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Libellulidae) Dragonflies Rely Mainly on Significantly Important in the Set

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Libellulidae) Dragonflies Rely Mainly on Significantly Important in the Set Odonalohgica 13 (3): 335-350 September I. 1984 Visual stimuli releasing attack of a territorialmale in Sympetrum (Anisoptera: Libellulidae) L.I. Frantsevich and P.A. Mokrushov Schmalhausen Institute of Zoology, Academy of Sciences of the Ukrainian SSR, Lenin Str. 15, USSR-252000 Kiev-30 Received May 24, 1983 / Revised and Accepted October 25, 1983 released rival male Visually responses to a were studied in territorial males of 5. S. and sanguineum, vulgatum, S. flaveolum S. pedemontanumusing a fishing-rod to present synthetic models. Film discs with the dark body centre and the concentric turned wing spot (if necessary) out to be most effective. Position and presence ofthe difference. abdomen made no The species devoid of wing patterns (S. sanguineum, S. other vulgalum, S. danae) are unable to recognize each by appearance. Probably, they utilize other means of interspecific isolation. The species decorated with wing distinguish spots certainly conspecific males by the specific feature of pattern: an orange pericentral fleck in 5. flaveolum, a subapical band in S.pedemonlanum. The shows Contrast in UV in S. wing pattern stronger than long wave rays. sanguineum showed moderate aggression against patterned models, but was less selective to various models than the The ofthe neural two patterned species. properties perceptor detecting the configurationalvisual stimulus are discussed. Distant recognition of the its conspecific mate by pattern is more reliable if the latter is restricted to basal flecks and bands, while odonate or apical thawing middle remains clear. Most spe- cies with wing decoration support this rule. INTRODUCTION Dragonflies rely mainly on their visual sense in their interaction with the environment. Visual stimuli are significantly important in the set of isolating from mechanical The of factors, ecological to ones. role the visible pattern in male-female recognition has been investigated by many authors. There are, fewer however, experiments and fieldobservations concerning the problem ofthe territorial male its recognizing rival. (Tab. I). The distant recognition of a conspecific rival is energetically advantageous, for the male avoids useless fights with strange species. Some observers stated that territorial odonate males lack 336 L.l. Frantsevitch & P.A. Mokrushov Table I Visible features of a conspecific male for dragonflies Meaningfulfeatures Species Method Body Body Wing mo- Wing co- Wingdis-dis- Abdomen References size loration colourcolour vementsvenants loration play display HetaerinaHelaerina americana. field observation JOHNSON, 1961 H. titiatilia + CalopteryxCalopieryx painting of + HEYMER, 1973 live insects Libellulapulchella field observation + PEZALLA, 1979 L.L luctuosa,lucluosa. ErythemisErvlhemis painted insects ANDREW,ANDREW. 19661966 on a rod + + + simplicicollis on a fishing Leucorrhinia dubia.dubia. live insects on PAJUNEN, 1962. 1964 L.L rubicunda a fishing rod + + L.L caudalis field observation + + PAJUNEN. 1964 field OrthetrumOnhe!rum albistylumalbislylum field observation + ITO,1TO, I960 1955 PerithemisPerilhemis tenera painting of JACOBS,JACOBS. live insects + PlathemisPlalhemis lydiaIvdia painting oflive JACOBS 1955; insects, field CAMPANELLACAMPANELLA& WOLF. 1974 observations + + + Nesciothemis painted insects PARR & PARR, 1974 nigeriensis on a fishing rod + ++ PachydiplaxPachvdiplax field observation. JOHNSON 1962; longipennis spectrophotometry ++ ROBEY,ROBEY. 1975 1983 CorduliaCordulia aenea dead insects flight abdomen UBUUBUKATA,KATA. 1983 on a rod direction shapeshape discrimination of reactions proper specific features and perform aggressive of families: viridis against dragonflies other species, genera, and Lestes (DREYER, 1978), Tanypteryx hageni (CLEMENT & MEYER, 1980), Cordulia aenea (UBUKATA, 1975), Leucorrhinia caudalis (PAJUNEN, 1964), Libellula quadrimaculata (ZA1KA, 1977), Orthetrum cancellatum (KRONER, 1977), Micrathyria atra (MAY, 1980), Pachydiplax longipennis (ROBEY, 1975) etc. of for Anisopterans the genus Sympetrum seem to be a promising subject studying visual releasers of male aggression. The site of observation was 5 of that viz. S. populated by species genus, S. sanguineum (Mull.), vulgatum (L.), S. flaveolum (L.), S. danae (Sulz.) and S. pedemontanum (Allioni). Their find seasonal and spatial distributions overlap broadly. It was possible to some places where males of 4-5 species establish their territories at the same time. S. sanguineum males share the habitat with males of each species. METHODS Field experiments were carried out in the vicinity of Kiev, from July to September 1981-1982, from 12.30 hrs until 16.30 hrs local time on sunny days. Presenting the models of dragonflies by the fishing-rod technique (ST. QUENTIN, 1934), we scored 5054 models. All models made of such responses to 72 different were synthetics as resistors, of stained film. coloured pieces insulation, and transparent or acetate certain times, from male another, then The observer showed a model up to ten proceeding one to in changed it to another one. A specific model, standard for the given species, was included each set of experiments to check the seasonal and weather variation in male activity. The difference in Visual stimuli in male Sympetrum 337 different models the 2 test. Models within 0.5-2.0 m responses to was analyzed using x were presented males into four of a resident male. Responses of fall types, classified as: the model Attack: model within 5 aerial manoeuvresof (1) a was approachedto cm, up to aclash; were repeated. themodel to distance of than 10-15 then (2) Survey: a straight exploratory flight to up a more cm, in-tandem return. Territorial males approached models frombelow. Attempts to attain an position were witnessed only twice. been (3) Escape: flight from the territory without any attempt to return. Escape could have elicited also by the movements of the observer. (4) Indifference: no take off. Transmission spectra ofdragonfly wings were recorded by the Specord-Zeiss spectrophotometer. UV fluorescent under black Wing transmission in was visualized by placing wings ona yellow plate light. from Bouin fixed Directions of surfaces and Eye segments were excised by hand preparations. with under dissection axial structures were measured a goniometer a microscope. RESULTS SPATIAL AND TEMPORAL DISTRIBUTION OF SPECIES in According to several years’ observations, Sympetrum imagines emerge the following order: S. flaveolum, S. sanguineum, S. pedemontanum, S. danae, and S. vulgatum. S. flaveolum territo- rial males are the first to undergo their maturation (at the end of July), and almost without delay followed they are by S. sangui- neum and 5. pedemondanum. The males of S. vulgatum start mating at the end of August. All five active in species are Septem- ber and survive till the first frosts in October, though the relative abundance of 5. Jlaveolum and S. pedemontanum apparently de- of creases at the end the flying season. 5. sanguineum occupies territo- ries at the side of water, in bogs, in wet and in dry riverside meadows, Fig. I. Height of perching above the ground (cm) in the first three habitats being territorial males of S. sanguineum and S. flaveolum. shared with S. fiaveolum. The (Temperature 21-22°; — 170 and 62 records; — the of areas of the histograms are normalized to unity). males 5. fiaveolum are more aggressive. They reside présuma- 338 L.I. Frantsevitch & P.A. Mokrushov the surface bly on or just above the ground. S. sanguineum males tend to perch somewhat higher on the grass oron shrubs (Fig. 1 ). When the weather is cool they descend to the surface. The males of S. pedemontanum are rather scarce. They gain territories on the sides of clear slow runnels or on the vegetation above the where reside of water, S. sanguineum males as well. The males S. vulgatum territories in amidst occupy the paths dry bottom meadows; other Sympetrum them. failed find the sites where species rarely accompany Unfortunately we to territorial males of S. danae the and swarm, though mass emergence, hunting, oviposition of this species were easily observable. INTERSPECIFIC DIFFERENCES OF MALES The features of five listed in Sympetrum species are Table II. An experienced observer can identify the species of Sympetrum either settled or even on the wing, practically without error. Table il Characteristics of Synipelrum males observable in the field Species Body Wing Body Wing plate Wing pattern length (mm) spanspan (mm) colour colour negligible sanguineum 33.6 ± 1.2 55.0 ± 16 blood-redblood-red slightly dusky vulgatum 39.3 ± 2.02.0 58.8 ± 1.41.4 cherry-red brownish absent hind flaveolum 33.633.6 ± 0.7 57.3 ± 2.4 orange-red clear basal yellow spot on wingwing(27% ofofwing area) danae 31.731.7 ± 0.40.4 49.449.4 ± 1.3 blackblack clearclear absent pedemontanumpedemonianum 31.831.8 ± 0.60.6'< 53.7 ± 0.7 coral-red clearclear subapical brown band (12%(12% ofwingwing area)area) The histogram above Figure 2 illustrates the short-wave shift of colour vision in odonates re- lative Thus to man. one may expect that wings of dragonflies look different in UV rays and in visible transmission rays. The of shown spectra wing plates are in Figure 2, while Figure 3 illus- Fig. 2. Transmission spectra of male - Sympelrum wings: (A) S. itanue: (B) .S'. vulgalum: — (C) S. flaveolum. transpa- — S. rent part; (D) jlaveolwn. orange mark; — (E) .S. pedennmlanum,subapical band. — The bars above the graph show the spectral sensitivity maxima of dorsal in cited eye receptors odonates. by LAVOI E-DOR NIK et al. (19X1). Visual stimuli in male Sympelrum 339 Fig. 3. Sympetrum wings against the yellow fluorescent plate; (A): in yellow light; (B) in UV light in trales wing appearance UV and long wave rays. The wing decorationof S. flaveolum and S. pedemontanum looks much more for for contrasting a dragonfly than a human observer. Wings of.S. vulgatum and S. look dark with sanguineum compared the transparent wings of S. danae. BODY COLOUR Dragonfly models were prepared from insulation of different colours. They mimicked the of body shape 5. sanguineum: body length 34 mm. "thorax” size 12x6 mm, wing surface 10x40 mm. Responses ofS.JIaveolum and 5. sanguineum males to these models are shown in Figure 4.
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