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Preference of Three Scarab Beetle Species to Floral Cues

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Preference of Three Scarab Beetle Species to Floral Cues Materials Express 2158-5849/2020/10/1764/007 Copyright © 2020 by American Scientific Publishers All rights reserved. doi:10.1166/mex.2020.1817 Printed in the United States of America www.aspbs.com/mex Preference of three scarab beetle species to floral cues Hongfei Zhang1,∗,†, Weizheng Li2,†, Yan Zhang3, Guohui Yuan2, and Mingsheng Yang1 1College of Life Science and Agronomy, Zhoukou Normal University, Zhoukou 466001, Henan, PR China 2Department of Entomology, College of Plant Protection, Henan Agricultural University, Zhengzhou 450002, Henan, PR China 3School of Education Science, Zhoukou Normal University, Zhoukou 466001, Henan, PR China ABSTRACT The role of floral visual cues was studied in both sexes of three nocturnal scarab beetle species (Holotrichia oblita, Holotrichia parallela, and Anomala corpulenta). Flower patterns were designed using n-petal rose curve and radial gradient tools. BioassayIP: 192.168.39.151 of plain colored On: patterns Tue, 28 showedSep 2021 that 08:49:27 both sexes of H. oblita and H. parallela preferred yellow and white. In contrast,Copyright:A. corpulenta American showedScientific sexual Publishers differentiation in preferences. Comparison Delivered by Ingenta between given radial gradient patterns and their color components indicated that a radial gradient was nec- essary in both sexes of H. oblita rather than both sexes of H. parallela to elicit the highest response. Sexual Article differentiation was found in A. corpulenta. Among 4-, 8-, and 12-petaled patterns, the 4-petaled patterns were most preferred by all of the test insects, regardless of species and sex. Choice assays that provided both odor and visual cues suggest that olfaction may be the primary sensory modality in the three scarab species. Keywords: Scarab Beetle, Visual Cue, Rose Curve, Pattern. 1. INTRODUCTION ambient illumination is below 0.1 lx. Visual cues become Three scarab beetle species, Holotrichia oblita, H. paral- more important than olfactory cues (sex pheromone) once lela,andAnomala corpulenta, damage many crops seri- the males are in the vicinity of a pheromone source [2]. ously in arid areas in north China. Recently, our group A similar phenomenon was observed in a congeneric has reported a generic attractant blend (120 L phenylac- diurnal species H. loochooana loochooana during mate- etaldehyde and 80 L benzyl acetate, embedded in 8 mL searching [3, 4]. Miao et al. found that the copulation effi- 2% agaropectin) for their adults [1]. Further improvement ciency of A. corpulenta pairs significantly decreased after of trapping efficiency of this chemical attractant is depen- male compound eyes were blackened or the female elytra dent on the optimization of trap parameters. Both pheny- were blackened [5]. These experiments suggested that the lacetaldehyde and benzyl acetate are typical floral scents, three scarab species studied here may also see visible light suggesting that this attractant may be improved by apply- reflected by colored objects at dusk. However, informa- ing in flower-like traps. tion is lacking regarding to the responses of the scarabs to The role of visual cues in mate-searching behavior has visual cues in another ecological context, flower-visitation, been documented in several scarab species. For example, although some studies have compared the effect of trap males of a nocturnal species, H. kiotonensis, can discrimi- colors on field trapping of several scarab species [6–8]. nate between black and white cotton balls, even when the In this study, we investigated the role of floral visual cues in both sexes of H. oblita, H. parallela,andA. corpu- ∗Author to whom correspondence should be addressed. lenta, as well as the priority of visual and olfactory modal- †These two authors contributed equally to this work. ities. We tested the effect of the plain color, the radial 1764 Mater. Express, Vol. 10, No. 10, 2020 Preference of three scarab beetle species to floral cues Materials Express Zhang et al. gradient patterns consisting of two preferred colors, and cylinder was fitted tightly to the lower cylinder (Fig. 1). the corolla petal number factors on the choice response of Bioassays were conducted from 1,900 hours to 2,200 hours these beetles. After these tests, we determined the priority at night in the laboratory, in consistent with the natural of two sensory modalities by comparing the attractiveness flight peak time of these scarab adults. Unless otherwise of the corresponding preferred patterns with a common stated, the beetles were released one by one in the follow- olfactory cue in each sex of each species. Intra- and inter- ing experiments, and each beetle was tested only once. In species differences related to the response to these floral each test, a beetle was introduced from a hole (6 cm inter- cues are discussed. nal diameter) in the cover center of the upper cylinder. The choice response of the beetle was noted under dim illu- mination provided by sunlight-emitting diodes. When the 2. MATERIALS AND METHODS beetle crawled over any one flower model at the first time 2.1. Insects and stay there for at least 5 seconds, it was considered as Scarab beetles were collected by light traps at the Kaifeng “choice.” When it did not crawl towards any option within Experiment Station, China Agricultural University. The three minutes, or crawled without a certain direction, it following morning, they were separated into six treatment was recorded as “no choice” and discard from statistical groups by species and sex. The beetles ready for use were analysis. Sexes and species were tested separately in all reared on fresh Siberian elm leaves in plastic boxes. of the experiments. The apparatus was dismantled after every five individuals were tested, and its component parts 2.2. Preparation of Floral Patterns were thoroughly cleaned using absolute alcohol, followed We used “n-petaled Rose Curve” and radial gradient tool by rinsing in distilled water, to remove any residual odors. to simulate floral patterns. A rose curve can be expressed x = n× by a pair of Cartesian parametric equations: cos 2.4. Plain Color Patterns y = n × sin , cos cos .Ifn is an integer, the curve We tested the preference of scarab beetles to eight plain- will be rose-shaped with 2n petals if n is even, and n petals colored flower models. The eight flower models, as men- Article if n is odd. The preparation procedure was: (1) create an tioned above, were randomly arranged on the bottom of the = arithmetic progression of (initialIP: 192.168.39.151 term 0.01, common On: Tue, 28cylinder. Sep 2021 The positions08:49:27 of the models changed randomly = = difference 0.01, and total term numberCopyright:800) American and cal- Scientificbetween Publishers replicates, to rule out possible unintended effects culate the corresponding x and y values in MicrosoftDelivered Excel by Ingentaon the beetles’ choices. Each sex within each species was X Y 2007; (2) plot an – scatter graph with a smooth curve, replicated with 120 individuals. remove data markers and gridlines; (3) paste the scatter graph on a 32-bit canvas with suitable size in Adobe Pho- 2.5. Radial Gradient Patterns toshop CS 5; (4) RGB values of the eight plain colors used Based on plain color preference, we prepared a variety in this study are as follows: red (R = 255, G = B = 0), of radial gradient patterns (Fig. 2) and tested the prefer- green (R = B = 0, G = 255), blue (R = G = 0, B = 255), ences of each sex within each species to these patterns yellow (R = G = 255, B = 0), cyan (R = 0, G = B = 255), under different conditions presented with two, three, or six magenta (R = G = 255, B = 0), black (R = G = B = 0), options. The number of replicates under various conditions and white (R = G = B = 255). A radial gradient pattern can be deducted from the “choice frequency” columns in consisted of two colors (one as the central marking color Tables II–IV. A total of six models were arranged. Under and the other as the corolla petal color) selected from the eight solid colors was painted with radial gradient tool; (5) print flower models on a Ricoh Aficio MPC 5501 inkjet color printer (Ricoh Imagine Change, Shanghai, China). d For simplicity, we named a radial gradient pattern from a the central marking color to the corolla petal color (e.g., yellow–white). All of the models were 3 cm in diameter. b 2.3. Bioassays The bioassay apparatus consisted of two Perspex cylinders (5 cm height × 40 cm ID). Both cylinders had transparent c covers but lacked bottoms. The lower cylinder was placed on a block of green floral foam (50 cm × 50 cm × 3cm). Fig. 1. Bioassay apparatus used for scarab beetle tests. Left: Break- down drawing of the apparatus. (a) upper cylinder; (b) lower cylinder; We used green floral foam to simulate the common back- (c) floral foam; and (d) beetle release position. The two arrows indicate ground of flowers encountered by these beetles in nature. the connection of the three parts of the apparatus. Right: Top view of Floral models were placed on the floral foam at equal inter- flower model arrangements on the floral foam. Dotted ring indicates the vals along the periphery of the lower cylinder. The upper fitted position of the lower cylinder. Mater. Express, Vol. 10, 2020 1765 Materials Express Preference of three scarab beetle species to floral cues Zhang et al. 2.6. Corolla Petal Numbers To determine the effect of corolla petal numbers on bee- tle preference, we designed four, eight, and twelve petals for the corresponding optimal patterns for each sex of each species, and tested them under conditions with three options. A total of six models, two for each type of pat- tern, were randomly arranged (N = 150). In these tests, the contour length of all the flower models kept consistent (3 cm diameter).
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