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CONSPICUOUSNESS of BAND-WINGED GRASSHOPPERS to PREDATORS and CONSPECIFICS (ORTHOPTERA: ACRIDIDAE: OEDIPODINAE) a Senior Thesis P

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CONSPICUOUSNESS of BAND-WINGED GRASSHOPPERS to PREDATORS and CONSPECIFICS (ORTHOPTERA: ACRIDIDAE: OEDIPODINAE) a Senior Thesis P CONSPICUOUSNESS OF BAND-WINGED GRASSHOPPERS TO PREDATORS AND CONSPECIFICS (ORTHOPTERA: ACRIDIDAE: OEDIPODINAE) A Senior Thesis Presented to The Faculty of the Department of Organismal Biology and Ecology Colorado College Bachelor of Arts Degree in Biology BY Brae A. Salazar May, 2018 Approved by: ____________________________________________ Dr. Brian Linkhart Primary Thesis Advisor ____________________________________________ Dr. Emilie Gray Secondary Thesis Advisor ABSTRACT In band-winged grasshoppers (subfamily Oedipodinae), the variety of hindwing colors—ranging from blue to red—is both striking and unexplained. Hindwing color can vary both within and between species. However, the functional significance, if any, of this variation is unknown. Notably, the colorful hindwings are revealed only in flight, and remain hidden in stationary individuals. Although experimental evidence is lacking, this flash of color has been proposed to 1) startle potential predators, 2) to signal the quality of a potential mate, and 3) to enhance species recognition. To elucidate their potential function(s), here I measure the spectral and spatial characteristics of the hindwing patterns in 6 different band-winged species. I then model how an avian predator or potential mate might view grasshopper wings at behaviorally relevant distances. These data suggest that there is a rapid change in conspicuousness as a grasshopper moves from rest to flight regardless of the color vision of the receiver. However, there is little within species variation in coloration or wing patterning. My results indicate that while hindwings 1) may function as a protean defense against avian predators, 2) it is unlikely that they serve as a signal of mate quality, although, they 3) may deliver enough information for species recognition. This research helps to elucidate evolutionary relationships leading to the diversification of behavior, visual systems, and coloration within band-winged grasshoppers. Key Words: Oedipodinae, band-winged grasshopper, visual signaling, morphological variation, communication 2 INTRODUCTION Ecology and evolution are driven by studies that explore morphological variation. Morphological variation can result from numerous selective pressures including those of 1) predation, 2) sexual selection for interspecific discrimination, and/or 3) sexual selection for intraspecific discrimination of mate quality. Alternatively, neutral evolution can also lead to variable morphological outcomes. Nonetheless, we often observe morphological variation under influence of natural selection (Badyaev and Foresman 2000; Van Valen 1965; Caumul and Polly 2005). For example, in the passion-vine butterfly, wing coloration and pattern are correlated with success in both predator deterrence and mate attraction (Finkbeiner et al. 2014). Additionally, rhinoceros beetles exhibit a large diversity of horns; structurally specialized horns provide a competitive advantage in male to male combat (McCullough et al. 2014). Many species of bird exhibit plumage variability as well. This variability often signals status and aids in mate attraction (Rohwer 1975; 1977). Not only can morphological variability lead to increased functional efficiency, it can also provide information about the identity and quality of an individual as both a potential target for predation and/or courtship. However, potential predators and conspecifics cannot respond to these cues unless they are able to receive and discriminate between the variants present. Spatial and color vision vary greatly between animals (Cronin 2014; Land and Nilsson 2002; Eaton and Johnson 2007); visual signals that are conspicuous to one 3 receiver may be inconspicuous or cryptic to another (Speiser et al. 2013; Brandley et al. 2016). This visual sensory differentiation may be present both within and between species (Munz 1958; Lythogoe 1968; Menzel 1979; Jacobs 1984; Kornicker and Harrison-Nelson 1997; Marshall et al. 1999; Petrowitz et al. 2000; Briscoe and Chittka 2001; Frentiu and Briscoe 2008; Gumm et al. 2012; Brandley et al. 2013). Functionally, visual differentiation allows for a single color or structure to relay varying degrees of information to multiple perceivers (Brandley et al. 2013). This color or structure may act as a signal, and thereby, is under selection due to the signal’s behavioral effect on potential receiver(s) (Cronin et al. 2014). Therefore, the study of visual systems is important for investigating morphological evolution through sensory differentiation. If a potential predator can discriminate between intra/interspecific variants of a signal, the signal may function as deterrence to predators. If a conspecific can only detect interspecific variants, then the signal may serve for species recognition. Finally, if a conspecific can detect intraspecific variants, then a signal of mate quality may be present. Otherwise, the inability of a relevant receiver to discriminate between signals would suggest no functional significance in a given role. These variations, then, are considered incidental effects rather than evolved functions (Otte 1974; Gould and Lewontin 1979). Comparing the perception of signals between predators and conspecifics is a key tool for investigating selection forces acting on the diversification of morphology. Often, it is evolutionarily illuminating to explore the communication systems of organisms of interest via spatial and spectral visual modeling (Eaton 1987; Cronin et al. 2014; Stoddard and Prumm 2008; Brandley et al. 2016; Endler et al. 2001). This approach 4 can elucidate the source of pressure(s) leading to inter/intraspecific morphological variation. In band-winged grasshoppers (subfamily Oedipodinae), the variety of hindwing colors—ranging from blue to red—is both striking and unexplained (Figure 1). Hindwing color can vary both within and between species (Otte 1981; Schennum and Willey 1979). Additionally, their hindwings often include a black band of varying size and form. While most species have a small black band extending across the outer margins of the wing, some species have broad bands that fill only the inner margins. The functional significance, if any, of this variation is unknown. Notably, the colorful hindwings are revealed only in flight (Otte 1981). In stationary individuals, their often cryptically colored forewings conceal the hindwings and render the animal inconspicuous (at least to a human observer). When approached by a threat, many species of band-winged grasshoppers have been shown to prefer escape via flight (Cooper 2006; Bateman and Fleming 2014; Kral 2010). To a human observer, the striking appearance of their wings during flight is anything but cryptic. Therefore, previous studies suggest that band-winged grasshoppers’ predator flight response is a form of protean defense—the sudden conspicuous flash of their colored wings and unpredictable flight patterns are a method in which they can momentarily disorient a pursuing predator (Willey and Willey 1969; Humphries and Driver 1970). Despite these claims, no study has experimentally examined whether or not protean behavior decreases the risk of predation in band-winged grasshoppers. Alarmingly, it is only an assumption based solely on human spatial 5 and color vision that band-wings shift from conspicuous to inconspicuous as they transition from flight to rest. Figure 1: Variations in hindwing pattern and color. Pictured from left to right: Disossteira carolina, Arphia conspersa, Trimerotropis verruculata, Arphia pseudonietana, Chortophaga viridifasciata, and Spharagemon equale. In addition to having predator deterrence functions, naturalists have suggested that the hindwing coloration may have diversified under sexual selection (Otte 1970; Willey and Willey 1969). Males have been observed to make short advertising flights in which they flash their hindwings and crepitate (Otte 1981, 1970; Capinera et al. 2001; Niedzlek-Feave 1995; Willey and Willey 1969). Otte (1970) reports that these courtship flights are acoustically and visually conspicuous, causing attraction of the opposite sex. However, results from a field study conducted with Chortophaga viridifasciata postulate that crepitating courtship flights may function not to attract females, but to signal a male’s arrival in a new area. If a female becomes more active in response to the flight, then the male has a greater chance at locating her. Although this hypothesis is suggested, no data indicates an increase in female activity following the courtship flight of a male (NiedZlek-Feave 1995). Additionally, a number of species of band-winged grasshoppers exhibit red hindwings. Notably, a long-wavelength-sensitive photoreceptor has yet to have been detected in any species of band-winged grasshoppers, which is used to detect red coloration 6 (Briscoe and Chittka 2001). Therefore, it would appear erroneous that hindwing coloration has evolved strictly under sexual pressure. If, however, conspecifics can discriminate between interspecific variants, then the signal may serve for species recognition. On the other hand, if females assess the quality of a potential mate via courtship flight, then significant differences in color and/or hindwing structure must 1) be present within a species, and 2) be detected by a conspecific at behaviorally relevant distances. Here, I examine how potential avian predators and conspecifics view six band- winged species (Chortophaga viridifasciata, Dissosteira carolina, Arphia pseudonietana, Arphia conspersa, Spharagemon
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