Coenagrionidae) Ceriagrion Selys and Nipponicum Living in Old

Odonalologica 17(3): 195-204 September I. 1988 Adult ecology of CeriagrionmelanurumSelys and C. nipponicumAsahina (Zygoptera: Coenagrionidae) 1. diurnal variations in reproductive behaviour K. Mizuta Hiroshima Agricultural College Saijo, Higashi-Hiroshima, Hiroshima Prefecture, 724, Japan Received November 3, 1986 / Revised and Accepted April 18, 1988 The male tactics of mate-locating are the "searching”type in C. melanurum,and of ”sit-and-wait” in the type C. nipponicum. The male of C. melanurum actively makes flights for females searching during a limited period within a day and mates only during this period, while the male of C nipponicum mates duringa longer period within a day. When it becomes cloudy, the reproductive activities ofthe male of C. melanurum sharply go down, whereas the reproductive activities of C. nipponicum much are not affected so as those of C. melanurum. INTRODUCTION Ceriagrion melanurum Selys and C. nipponicum Asahina are damselflies in old and marshes with living ponds swamps or abounding emergent plants. The adult of C. is about halfof weight nipponicum one that of C. melanurum. but both abdomen and of the two wing lengths species are almost the same, being about 30 mm and 20 mm, respectively (ISHIDA, 1969). AOYAGI (1973) studied the adult behaviourand postures of C. melanurum. his the of of According to report, mode mate-locating this species is searching for females by males. On the other hand, it is reported that mature males of C. territories nipponicum adopt by perching on emergent plants (MIYAZAKI & C. MURAKI 1984). Thus, though melanurumand C. nipponicum belong to the same are genus and of similar size, the mate-locating modeof the male seems to differ in each in species. The mode C. melanurum would be regarded as the and that in C. ’’searching” type nipponicum as the ”sit-and-wait” type. Detailed studies have not been published on the adult activities of either species. Concentrating on the mate-locating tactics of both species, I have ob- 196 K. Mizuta served and described the time budgets and diurnal activities in mature adults. METHODS The from the research was conducted end of July till the middle ofAugust, 1965, at Midorogaike, Kyoto (N 35° 3.3', E 135° 46.3', about 80 m above sea level), and from the end of Julytill the middle of Iwanohimenomikoto Nara 80 August, 1969, at Mausoleum, (N 34° 41.9', E 135° 48.2', about m above sea level). During the research periods, the sun rose and set at about 5:00 — 5:15 h and 18.45-19.00 h JST, respectively. Both species occur in the two places. The individuals field research was conducted on mature under conditions using three methods as ofthe follows: (I) Hourly fluctuations time budgets of activities; The flights and their durations were observed 5 min. sometimes individual. individuals over or longer, per Feedingand copulating were the excluded from the observation. Time measurements of flights were made to unit of 0.1 s by a I the scale and measured and stopwatch. traced flight route on a map (the of which is 1:50) its length, then calculated the average speed of flight dividing the length of the flight route by the flight duration. Light intensities were measured by a Toshiba Photocell llluminometer SPI-5; — (2) Counts of ovipositing individuals; The ovipositing tandem pairs flying over a certain area (about 3 in of Iwanohimenomikoto m x 5 m) the pond Mausoleum were counted every hour. The study carried 13 and 1969; — Tandem males tethered was out on August 12, 15, (3) response by to In order the and sexual tandem females: to study presence intensity of drives of males, response was of male be tested. observed by presenting a tethered female 20-50 cm in front a to The above observations were made mainly on fine days. RESULTS The activities could roughly be grouped as follows: (a) Perching; — (b) Simple flight: a short and briefflight for unknownreason (Tab. I). A return to Table I Duration of flight of each type No. of ± SD Species & type of flight samples xx ± SD Range observed (sec) (sec) C. melanurum Simple flight $ 234 2.I± l.l1.1 0.5-70.5- 7 Simple flight 9$ 90 2.4± 1.5 0,5-100.5-10 Feeding flight <5 no110 3.3± 2.0 0.5-11 Feeding flight 9$ 92 3.I± 1.8 0.5-80.5- 8 Searching flight <5Q 1085 22.0±I3.922.0± 13.9 1-90 C. nipponicum Simple flight$ 297 1.8± 1.0 0.5-80.5- 8 Simple flight 9$ 227111 1.5± 0.8 0.5- 6 Feeding flight$ 83 3.l±l.83.l± 1.8 0.5-10 Feeding flight9$ 171 3.l± 2.1 0.5-120,5-12 Investigatory flight <5 184 4.5±4.5± 2.8 0.5-17 Ecology of Ceriagrion melanurum and C. nipponicum 197 after often This the original perch such a flight was observed. corresponds to the "normal described — flight” as by AOYAGI (1973); (c) Feeding flight: a dashing flight to catch small insects flying near the perch (Tab. 1). This activity couldbe observed other of during types flights as well, when a dashing flight was often made to insects in mud leaf. perching or, some cases, to spots on a The catch rate (the ratio of the number ofsuccessful catches to that offeeding flights) in C. melanurumwas investigated. The result was 0.11 (= 7/ 64) in the maleand 0.12 (= 13/112) in the female; — (d) Escape flight: When a large anisopteran, or an insect it net, were close to an individual, made a hasty and linear escape flight. When these appeared suddenly, the damselfly sometimes dropped downfolding Fig. I. Frequency distributionof durations of searching flight in the male of C. melanurum (top) and investigatory flight in the male of C. nipponicum(bottom). the This of wings instantly. type response was not observed during the study of time of — budgets activities; (e) Searching flight: a flight made by males of C. melanurum for females. apparently searching The males made a zig-zag flight at the height ofabout 50 their bodies and down 10-20 cm, shaking up among or cm above the surrounding bushes. They mainly flew forwards but sometimes hovered. When a femalewas found,they dashed towards it. The durationofflight with was long compared other types of flights (Tab. 1, Fig. 1). The flight speed measured in 5 flights lasting over seconds was 20.5 ± 5.0 cm/s (x ± SD, n=45). M1ZUTA (1974) termed this type of flight "sexual flight”; — (f) Investigatory 198 K. Mizuta of flight: a type flight madeby males ofC. nipponicum. The males ofthis species, when finding damselflies passing about 50 cm from their perch sites, made this flight and approached them at a distance of 5-10 cm and then hovered fora short time The (Tab. 1, Fig. I). investigatory flights observed are broken down as shown in Table II. This type of flight couldbe considered to be sexual, since when the males have approached the flying insects and found them to be females, they try to form a tandem. In many cases, however, interactions occurred involving than three male in which males more individuals, approached each other at a distance of about 5-10 cm and hovered almost ’’face to face”. As a result of these interferences among males, their perches are defended in some cases but, in other taken over other males. of cases, by The male C. nipponicum shifts from one another within in perch to a day, defending each turn. Consequently, the males out at least about 50 from other individuals. get spaced cm away The time between finding and gripping Table II a female in tandem, i.e. after starting the Objects for which the male C. nipponicum in the of investigatory flight case C. nip- made investigatory flights ponicum, differs between C. melanurum Frequency Objects and C. nipponicum. An experiment in observed which a tethered fluttering female was C. nipponicum.nipponicum, <3$ 152 presented at a distance of 50 cm showed C. nipponicum 2 12 (8)* that the above mentionedtime was 0.85 ± C. melanurum,melanurum. $ 17 0.49 ± S s (x D, n=23) and 2.52 ± 1.51 s (x C. melanurum. 2$ 5 ± SD, n=35) in C. melanurum and C. C. nipponicum. tandem 1 C. melanurum. tandemtandem 3 nipponicum, respectively, when the male Copera annulata,annulala, $ 2 succeeded in forming a tandem.The male • of C. melanurum the female in- Successful tandems grasped stantly whereas the male of C. nip- did after ponicum so hovering for a short time about 10 cm diagonally above her. in Copulation both species is of the long duration type as defined by CORBET (1962). In case of C. melanurum, the duration of the wheel position was 37.2 ± 12.7 min (x ± SD, n=6, range; 18-53 min). Only two measurements were made for C. nipponicum (32 min and 48 min). Oviposition is conducted in a male-upright tandem position. The site used for is oviposition communaland several pairs in tandem were often observed ovipo- as close about 10 from each other. The time siting as cm away during which a tandem pair stayed at the oviposition site was foundto be about 2 hours (range: 0.5-4 hr, n=35 in C. melanurum and n=15 in C. nipponicum). The male and female individuals in both species copulated and oviposited, in general, only once a day. Moreover in both species some individuals, both male and female, copulated and oviposited every day for several consecutive days (Mizuta, unpublished).

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