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Diptera: Syrphidae) ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Entomologie heute Jahr/Year: 2018 Band/Volume: 30 Autor(en)/Author(s): Neimann Alexander, An Lina, Lunau Klaus Artikel/Article: The Yellow Specialist: Colour Preferences and Colour Learning of the Hoverfly Eristalis tenax (Diptera: Syrphidae). Der Gelbspezialist: Farbpräferenzen und Farbenlernen der Schwebfliege Eristalis tenax (Diptera: Syrphidae) 27-44 Colour preferences and colour learning of the hoverfl y Eristalis tenax 27 Entomologie heute 30 (2018): 27-44 The Yellow Specialist: Colour Preferences and Colour Learning of the Hoverfly Eristalis tenax (Diptera: Syrphidae) Der Gelbspezialist: Farbpräferenzen und Farbenlernen der Schwebfliege Eristalis tenax (Diptera: Syrphidae) ALEXANDER NEIMANN, LINA AN & KLAUS LUNAU Summary: The hoverfl y Eristalis tenax (Syrphidae, Diptera) has a pronounced preference for yel- low fl owers like many other members of the syrphid family. Experiments testing landing behaviour and proboscis extension indicate that E. tenax has an innate preference for yellow colours. Little is known about colour learning in E. tenax and about colour parameters determining the fl ies’ colour preference. This study focuses on the colour preference and colour learning regarding the role of UV-refl ection properties for visiting yellow fl owers. In multiple choice experiments with artifi cial fl owers inexperienced and naïve E. tenax fl ies showed a clear preference for yellow colours, but no preference in landing behaviour for UV-absorbing or UV-refl ecting yellow artifi cial fl owers. In ad- dition, the fl ies also chose bright UV-absorbing non-yellow artifi cial fl owers for landing. The fl ies extended their proboscis preferably towards dark and UV-absorbing yellow colour patches. These results suggest that brightness has a so far unknown infl uence on colour choice both for landing and proboscis extension. The colour preferences of E. tenax seem to be fi ne-tuned to yellow fl owers displaying an ultraviolet refl ection pattern. Keywords: Eristalis tenax, dronefl y, colour preference, fl ower visitation Zusammenfassung: Die Schwebfl iege Eristalis tenax (Syrphidae, Diptera) hat eine ausgesprochene Präferenz für gelbe Blüten wie viele andere Schwebfl iegen auch. Experimente zum Landeverhal- ten und zur Rüsselrektion belegen eine angeborene Präferenz von E. tenax für gelbe Farben. Nur wenig ist bekannt über Farbenlernen bei E. tenax und über die Eigenschaften von Farben, die die Farbpräferenz bestimmen. Diese Studie beschäftigt sich mit Farbpräferenzen und Farbenlernen und den Einfl uss von UV-Refl exion auf den Besuch von gelben Blütenattrappen. In Mehrfachwahlex- perimenten mit unerfahrenen und naiven E. tenax zeigten die Fliegen eine klare Präferenz für gelbe Farben, aber keine Präferenz für UV-refl ektierende oder UV-absorbierende gelbe Blütenattrappen. Zusätzlich wurden helle, UV-absorbierende, nicht-gelbe Blütenattrappen für die Landung gewählt. Die Fliegen zeigten die Rüsselreaktion bevorzugt zu dunklen, UV-absorbierenden Farbfl ecken. Diese Ergebnisse belegen einen bislang unbekannten Einfl uss der Helligkeit auf die Farbwahl bei Landung und Rüsselreaktion. Die Farbpräferenzen von E. tenax scheinen fein abgestimmt zu sein auf den Besuch von gelben Blüten mit einem UV-Muster. Schlüsselwörter: Eristalis tenax, Mistbiene, Farbpräferenz, Blütenbesuch 1. Introduction plant (VON FRISCH 1914; KELBER & OSORIO 2010; HOPKINS & RAUSHER 2012; LUNAU & Many fl ower visitors use visual colour cues MAIER 1995; VAN DER KOOI et al. 2018). The of fl owers to identify and fi nd their food dronefl y Eristalis tenax (Syrphidae, Diptera) Entomologie heute 30 (2018) 28 ALEXANDER NEIMANN, LINA AN & KLAUS LUNAU is a Batesian mimic of the Western honeybee of the fl ies (BASTIAN 1986; GILBERT 1986). Apis mellifera (GOLDING & EDMUNDS 2000; Th e fl ies take nectar from fl owers for energy LUNAU 2011). Honeybees and dronefl ies visit supply (WOODCOCK et al. 2014). fl owers for nectar and pollen (SRINIVASAN Previous studies demonstrated that E. & GUY 1990). The imagines are usually tenax fl ies respond to light stimuli while observed from March to October visiting flying and walking (MAST 1923), have fl owers or laying eggs at manure heaps and colour vision and possess a preference eutrophic waters (GILBERT 1986). The larvae for yellow colours (ILSE 1949). The fl ies exhibit a saprophagous live style in polluted exhibit fl ower constancy also for fl owers waters, muddy pools and liquid manure of other colours than yellow (KUGLER and are considered to be an indicator for 1951). E. tenax exhibits an innate probos- polysaprobic water quality (KOLKWITZ & cis refl ex towards yellow colours (LUNAU MARSSON 1909; CHAPMAN 1996). Because of 1987; STERNKE-HOFFMANN & LUNAU 2015). the extended larval anal breathing tube, the Pollen, anthers and fl ower marks of ma ny larvae of E. tenax are called rat-tailed mag- entomophilous fl owers visited by E. tenax gots. The imagines of E. tenax are consid- refl ect wavelengths longer than 510nm and ered as important pollinators in greenhouse absorb UV-light (LUNAU & WACHT 1997a). agriculture (JARLAN et al. 1997). In the wild Yellow and UV-absorbing pollen, anthers the fl ies usually visit yellow and white fl ow- and fl ower marks trigger a spontaneous ers for pollen and nectar (GILBERT 1986; DE proboscis refl ex in newly emerged and BUCK 1990). The freshly hatched imagines inexperienced E. tenax imagines (LUNAU depend on pollen in the fi rst place. The 1988; LUNAU & WACHT 1994, 1997a). This nutrients contained therein are essential for response to yellow colours is innate and the development of the reproduction system cannot be modifi ed by training (LUNAU Fig. 1: Spectral sensitivity of the R8y photoreceptor type of Eristalis tenax (after TSUKAHARA & HORRIDGE 1977; dashed line) and spectral effi ciency function of the innate proboscis extension towards monochromatic test stimuli (solid line). Both functions are normalized at the maximum. Modifi ed from LUNAU & WACHT (1994). Abb. 1: Spektrale Empfi ndlichkeit des R8y-Photorezeptortyps von Eristalis tenax (nach TSUKAHA- RA & HORRIDGE 1977; gestrichelte Linie) und spektrale Reizwirksamkeitskurve der angeborenen Rüsselreaktion auf monochromatische Teststimuli (durchgezogene Linie). Beide Funktionen sind normalisiert auf das Maximum. Modifi ziert nach LUNAU & WACHT (1994). Colour preferences and colour learning of the hoverfl y Eristalis tenax 29 1988; LUNAU et al. 2018). The innate pro- fl ower visiting insects (LUNAU & MAIER boscis refl ex is triggered by monochro- 1995; VAN DER KOOI et al. 2018) because matic light stimuli in the small range of they need to detect fl owers of potential wavelength between 520nm and 600nm foodplants and to discriminate between (LUNAU & WACHT 1994) and inhibited by fl owers of different plant species. Honey- admixed blue and ultraviolet light (LUNAU bees (A. mellifera) and bumblebees (Bombus & WACHT 1997b). These results suggest terrestris) are known for their colour prefer- that the stimulation of the retinula cell ence and their abilities of colour learning R8y is involved in this response to green and colour discrimination (ROHDE et al. and yellow monochromatic light (LUNAU 2013). & WACHT 1994; Fig. 1) stimuli similar to Many yellow fl owers display a so-called reactions known as wavelength-specifi c UV bull’s eye, i.e. an ultraviolet refl ection behaviour (CRONIN et al. 2014). pattern with a UV-absorbing centre part The compound eyes of fl ies consist of (LUNAU 2007). From experiments with hundreds to thousands ommatidia. All om- honeybees and bumblebees it is known matidia possess eight photoreceptor cells. that they respond to this kind of fl oral Although colour vision in fl ies is largely colour pattern. Bees do not discriminate understudied, most studies agree to as- between yellow fl owers with or without sume to separate visual subsystems, which a UV-absorbing fl ower centre, but the are a colour-blind neural superposition fi rst physical contact with the fl ower by subsystem and a tetra variant colour vision means of the antennae is targeting to the subsystem (for review see LUNAU 2014). UV-absorbing area (PAPIOREK et al. 2016). The six peripheral receptor cells are labelled Similar studies for hoverfl ies are missing. R1-R6, have identical spectral sensitivity, This study aims to investigate how E. tenax are organized in a neuronal superposition fl ies respond to yellow colours when land- system and are responsible for motion vi- ing and when extending the proboscis with sion (LUNAU & WACHT 1994; LUNAU 2014). particular reference to the UV-refl ection The two interior receptor cells R7 and R8 properties. For this purpose the innate are arranged as a tandem and termed yel- preference for single-coloured artifi cial low (R7y and R8y) or pale (R7p and R8p) fl owers varying in brightness and colour in regard to the appearance of the omma- hue are studied as well as colour learning tidia in orthodromic illumination (HARDIE experiments. The data analysis is based on 1986; HARDIE & KIRSCHFELD 1983) and are the spectral refl ectance properties of the responsible for colour vision (TROJE 1993; tested colours and to a lesser extent based KELBER 2001; KELBER et al. 2003; LUNAU on the colour space of fl ies. The colour 2014). The retinula cell tandems, R7y/P8y, vision model of fl ies (TROJE 1993) that resp. R7p/R8p, are arranged with the R7 predicts that fl ies can discriminate only retinula cell located distal to R8 retinula cell. four colour categories, has been success- It is assumed that E. tenax fl ies thus possess fully applied for understanding of fl ower tetrachromatic colour vision. R7p is most colour choice in Eristalis fl ies (ARNOLD et sensitive to ultraviolet light, whereas R8p is al. 2009; BERGAMO et al. 2018; GRAY et al. sensitive in the blue range of wavelength. 2018). Spontaneous choice experiments R7y is sensitive to ultraviolet and blue light, with yellow artifi cial fl owers displaying a whereas R8y is sensitive to green light (LU- real or, alternatively, an inverse bull’s eye NAU & WACHT 1994).
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