecol Entomol 20:303–310

    Article  Google Scholar 

  • Arakaki N, Wakamura S (2000) Bridge in time and space for an egg parasitoid—kairomonal use of trace amount of sex pheromone adsorbed on egg mass scale hair of the tussock moth, Euproctis taiwana (Shiraki) (Lepidoptera: Lymantriidae), by egg parasitoid, Telenomus euproctidis Wilcox (Hymenoptera: Scelionidae), for host location. Entomol Sci 3:25–31

    Google Scholar 

  • Arakaki N, Wakamura S, Yasuda T, Yamagishi K (1997) Two regional strains of a phoretic egg parasitoid, Telenomus euproctidis (Hymenoptera: Scelionidae), that use different sex pheromones of two allopatric tussock moth species as kairomones. J Chem Ecol 23:153–161

    Article  CAS  Google Scholar 

  • Arakaki N, Yamazawa H, Wakamura S (2011) The egg parasitoid Telenomus euproctidis (Hymenoptera: Scelionidae) uses sex pheromone released by immobile female tussock moth Orgyia postica (Lepidoptera: Lymantriidae) as kairomone. Appl Entomol Zool 46:195–200

    Article  CAS  Google Scholar 

  • Baker R, Herbert RH, Howse PE, Jones OT, Francke W, Reith W (1980) Identification and synthesis of the major sex pheromone of the olive fly, Dacus oleae. J Chem Soc Comm 1:52–54

    Article  Google Scholar 

  • Beale MH, Birkett MA, Bruce TJA, Chamberlain K, Field LM, Huttly AK, Martin JL, Parker R, Phillips AL, Pickett JA, Prosser IA, Shewry PR, Smart LE, Wadhams LJ, Woodcock CM, Zhang Y (2006) Aphid alarm pheromone produced by transgenic plants affects aphid and parasitoid behavior. Proc Natl Acad Sci Am 103:10509–10513

    Article  CAS  Google Scholar 

  • Benelli G (2014) Aggressive behavior and territoriality in the olive fruit fly, Bactrocera oleae (Rossi) (Diptera: Tephritidae): role of residence and time of day. J Ins Behav 27:145–161

    Article  Google Scholar 

  • Benelli G, Canale A (2012a) Learning of visual cues in the fruit fly parasitoid Psyttalia concolor (Szépligeti) (Hymenoptera: Braconidae). Biocontrol 57:767–777

    Article  Google Scholar 

  • Benelli G, Canale A (2012b) Do Psyttalia concolor (Hymenoptera: Braconidae) males gain in mating competitiveness from being courted by other males while still young? Entomol Sci 15:257–260

    Article  Google Scholar 

  • Benelli G, Canale A (2013a) Male-male sexual behavior in the parasitic wasp Psyttalia concolor. J Ins Sci 13:25

    Google Scholar 

  • Benelli G, Canale A (2013b) Do tephritid-induced fruit volatiles attract males of the fruit flies parasitoid Psyttalia concolor (Szépligeti) (Hymenoptera: Braconidae)? Chemoecology 23:191–199

    Article  CAS  Google Scholar 

  • Benelli G, Bonsignori G, Stefanini C, Canale A (2012) Courtship and mating behaviour in the fruit fly parasitoid Psyttalia concolor (Szépligeti) (Hymenoptera: Braconidae): the role of wing fanning. J Pest Sci 85:55–63

    Article  Google Scholar 

  • Benelli G, Gennari G, Francini A, Canale A (2013a) Longevity costs of same-sex interactions: first evidence from a parasitic wasp. Invert Biol 132:156–162

    Article  Google Scholar 

  • Benelli G, Gennari G, Canale A (2013b) Host discrimination ability in the tephritid parasitoid Psyttalia concolor (Hymenoptera: Braconidae). J Pest Sci 86:245–251

    Article  Google Scholar 

  • Benelli G, Revadi S, Carpita A, Giunti G, Raspi A, Anfora G, Canale A (2013c) Behavioral and electrophysiological responses of the parasitic wasp Psyttalia concolor (Szépligeti) (Hymenoptera: Braconidae) to Ceratitis capitata-induced fruit volatiles. Biol Control 64:116–124

    Article  CAS  Google Scholar 

  • Benelli G, Bonsignori G, Stefanini C, Raspi A, Canale A (2013d) The production of female sex pheromone in Bactrocera oleae (Rossi) young males does not influence their mating chances. Entomol Sci 16:47–53

    Article  Google Scholar 

  • Benelli G, Pacini N, Conti B, Canale A (2013e) Following a scented beetle: larval faeces as a key olfactory cue in host location of Stegobium paniceum (Coleoptera: Anobiidae) by Lariophagus distinguendus (Hymenoptera: Pteromalidae). Chemoecology 23:129–136

    Article  Google Scholar 

  • Benelli G, Daane KM, Canale A, Niu CY, Messing RH, Vargas RI (2014) Sexual communication and related behaviours in Tephritidae: current knowledge and potential applications for Integrated Pest Management. J Pest Sci. doi:10.1007/s10340-014-0577-3

    Google Scholar 

  • Borges M, Costa MLM, Sujii ER, Cavalcanti MG, Redígolo GF, Resck IS, Vilela EF (1999) Semiochemical and physical stimuli involved in host recognition by Telenomus podisi (Hymenoptera: Scelionidae) toward Euschistus heros (Heteroptera: Pentatomidae). Physiol Entomol 24:227–233

    Article  Google Scholar 

  • Bruce MJ, Herberstein ME, Elgar MA (2001) Signalling conflict between prey and predator attraction. J Evol Biol 14:786–794

    Article  Google Scholar 

  • Bruni R, Sant’Ana J, Aldrich JR, Bin F (2000) Influence of host pheromone on egg parasitism by scelionid wasp: comparison of phoretic and nonphoretic parasitoids. J Ins Behav 12:165–173

    Article  Google Scholar 

  • Canale A, Benelli G (2012) Impact of mass-rearing on the host-seeking behaviour and parasitism by the fruit fly parasitoid Psyttalia concolor (Szépligeti) (Hymenoptera: Braconidae). J Pest Sci 85:65–74

    Article  Google Scholar 

  • Canale A, Carpita A, Conti B, Canovai R, Raspi A (2012) Effect of age on 1,7-dioxaspiro-[5.5]-undecane production in both sexes of olive fruit fly, Bactrocera oleae (Rossi) (Diptera Tephritidae). IOBC Bull 72:219–225

    Google Scholar 

  • Canale A, Benelli G, Lucchi A (2013a) Female-borne cues affecting Psyttalia concolor (Hymenoptera, Braconidae) male behavior during courtship and mating. Insect Sci 20:379–384

    Article  PubMed  Google Scholar 

  • Canale A, Benelli G, Lanzo F, Giannotti P, Mazzoni V, Lucchi A (2013b) The courtship song of fanning males in the fruit fly parasitoid Psyttalia concolor (Szépligeti) (Hymenoptera: Braconidae). Bulletin Entomol Res 103:303–309

    Article  CAS  Google Scholar 

  • Canale A, Germinara SG, Carpita A, Benelli G, Bonsignori G, Stefanini C, Raspi A, Rotundo G (2013c) Behavioural and electrophysiological responses of the olive fruit fly, Bactrocera oleae (Rossi) (Diptera: Tephritidae), to male- and female-borne sex attractants. Chemoecology 23:155–164

    Article  CAS  Google Scholar 

  • Carpita A, Canale A, Raffaelli A, Saba A, Benelli G, Raspi A (2012) (Z)-9-tricosene identified in rectal gland extracts of Bactrocera oleae males: first evidence of a male-produced female attractant in olive fruit fly. Naturwissenschaften 99:77–81

    Article  CAS  PubMed  Google Scholar 

  • Colazza S, Salerno G, Wajnberg E (1999) Volatile and contact chemicals released by Nezara viridula (Heteroptera: Pentatomidae) have a kairomonal effect on the egg parasitoid Trissolcus basalis (Hymenoptera: Scelionidae). Biol Control 16:310–317

    Article  Google Scholar 

  • Conti E, Salerno G, Bin F, Williams HJ, Vinson SB (2003) Chemical cues from Murgantia histrionica eliciting host location and recognition in the egg parasitoid Trissolcus brochymenae. J Chem Ecol 29:115–130

    Article  CAS  PubMed  Google Scholar 

  • Dweck HKM, Svensson GP, Gündüz EA, Anderbrant O (2010) Kairomonal response of the parasitoid, Bracon hebetor Say, to the male-produced Sex pheromone of its Host, the greater waxmoth, Galleria mellonella (L.). J Chem Ecol 36:171–178

    Article  CAS  PubMed  Google Scholar 

  • El-Sayed AM (2014) The pherobase: database of insect pheromones and semiochemicals, http://pherobase.com

  • Endler JA (1978) A predator’s view of animal color patters. Evol Biol 11:319–364

    Article  Google Scholar 

  • Endler JA (1991) Variation in the appearance of guppy color patterns to guppies and their predators under different visual conditions. Visual Res 31:587–608

    CAS  Google Scholar 

  • Fatouros NE, Huigens ME, van Loon JJA, Dicke M, Hilker M (2005) Chemical communication: butterfly anti-aphrodisiac lures parasitic wasps. Nature 433:704

    Article  CAS  PubMed  Google Scholar 

  • Fatouros NE, Dicke M, Mumm R, Meiners T, Hilker M (2008) Foraging behavior of egg parasitoids exploiting chemical information. Behav Ecol 19:677–689

    Article  Google Scholar 

  • Feener DH, Jacobs LF, Schmidt JO (1996) Specialized parasitoid attracted to a pheromone of ants. Anim Behav 51:61–66

    Article  Google Scholar 

  • Fiaboe MK, Smith H, Borgeweister C, Chabi-Olaye A, Gounou S, Schulthess F (2003) Sesamia calamistis calling behavior and its role in host finding of egg parasitoids Telenomus busseolae, Telenomus isis, and Lathromeris ovicida. J Chem Ecol 29:921–929

    Article  CAS  PubMed  Google Scholar 

  • Fisher IG, Tyman JHP (1998) The synthesis of long chain dialkylalkynes, dialkyldiynes and their hydrogenation to monoenes and dienes. Synth Comm 28:1323–1338

    Article  CAS  Google Scholar 

  • Frenoy C, Durier C, Hawlitzky N (1992) Effect of kairomones from egg and female adult stages of Ostrinia nubilalis (Hübner) (Lepidoptera: Pyralidae) on Trichogramma brassicae Bezdenko (Hymenoptera, Trichogrammatidae) female kinesis. J Chem Ecol 18:761–763

    Article  CAS  PubMed  Google Scholar 

  • Glinwood RT, Powell W, Tripathi CPM (1998) Increased parasitization of aphids on trap plants alongside vials releasing synthetic aphid sex pheromone and effective range of the pheromone. Biocontrol Sci Technol 8:607–614

    Article  Google Scholar 

  • Glinwood RT, Du YJ, Powell W (1999) Responses to aphid sex pheromones by the pea aphid parasitoids Aphidius ervi and Aphidius eadyi. Entomol Exp Appl 92:227–232

    Article  Google Scholar 

  • Hardie J, Nottingham SF, Powell W, Wadhams LJ (1991) Synthetic aphid sex pheromone lures female aphid parasitoids. Entomol Exp Appl 61:97–99

    Article  CAS  Google Scholar 

  • Hardie J, Hick AJ, Höller C, Mann J, Merritt L, Nottingham SF, Powell W, Wadhams LJ, Witthinrich J, Wright AF (1994) The responses of Praon spp. parasitoids to aphid sex pheromone components in the field. Entomol Exp Appl 71:95–99

    Article  CAS  Google Scholar 

  • Hedlund K, Vet LEM, Dicke M (1996) Generalist and specialist parasitoid strategies of using odours of adult drosophilid flies when searching for larval hosts. Oikos 77:390–398

    Article  Google Scholar 

  • Hilker M, Blaeske V, Kobs C, Dippel C (2000) Kairomonal effects of sawfly sex pheromones on egg parasitoids. J Chem Ecol 26:2591–2601

    Article  CAS  Google Scholar 

  • Huigens ME, Fatouros NE (2013) A hitch-hiker’s guide to parasitism: the chemical ecology of phoretic insect parasitoids. In: Wajnberg E, Colazza S (eds) Chemical ecology of insect parasitoids. John Wiley & Sons, Ldt., Chichester, p 86–111

  • Huigens ME, Pashalidou FG, Qian MH, Bukovinszky T, Smid HM, van Loon JJ, Dicke M, Fatouros NE (2009) Hitch-hiking parasitic wasp learns to exploit butterfly antiaphrodisiac. Proc Natl Acad Sci USA 106:820–825

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Huigens ME, Woelke JB, Pashalidou FG, Bukovinszky T, Smid HM, Fatouros NE (2010) Chemical espionage on species-specific butterfly anti-aphrodisiacs by hitchhiking Trichogramma wasps. Behav Ecol 21:470–478

    Article  Google Scholar 

  • Issacs R (1994) Studies on the chemical ecology of the black bean aphid, Aphis fabae. Ph.D. Thesis, University of London

  • Kimani-Njogu SW, Trostle MK, Wharton RA, Woolley JB, Raspi A (2001) Biosystematics of the Psyttalia concolor species complex (Hymenopera: Braconidae: Opiinae): the identity of populations attacking Ceratitis capitata (Diptera: Tephritidae) in coffee in Kenya. Biol Control 20:167–174

    Article  Google Scholar 

  • Kyparissoudas DS (1987) The occurrence of Encarsia perniciosi in areas of northern Greece as assessed by sex pheromone traps of its host Quadraspidiotus perniciosis. Entomol Hell 5:7–12

    Google Scholar 

  • Leal WS, Higushi H, Mizutani N, Nakamori H, Kadosawa T, Ono M (1995) Multifunctional communication in Riptortus clavatus (Heteroptera: Alydidae): conspecific nymphs and egg parasitoid Ooencyrtus nezarae use the same adult attractant pheromone as chemical cue. J Chem Ecol 211:973–985

    Article  Google Scholar 

  • Levi-Zada A, Nestel D, Fefer D, Nemni-Lavy E, Deloya-Kahane I, David M (2012) Analyzing diurnal and age-related pheromone emission of the olive fruit fly, Bactrocera oleae by sequential SPME-GCMS analysis. J Chem Ecol 38:1036–1041

    Article  CAS  PubMed  Google Scholar 

  • Lilley R, Hardie J, Wadhams LJ (1994) Field manipulation of Praon populations using semiochemicals. Nor J Agric Sci Suppl 16:221–226

    Google Scholar 

  • Lloyd JE (1997) Firefly mating ecology, selection and evolution. In: Choe JC, Crespi BJ (eds) The evolution of mating systems in insects and arachnids. Cambridge University Press, Cambridge, pp 184–192

    Chapter  Google Scholar 

  • Lloyd JE, Wing SR (1983) Nocturnal aerial predation of fireflies by light-seeking fireflies. Science 222:634–635

    Article  CAS  PubMed  Google Scholar 

  • Mangold JR (1978) Attraction of Euphasiopteryx ochracea, Corethreella sp. and gryllids to broadcast songs of the southern mole cricket. Fla Entomol 61:57–61

    Article  Google Scholar 

  • Mazomenos BE, Haniotakis GE (1981) A multicomponent female sex pheromone of Dacus oleae Gmelin: isolation and bioassay. J Chem Ecol 7:1561–1573

    Article  Google Scholar 

  • Mazomenos BE, Haniotakis GE (1985) Male olive fruit fly attaction to synthetic sex pheromone components in laboratory and field tests. J Chem Ecol 11:397–405

    Article  CAS  PubMed  Google Scholar 

  • Mbata GN, Shu S, Phillipis TW, Ramaswamy SB (2004) Semiochemical cues used by Pteromalus cerealellae (Hymenoptera: Pteromalidae) to locate its host, Callosobruchus maculatus (Coleoptera: Bruchidae). Ann Entomol Soc Am 97:353–360

    Article  CAS  Google Scholar 

  • McClain DC, Rock GC, Woolley JB (1990) Influence of trap color and San Jose scale (Homoptera: Diaspididae) pheromone on sticky trap catches of 10 aphelinid parasitoids (Hymenoptera). Environ Entomol 19:926–931

    Google Scholar 

  • McGregor R, Henderson D (1998) The influence of oviposition experience on response to host pheromone in Trichogramma sibericum (Hymenoptera: Trichogrammatidae). J Ins Behav 11:621–632

    Article  Google Scholar 

  • Micha SG, Wyss U (1996) Aphid alarm pheromone (E)-β-farnesene: a host finding kairomone for the aphid primary parasitoid Aphidius uzbekistanicus (Hymenoptera: Aphidiinae). Chemoecology 7(132):139

    Google Scholar 

  • Milonas P, Mazomenos BE, Kostantopoulou MA (2009) Kairomonal effect of sex pheromone components of two lepidopteran olive pests on Trichogramma wasps. Insect Sci 16:131–136

    Article  Google Scholar 

  • Mizutani N (2006) Pheromones of male stink bugs and their attractiveness to their parasitoids. Japan J Appl Entomol Zool 50:87–99

    Article  CAS  Google Scholar 

  • Noldus LPJJ, van Lenteren JC, Lewis WJ (1991) How Trichogramma parasitoids use moth sex pheromones as kairomones: orientation behaviour in a wind tunnel. Physiol Entomol 16:313–327

    Article  Google Scholar 

  • Powell W (1999) Parasitoid hosts. In: Hardie J, Minks AK (eds) Pheromones of non-lepidopteran insects associated with agricultural plants. CAB International, Wallingford, pp 405–427

    Google Scholar 

  • Powell W, Hardie J, Hick AJ, Höller C, Mann J, Merritt L, Nottingham SF, Wadhams LJ, Witthinrich J, Wright AF (1993) Responses of the parasitoid Praon volucre (Hymenoptera: Braconidae) to aphid sex pheromone lures in cereal fields in autumn: implications for parasitoid manipulation. Eur J Entomol 90:435–438

    Google Scholar 

  • Powell W, Pennacchio F, Poppy GM, Tremblay E (1998) Strategies Involved in the location of hosts by the parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae: Aphidiinae). Biol Control 11:104–112

    Article  Google Scholar 

  • Reddy GVP, Holopainen JK, Guerrero A (2002) Olfactory responses of Plutella xylostella natural enemies to host pheromone, larval frass, and green leaf cabbage volatiles. J Chem Ecol 28:131–143

    Article  CAS  PubMed  Google Scholar 

  • Ruther J, Steiner S (2008) Costs of female odour in males of the parasitic wasp Lariophagus distinguendus (Hymenoptera: Pteromalidae). Naturwissenschaften 95:547–552

    Article  CAS  PubMed  Google Scholar 

  • Ruther J, Meiners T, Steidle JLM (2002) Rich in phenomena lacking in terms—a classification of kairomones. Chemoecology 12:161–167

    Article  Google Scholar 

  • Samways MJ (1988) Comparative monitoring of red scale Aonidiella aurantii (Mask.) (Hom., Diaspididae) and its Aphytis spp. (Hym., Aphelinidae) parasitoids. J Appl Entomol 105:483–489

    Article  Google Scholar 

  • Schöller M, Prozell S (2002) Response of Trichogramma evanescens to the main sex pheromone component of Ephestia spp. and Plodia interpunctella, (Z, E)-9,12-tetra-decadenyl acetate (ZETA). J Stor Prod Res 38:177–184

    Article  Google Scholar 

  • Silva CC, Moraes MCB, Laumann RA, Borges M (2006) Sensory response of the egg parasitoid Telenomus podisi to stimuli from the bug Euschistus heros. Pesq Agropec Bras 41:1093–1098

    Article  Google Scholar 

  • Soper RS, Shewell GE, Tyrrell D (1976) Colcondamyia auditrix nov. sp. (Diptera: Sarcophagidae), a parasite which is attracted by the mating song of its host, Okanagana rimosa (Homoptera: Cicadidae). Can Entomol 108:61–68

    Article  Google Scholar 

  • Steidle JLM, Steppuhn A, Ruther J (2003) Specific foraging kairomones used by a generalist parasitoid. J Chem Ecol 29:131–143

    Article  CAS  PubMed  Google Scholar 

  • Sternlicht M (1973) Parasitic wasps attracted by the sex pheromone of their coccid host. Entomophaga 18:339–342

    Article  Google Scholar 

  • Zuk M, Kolluru R (1998) Exploitation of sexual signals by predators and parasitoids. Quart Rev Biol 73:415–438

    Article  Google Scholar 

Download references

Acknowledgments

We would like to thank three anonymous reviewers for their comments on an earlier version of the manuscript, Augusto Loni for help in P. concolor mass-rearing, Serena Geri and Giulia Giunti for assistance during experiments. This research was financially supported by M.I.U.R. (Ministry of Education, University and Research, Italy)—P.R.I.N. 2007.

Author information

Authors and Affiliations

  1. Insect Behaviour Group, Department of Agriculture, Food and Environment, University of Pisa, via del Borghetto 80, 56124, Pisa, Italy

    Giovanni Benelli, Sandro Simoncini, Alfio Raspi & Angelo Canale

  2. Department of Chemistry and Industrial Chemistry, University of Pisa, via Risorgimento 35, 56100, Pisa, Italy

    Adriano Carpita

Authors
  1. Giovanni Benelli
    View author publications

    You can also search for this author in PubMed Google Scholar

  2. Adriano Carpita
    View author publications

    You can also search for this author in PubMed Google Scholar

  3. Sandro Simoncini
    View author publications

    You can also search for this author in PubMed Google Scholar

  4. Alfio Raspi
    View author publications

    You can also search for this author in PubMed Google Scholar

  5. Angelo Canale
    View author publications

    You can also search for this author in PubMed Google Scholar

Corresponding author

Correspondence to Giovanni Benelli.

Additional information

Communicated by N. Desneux.

Rights and permissions

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Benelli, G., Carpita, A., Simoncini, S. et al. For sex and more: attraction of the tephritid parasitoid Psyttalia concolor (Hymenoptera: Braconidae) to male sex pheromone of the olive fruit fly, Bactrocera oleae . J Pest Sci 87, 449–457 (2014). https://doi.org/10.1007/s10340-014-0595-1

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s10340-014-0595-1

Keywords

Search

Navigation