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Azimuth Orientation of the Dragonfly (Sy Mpetrum)

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Azimuth Orientation of the Dragonfly (Sy Mpetrum) Azimuth Orientation of the Dragonfly (Sy mpetrum) MITUHIKO HISADA Hokkaido University ELATIVELY LITTLE IS KNOWN about the ies of the dragonfly (Sympetrum costiferum, R migratory behavior of the dragonflies. Kennedy, ref. 7, Cratilla calverti, ref. 8). Sole summer populations of Anax junius in However, the factor or factors determining Canada are thought to be maintained by mi- orientation are yet to be convincingly demon- gration from the southern range of their dis- strated. tribution (refs. 1 and 2). Other species, such While collecting the dragonfly Sympetrum as Sympetrum rubicundulum (ref. 3) in species in the field as an experimental mate- North America, and Sympetrum striolatum rial, we noted a peculiar tendency of the (ref. 4) and Aeshna mixta (ref. 5) in Eu- alighting individuals to take a particular rope, have been recorded as flying south in direction relative to the Sun. This phenom- early autumn. If these northward and south- enon attracted our attention because of possi- ward movements of the dragonflies are, as ble connection to the migratory behavior as they appear to be, unidirectional and well well as to the dorsal light reaction of the oriented, then it becomes of interest to find species that we have already partly described out the factor or factors determining the ori- (ref. 9). Cursory field observation revealed entation mechanism in migration. that the direction of orientation is different In the past only a few remarks have been at different times of the day as it appeared to made on the orientation of the settling dra- be somehow related to the displacement of gonflies. Adults of Sympetrum sanguineum the azimuth of the Sun. are reported to settle in large numbers on Directional phototactic orientation of telegraph wires or fences. The orientation of many kinds of animals, both in the field and these settling individuals is said to be very in the laboratory, has been amply described constant, being related to the direction of (refs. 10 to 12). There are, however, a small incident illumination, so that they face away number of observations on the orientation of from the darkest part of their immediate sur- alighting insects. The contribution of the dor- roundings (ref. 6).This habit of settling en sal light reaction to the equilibrium reaction masse on telegraph wires and adopting a con- of the dragonfly Anax imperator has been stant orientation is also shown by other spec- analyzed in detail 4y Mittelstaedt (ref. 13), 511 512 ANIMAL ORIENTATION AND NAVIGATION mainly in free flying animals. Some other ob- sunlight from the nearby substratum, data servations exist, but they place emphasis on were collected from the dragonflies alighting the reaction of various insects either around exclusively on tops of bamboo sticks project- their longitudinal axis (banking response) or ing about 2 m above the ground. It is of transverse axis (pitching response) when the interest that there were no directional devia- light source is placed around those axes (refs. tions of the individuals when the tops were 14 to 16). lower than 2 m. However, there were direo Only one report exists on the azimuth ori- tlonal deviations when the dragonflies were entation of the dragonfly in the horizontal on flat surfaces of high reflectance, such as a plane. Thus Jander (ref. 17) demonstrated white-painted signpost. the response of Calopteryx splendens, illumi- Around the end of September and early nated from various directions in the horizon- October, the daily active phase of the dra- tal plane; but again the emphasis was on the gonfly appears to start at about 8:OO a.m. change of posture, not on the change of and continues almost to sunset, the general direction. level of activity being greatly affected by the In this paper, evidence is presented of atmospheric temperature and the amount of directional orientation of the alighting dra- sunshine. During warm, but not too hot, gonfly relative to the azimuth of the Sun, and brighter, and less windy weather the number an indication is given of the contribution of of individuals appearing in the field and the wind direction to this orientation. Some alighting is more abundant. On the other preliminary experiments to seek out a possi- hand, with winds of more than 2 to 3 m/sec, ble operative receptor for this orientation will there is a reduction in number of individuals also be noted. that can be observed. Thus the measurements Observed individuals belong to genus were performed mainly on relatively calm Sympetrum (Libellulidae, Odonata) ,of which days. more than 95 percent are Sympetrum fre- Azimuth bearing of the individuals alight- quens Selys, with a very few being S. darwin- ing and the azimuth of the Sun were both ianum S. and S. infuscatum S. However, no measured with the aid of a compass attached particular difference in behavior was noticed to a clinometer for geological survey. Wind among those species. velocity and direc~onwere measured with the aid of a thermistor-type anemometer. FIELD OBSERVATION Alighting individuals were approached from the rear, and the azimuth readings of General Behavior their longitudinal body axes were measured through the sight attached to the clinometer. Field observations were performed in an In each run of the measurement, 20 to 35 open field on the university campus, which individuals could be observed alighting on measures about 30 m in east-west direction the tops of bamboo sticks that were separated and about 100 m in north-south direction, from each other by at least 2 m. The read- and is surrounded with low bushes and bor- ings were taken as fast as possible, mostly dered with some trees about 5 m high at the within less than 10 min, and at the same north-east corner, the nearest of which stands time, the direction of the wind and also the about 30 m apart from the field’s edge. wind speed were recorded together with the To avoid possible interference by reflected azimuth of the Sun and air temperature. SESSION V: SENSORY MECHANISMS-OTHER 513 Dragonfly alighting appears to be per- appear as a jerking movement of the head formed in at least four distinct phases. First and also a slight depression of the wings. If the dragonfly will approach the stick from a the dragonfly is startled with a stronger stimu- direction that is usually slightly deviated lus, it will take off but will often return re- from the final orientation it will take after peatedly to the same spot through the almost settling. Second, after a short hovering over identical course and alight again after a short the stick, it will alight. Third, while the while. wings are still stretched horizontally, it will At noon of a hot day (27" C) ,one individ- make a final adjustment of the orientation by ual was found to be taking a very peculiar stepping around on the tip of the stick. Dur- posture while alighting on the ground: Its ing this phase, the head is still pivotted abdomen pointed upward almost in the around. Finally, after a short while, the wings direction of the rays of the Sun (close to 45 will be slightly drooped with the head held deg). In this manner it presented the mini- fast until startled by a moving object. It is mum surface to the direct rays of the Sun, rather curious to observe that when the dra- while the wings were drooping to cover the gonfly is startled, the first sign of alerting will thorax. This kind of posture has been ob- served in tropical species (refs. 18 and 19). Also around sunset, on a calm and rather cool day (15" C), many individuals were found to be perched headup on the vertical surfaces of nearby walls to expose their dorsal surfaces to the direction of the Sun. Other- wise, all individuals observed under the direct sunshine in the field have been observed to take a horizontal alighting posture, and their orientation was undoubtedly within a certain azimuth angle. Orientation to the Sun's Azimuth Four runs of the observation were per- formed in one day within their active phase FIGURE 1. Changes in orientation direction of to see the orientation change relative to the alighting dragonflies relative to the Sun's azimuth movement of the Sun. The measurements observed on one day. Azimuth is shown in heavy were performed on October 1, 1966, a quite arrows and wind direction in dotted arrows. calm and warm day. The air temperature North is upward. (A) Time of observation: 940-9:50 a.m. Overcast: cloudless. Azimuth SE around the measured dragonflies stayed 23 to 25-23" (measured from due south). Wind NE, 27 C ; there was no overcast nor haze during 0.2 m/sec max. Air temperature at 2 m above the daytime. The first measurement (fig. 1A) ground 23°C. (B) 1208-12:20 p.m. Cloudless. was performed at 9:40 to 9:50 a.m., with the Sun: SW 22-26". Wind SW, 0.2-0.6 m/sec. Sun's azimuth reading at 25 to 23"SE. The 27" C. (C) 2:30-2:50 p.ni. Cloudless. Sun: SW 65-71". Wind S, 1 m/sec max. 25" C. (D) 418- measurements were done on 22 individuals, 420 p.m. Cloudless. Sun: SW 82-85". Wind S, all of which alighted on the tips of the bam- 1.5 m/sec max. 24" C. boo sticks. The wind velocity was around 0.2 514 ANIMAL ORIENTATION AND NAVIGATION m/sec and from the northeast. Two groupings of dragonflies, both facing almost at right angles to the azimuth of the Sun were clearly recognized.
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