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Behavioral Adaptations of Cryptic Moths. Iv. Preliminary Studies on Species Resembling Dead Leaves

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Behavioral Adaptations of Cryptic Moths. Iv. Preliminary Studies on Species Resembling Dead Leaves JOURNAL OF THE LEPIDOPTERISTS' SOCIETY --- - - ------ Volume 23 1969 Numher 4 BEHAVIORAL ADAPTATIONS OF CRYPTIC MOTHS. IV. PRELIMINARY STUDIES ON SPECIES RESEMBLING DEAD LEAVES RONALD R. KEIPER Dept. Biology, Pennsylvania State University, Mont Alto, Pennsylvania A number of experimental studies have demomtrated that cryptic coloration conveys a selective advantage to animals by enabling them to avoid detection by predators (Young, 1916; Carrick, 1936; Turner, 1961). In addition to morphological adaptations, behavior of cryptic animals is also important. An animal, even if it is cryptically patterned, must be able to choosc an appropriate background and to orient itself so that the cryptic pattern attains maximum effectiveness. Several studies have shown that certain species of cryptic moths, which normally rest on tree trunks, are capable of selecting backgrounds which tend to match the reflectance of their forewings (Kettlewell, 1955; Sargent, 1966), and that these moths will adopt <l,ppropriate attitudes upon these backgrounds (Sargent, 1969). Therc are other moth species which rest in or on other substrates and which seem to be colored or patterned to match these substrates. This paper presents some early results from studies designed to examinc the beha vior of moths which resemble and rest in dead leaves. FIELD STUDIES Two techniques were used in studying the natural resting habits of these moths. The first of these simply involved ~;earching an area of forest floor covered with dead leaves. When a resting moth was located, a photograph was taken, along with notes concerning the resting attitude. Occasionally moths were flushed before being photographed. These were then followed to subsequent resting places and appropriate notes were taken. The second technique involved releasing moths in wooded areas, and following them to their resting places. These moths had been captured 206 KEIPER: Moths resemble dead leaves Vol. 23, no. 4 TABLE 1. FIELD OBSERVATIONS OF SOME MOTHS WHICH REST IN DEAD LEAVES Species1 Remarks Geometridae: Euchlllcna spp. Hubner On top of dead leaves, often with one or (0, 6) both wings raised and resting against some object (e.g., tree trunk, rock) Metarl'hanthis duaria Guenee Flat on top of leaves with wings spread (0, 3) H yperitis alienaria Herrich-Schaeffer Flat on top of leaves with wings spread (4, 2) Sabulodes transversata Drury Flat on top of leaves with wings spread (6, 0) Abbotana clematllria Smith and Abbot Flat on top of leaves with wings spread (0, 8) Nematocampa filamentaria Guenee Flat on top of leaves with wings spread (7, 0) Noctuidae: Parallelia bistriaris Hubner Often "head down" in leaves, wings (28, 6) closed Panopoda rufimago Hubner Tends to crawl at least partly under (0, 4) leaves, wings closed Zanclognatha crura lis Guenee In or on top of leaves, wings closed (50, 7) Chytolita morbidalis Guenee Flat on top of leaves, wings closed (llO, 10) Renia discolomlis Guence In or on top of leaves, wings closed (10, 2) 1 Numbers in parentheses: (previously undisturbed individuals, rell~ased individuals) at lights or "sugar," kept overnight in experimental boxes, and released the following morning. Here, as with the undisturbed moths, photo­ graphs and notes were taken. The two techniques produced similar results for a number of species and these observations are presented in Table 1. These results suggest that a number of brownish, leaf-like moths are able to select appropriate backgrounds and further, to orient with respect to these backgrounds so as to increase their crypsis. EXPERIMENTAL STUDIES Several species were released into an experimental box in an attempt to determine if the moths would exhibit preferences for certain test substrates. The experimental box (24 inches square by 19 inches high) was constructed of plywood. The floor of the box was equally divided into four quadrats (each 12 x 12 inches), and during these experiments, 1969 Journal of the Lepidopterists' Society 207 TABLE 2. SELECTION OF SUBSTRATES BY MOllIS IN AN EXPERIMENTAL APPARA TUS Number of individuals selecting substrates: pine dead dead Species needles grass leaves twigs P Parallelia bistriaris Hubner 3 6 11 3 ns Epizeuxis lubricalis Geyer 0 2 15 3 * Zanelognatha cruralis Guenee 3 3 10 5 ns Amphipyra pyramidoides Guenee 2 3 19 2 * * Denotes significant d eviations from chance selections. Analysis by chi square tests, Probability less than 0.01 ns Denotes n o significant deviation from chance selection. Probability greater tha n 0.05 each quadrat contained a different substrate. In these experiments, the substrates used were: dead necdles of white pine, dead grass stems, dead oak and maple leaves, and small twigs. Fresh substrate material was added nightly, and the box was rotated to eliminate any "position" effect. The box was covered with a pane of window glass and placed in a wooded area where a thick canopy excluded direct sunlight. Moths were introduced into the box by sliding the glass top to one side. The moths were collected at lights or "sugar" bait and immediately introduced into the experimental box, generally 5-10 moths of the same species per night. The following morning, between 0600-0800 E.S.T., the box was checked and the substrate selection by moths noted. The results of this experiment, which are summarized in Table 2, suggest that at least some of the species examined are capable of selecting appropriate substrates. However, the moths appear to fall into three groups with respect to their behavior in the box. Moths in the first group never selected a substrate but instead were always found sitting either on the glass or on the sides of the box (e.g., the geometrids Abbotarw clemataria Smith and Abbot and Nematocampa filamentaria Guenee) . The noctuids Parallelia bistl'iaris Hiibner and Zanclognatha cruralis Guenee are examples of moths which show the second type of behavior observed in the experimental box. Moths of this group select dead leaves, but also are frequently found resting on top of other substrates. From observation of both released and undisturbed moths, it appears that both species behave the same way in the wild, resting on top of leaves or other substrates if forced into areas where no dead leaves are present. Apparently these species choose any substrate which offers a relatively flat surface. P. bistl'iaris further enhances its crypsis by resting 208 KEIPER: Moths resemble dead leaves Vol. 23, no. 4 TABLE 3. SELECfION OF DIFFERENTLY COLORED SUBSTRATES (3 X 3 IN. CARDBOAHD SQUARES) IN AN EXPERIMENTAL BOX Number of individuals selecting substrates: Species Black Brown Green Yellow P Parallelia bistriaris H libner 4 1 2 1 Ep'izeuxis lubricalis Geyer 2 3 1 2 Amphipyra pyramidoides Guenee 12 5 5 4 with the head burrowed into the substrate so that the moth is resting almost perpendicular to it. This posture allows the lighter-colored edge of the wing to stick up from the substrate and thus serves to break up the pattern of the moth. On the other hand, the noctuids Epizeuxis lubricaUs Geyer and Amphipyra pymmidoides Guenee did not rest on top of any substrate, but instead were found resting on the plywood floor of the box beneath the layer of dead lcaves. This third type of behavior may not represent selection of dead leaves as such, but may only result from the moths choosing this substrate because it can be easily burrowed into and then effectively shields the moths from the light. In an attempt to learn more about the mechanisms governing these choices, several additional experiments were devisecl. The same box was used, but cardboard squares (3 X 3 inches) were presented instead of the previous natural substrates in an attempt to keep the texture of the substrates constant. In the first experiment, an attempt was made to determine the im­ portance of the color of the substrate. Accordingly, squares of four different colors were used: black, chocolate brown, dark green, and bright yellow. The results of this experiment are shown in Table 3. For A. pymmidoides, there were sufficient numbers for a chi square test, but no significant selection tendency was noted. The sample sizes for the other species were too small to analyze separately, but the results were tested across species. In both cases however, the results suggcsted that the color of the substrate was not important to the moths in chOOSing a substrate. In an attempt to determine the influence of the shapes of the substrate components on the choice of the moths, squares of white cardboard were used, but they were presented in four different shapes: flat squares ("flat"), squares folded in the middle and set on end ("tent"), squares folded in the middle and set on a side ("right angle"), and squares laid flat but set on top of a nail ("leaf") and thus raised off the floor of the 1969 Journal of the Lepidopterists' Society 209 TABLE 4. SELECTION OF SUBSTRATE COMPONENTS (3:< 3 IN. CARDBOARD SQUARES ) OF DIFFERENT SHAPES IN AN EXPERIMENTAL BOX Number of individuals selecting substrates: "Right Species "Tent" Angle" "Flat" "Leaf" p Parallelia bistriaris Hubner 5 6 4 5 ns Epizeuxis lubricalis Geyer 19 1 0 0 * Amphipym pyramidoides Gllenee 26 0 0 0 * * Denotes significant d eviations from chan ce selections. Analysis by chi square t ests, p robability less than 0.01 ns D enotes no significant deviation from chance selectio n. Probability grea ter than 0.05 box. Sufficient numbers were obtained for only three species, but fortunately these represent two of the previously discussed types. The results from this experiment are shown in Table 4. P. bistriaris was found about evenly distributed on all substrates, but was oriented somewhat differently on each. For example, on the "right angle" squares, this moth generally perched 'bead down" on the inside of the extended arm.
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