Odonatologica 15(3): 335-345 September I, 1986
A survey of some Odonata for ultravioletpatterns*
D.F.J. Hilton
Department of Biological Sciences, Bishop’s University,
Lennoxville, Quebec, J1M 1Z7, Canada
Received May 8, 1985 / Revised and Accepted March 3, 1986
series of 338 in families A museum specimens comprising spp. 118 genera and 16
were photographed both with and without a Kodak 18-A ultraviolet (UV) filter.
These photographs revealed that only Euphaeaamphicyana reflected UV from its
other wings whereas all spp. either did not absorb UV (e.g. 94.5% of the Coenagri-
did In with flavescent. onidae) or so to varying degrees. particular, spp. orange or
brown UV these wings (or wing patches) exhibited absorption for same areas.
However, other spp. with nearly transparent wings (especially certain Gomphidae)
Pruinose also had strong UV absorption. body regions reflected UV but the standard
acetone treatment for color preservation dissolves thewax particles of the pruinosity
and destroys UV reflectivity. As is typical for arthropod cuticle, non-pruinosebody
regions absorbed UV and this obscured whatever color patterns might otherwise be
visible without the camera’s UV filter. Frequently there is sexual dimorphismin UV
and and these role various of patterns (wings body) differences may play a in aspects
mating behavior.
INTRODUCTION
Considerable attention has been paid to the various ultraviolet (UV) patterns
exhibited by the Lepidoptera (e.g. SCOTT, 1973). Studies have shown (e.g.
RUTOWSKI, 1981) that differing UV-reflectance patterns are frequently used as
visual in various of behavior. few insect cues aspects mating Although a other
groups have been investigated for the presence of UV patterns (HINTON, 1973; POPE & HINTON, 1977; S1LBERGL1ED, 1979), little informationis available
for the Odonata. ROBEY (1975) noted that pruinose areas on the bodies of male
Erythemis simplicicollis, Libellula cyanea, L. incesta and Pachydiplax longi-
• Dr. the occasion of his 70th Dedicated to M.J. Westfall, Jr. on birthday. 336 D.F.J. Hilton
pennis reflected UV; so too did S1LBERGLIED (1979) for L. pulchella and
Plathemis (= Libellula) lydia, and ROBERTSON (1984) for L. luctuosa, L. pulchella, Pachydiplax longipennis and Plathemis lydia. In addition, SILBER-
GLIED (1979) observed UV reflection from the iridescent body regions of both
sexes of Argia fumipennis. In contrast to these studies which recorded UV
reflectance, SATO (1984) recorded UV absorbance in the wings of Calopteryx
C. and did FRANTSEVICH cornelia, virgo japonica Mnais pruinosa costalis, as
& MOKRUSHOV (1984) for
certain species of Sympetrum.
Members ofthe Odonata are noted for their excellent vision, and those species which have
been investigated are known to
have UV receptors (MENZEL,
1979), especially in the omma- tidia of the dorsal region ofthe eyes (EGUCH1, 1971; GOLD-
SMITH & BERNARD, 1974).
For these reasons it seemed particularly important to sur-
vey the Odonata in order to
determine the extent of UV
patterns in these insects.
METHODS
taken Photographs were of preser- ved specimens in the Odonata collec- tion at Bishop’s University. These
represented 338 species (plus 11
in 118 and 16 subspecies) genera
families ( Tab. 1). All non-Canadian species were obtained from other
and relied odonatologists I upon their 1-3. and ultraviolet of: determinations for species identifica- Figs Daylight (D) (U) patterns
(I) Tetragoneuria costalis male Davidius tion. (M); — (2)
nanus (M); (3) Gomphurusfraternus ( M). Photographic techniques were
adapted from ECUJCHI & MEYER-
-ROCHOW (1983). FERRIS (1972)
ME Pcntax-M and HI IT, (1977). The camera was a Pentax Super with a 50 mm 1:1.7 lens to which
the UV Kodak 18-A UV was attached (for photographs) a ("Wrattcn") filter. The film was Tri-X
Panchromatic 400 ASA. 337 Ultraviolet patterns in Odonata
RESULTS
Fourteen response patterns to UV light have been determined in this survey
in its indicated (Figs 1-14) and each species Table I has UV pattern-type by a
number the which matches its representing Figure most closely own pattern.
Pattern 1 (Fig. 1) has no UV absorbance or reflectance, patterns 2-. 13 (Figs 2-13) show varying degrees of UV absorbance, while pattern 14 (Fig. 14) shows UV
reflectance.
Certains trends can be seen.
Almost all the Coenagrioni-
dae [52/55 (94.5%)] did not
show any UV patterns (like
Fig. 1) and the same was true
to a lesser extent for the
Leucorrhiniinae [11/14
(78.6%)] and Libellulinae
[14/21 (66.7%)]. In contrast,
10/12 (83.3%) ofthe Calopte-
ryginae had a very dark UV-
-absorbing wing pattern (like
Fig. 13), probably due to the
dark pigment(s) typically pre-
sent in their wings. The effect
of wing pigment(s) as UV
absorbers can be seen in
species such as Mnaispruino-
sa in which dark-winged
males had a very dark UV
while pattern paler-winged
females had only a slight wing
darkening in UV. This ten-
dency for wing pigment(s) to
absorb UV was found in a 4-6. and ultraviolet of: Figs Daylight (D) (U) patterns (4)
number of species with fla- vinosa Boyeria (M); — (5) Dromogomphus spinosus fe-
vescent, orange or brown male (F); — (6) Gomphus melaenops (M).
wings or wing patches [e.g.
Lestes eurinus (like Fig. 5), Anax parthenope julius (Fig. 9), Nannophya pyg-
maea (like Fig. II), Sympetrum semicinctum (like Fig. II) and Tholymis
such Libellula tillarga (Fig. 12)]. Species as lydia or Rhyothemis fuligi-
nosa, with very dark wing patches, undoubtedly also absorbed UV in these
pigmented areas. However, they are already so dark in daylight that no change
was evident in UV and thus are considered in Table I not to show UV patterns 338 D.F.J. Hilton
(like Fig. 1). Many Gomphidae had dark UV wing patterns, particularly at
one-halfof the basal one-quarter to the wings (Figs 2, 3 & 7), even though their
in wings were transparent daylight. However, by holding the wings against a white background and looking very closely it was often possible to see a very faint flavescence which corresponded in position to the; UV-absorbing regions of the
in wings. Sexual dimorphism U V wing patterns was frequent. This might just bea small difference in wing darkening (e.g. Polycanthagyna melanictera)I or one sex could have dark in while remained a wing pattern UV the other sex’s wings
transparent [e.g. Gomphus
hodgesi (Fig. 10)].
Euphaea amphicyana was
unusual because it was the only
species to have wings that reflected UV. This occurred in
the basal two-thirds of the hind
wings (Fig. 14) and correspon-
ded with a deep-purple reflec-
tive region which became visi- ble against the very dark brown
wing pigment(s) only when the
wings were held at a particular
angle.
DISCUSSION
In contrast to many butter-
flies (e.g. SCOTT, 1973) (and except for Euphaea amphicy- ana) the Odonata listed in
Table I did not show UV reflectance. GHIRADELLA
(1984) and GHIRADELLA et al. (1972) have shown that UV reflectance in Lepidoptera is Figs 7-9. Daylight (D) and ultraviolet (U) patterns of:
due to optical interference (7) Gomphus pryeri (M); (X) Anax panybeus (M);
within the microscopic struc- (9) Anax parthenopejulius (F).
ture of wing scales. Certain
Coleoptera (POPE & HINTON, 1977)have UV reflectance primarily caused by
microtrichia pigments contained within and other setae but it may also be from dried cuticular secretions (some Tenebrionidacand Buprestidae) or subcuticular pigments (a few desert-dwelling Tenebrionidac). Because Odonata lack lepidopteran-type scales or the surface cuticular processes typical of some beetles. Ultraviolet patterns in Odonata 339
this be the few reflect UV. The may reason why so species major exceptions are those which exhibit species pruinosity. ROBERTSON (1984), using a spectrophotometer, determinedthat pruinosity strongly reflected UV. BYERS
(1975), ROBEY (1975) and SILBERGL1ED (1979) recorded the tendency for white or bluish pruinose areas to have an enhanced whiteness when
UV filter. I noticed photographed through a also this effect but did not designate it in as a separate pattern-type TableI. Both BYERS (1975) and S1LBERGL1ED
that UV reflectance of (1979) suggested pruinose areas might be a Tyndall effect and BYERS (1975) observed that when the wax particles which comprise the pruino- sity are removed by organic solvents the Tyndall effect is permanently destroyed. Due to the widespread practice by odonatologists of soaking captured Odonata in acetone for 24 hrs in order to preserve colors, many museum speci- mens will have lost their pruinosity and thus not reflect UV from these areas.
It is not known what pigments are responsible for the flavescent, orange or brown wings (or wing pat- ches) that strongly absorbed
UV in many Odonata. How-
in ever, plants, BREHM &
K RELL (1975) and THOM P-
SON et al. (1972) have determinedthat flavonoids in the nectar guides of some flowers are responsible for 10-12. (D) ultraviolet Figs Daylight and (U) patterns of: these regions absorbing UV. (10) Gomphus hodgesi (F + M); — (II) Sympetrum As noted SILBERGLIED Occidentale fasciatum (F + M); — (12) Tholymis tillarga (1979), arthropod cuticle ab- (F + M). sorbs UV. This is why
Odonata without pruinose regions had dark bodies in UV and their daylight color-patterns were obscured (Figs 1U-I4U).
One cannot generalize about the role UV patterns play in odonate behavior. However, it is obviously not a species or sex recognition factor for some (e.g. 340 D.F.J. Hilton
Table I
of Odonata which examined for the of Species were presence UV patterns
UV 22 2 . pattempattern type . UV „ , patternvpattem type Species'Species 1 Species'Species 1 3£ 9Ç 3 9
COENAGRIONIDAECOHNAGRIONIDAE E.£ hageni < lI I1
Amphicnemidinae !Ischnuraschnüra asiaticaasiatica lI I
AntiagrionAniiagrion gayi II I1 I.1. cervula 1
Pseudagrioninae I.1. elegans I I1
Ceriagrion latericumlalericum I.1. prognathaprognalha lI
ryukyuanum iI I1 I.1. pumilio lI
C. melanurum Ii I1 I.1. ramburii lI
C. nipponicumnipponicum Ii I. senegalensis Il I1
1. PseudagrionPseudagrionp. pilidorsum Ii I1 I. verticalisverlicalis lI I1
TelebasisTelehasis salva I Oxyagrion terminale I I1
Coenagrioninae Agriocnemidinae
I I1 lI I1 CercionCerdon calamorum \ Agriocnemis feminaoryzae
C. hieroglyphicum I I1 Mortonagrionselenionselenion lI I1
C. plagiosum Ii I1 Argiinae
C. sexlineatumsexlineatum iI I1 Argia moeslamoesta lI I
C. sieboldiisieholdii iI I1 A. oenea 4
Chromagrion conditumcondilum iI I1 A. pulla I1
Coenagrion angulatum iI I1 A. tezpi 13
C. armatumarmalum iI I A. violacea I1 I1
C. concinnumcuncinnum I I PLATYCNEMIDIDAEPLATYCNEM1DIDAE
C. hastulalumhaslulalum I I1 Calicnemidinae
C. puella iI I1 ■Coeticciaf.Coeliccia f. flavicaudaJlavicauda I1
C. pulchellumpulchellum iI I1 Platycnemidinae
C. resolutumresolulum I I1 Copera annulataannulala I1
C. vernale I I 1 C. marginipes I1
Erythromma najas I I1 C.C. tokyoensistokyoensis 2
Nehalennia gracilis iI I1 PlatycnemisPlalycnemisfoliaceafuliacea sasakii I1 I1
N. irene Ii I1 P. pennipes 4 55
N. speciosaspeciosa I I1 PSEUDOSTIGMATIDAE
PyrrhosomaPirrhosoma nymphulanymphula I I1 MecistogasterMecislogasierlinearislinearis 4
Ischnurinae Megaloprepus caerulatuscaerulalus I1
Acanthagrion gracile iI LESTIDAE
Aciagrion hisopa i1 Lestinae
A. migramigrai unitum iI I1 Archilestes grandis 5 4
AmphiagrionAmphiagrionsaucium iI I 1 ChalcolesiesChalcolesles viridisviridis 4 4
Cyanallagmainterruptainterrupta 4 4 Lestes congener I1 I1
Kn I1 I1 L.L d. Enallagmaallayma aspersum d. disjunctusdisjonctas I1 I1
E. I1 I1 L.L 1 horeale dryasdryas I I1
E.£ carunculatumcarunculaium I1 E.L eurinus 5 5
E.£ civilecivile I1 L./_ forcipatus I1 I1
E.£ cyathigerumcyathigcrum I1 L.£ japonicusjaponicus 4
E deseni /.. E. deserti yezoensis I1 L. rectangularisreelangularis I1
E. I1 I1 4 4 ebriumehrium L.L sponsa
E.£ exsulans I1 L./_ temporalis 4 4
and ' Genera species arc listed alphabetically;higher categories are according to Davies (19X1).
2 Numbers refer the to corresponding figures; a blank indicates this sex was not available for
examination. Ultraviolet in patterns Odonata 341
2 UV 2 patternpattern type UV pattern typetype’ Species-Species 1 Species 1
o<5 29 5S 92
LL. unguiculatus iII1 A. eremitaeremila I 1I
A. L. virens 4 4 cyanea 4 4
L. viridis I1 A. i. interruptainterrupla I1 I1
Sympecmatinae A. i. lineatalineal a I1 909
4 44 IndolestesIndolesles cyanea A. juncea I1 44
I. peregrinus I1 44 A. j.J. americana I1 5
Sympecma paedisca II1 1 A. multicolor 1
PSEUDOLESTIDAE A. nigronigroflavaflava 4 4
HypolestinaeHy polestinae A. palmata I1 1I
aculeataaculeata RhipidolestesRhipidolesies a. 4 1 A. sitchensis I1
R. aculeataaculeala hiraoi II1 1 A. subarctica I1 I1
R. okinawanus 4 A. lubercuUferatuberculitera I1 I1
CALOPTERYGIDAE A. umbrosa occidentalis I1 4
Calopteryginae A. u. umbrosa 1 4
CalopteryxCalopleryx amalumamatum 13 A. verticalisverticalis I1
C. atrataalrala 13 13 Anaciaeschna martini 5 5
C. Corneliacamélia 13 13 Anax juniusJunius 5 5
C. maculatum 13 13 A. longipes 6
C. 13 13 A. splendens A. n. nigrqfasciatusnigrofasciaius 5
C. virgo 13 13 A. panybeus 8
MatronaMalrona basilarishasilaris 13 13 A. parthenopepanhenopeJuliusjulius 9 9
M. basilarishasilaris japonica 13 Gynacanthajaponica 4 55
costalis Mnais 13 13 PolycanthagynaPoh canthagynamelanictera 5 4
M. pruinosa nawai 13 44 Brachytroninae
M. p.p. pruinosa 13 44 Aeschnophlebia anisoptera 13 I1
Neurobasis c. chinensis 13 13 A. longistigmalongisligma 55
Psolodesmus mandarinus BasiaeschnaBasiaeschna janatajanata 4
44 dorotheadorothea I1 Boyeria grafiana 4 4
VestalisVeslalis apicalis 4 1 B. maclachlani 11
Hetaerininae Hetaerininae B. vinosa 4
MnesareteMnesareie I1 1 pudica GGomphaeschnaomphaeschna furcillatafurcillaia I1
CHLOROCYPHIDAE Oligoaeschnapryeripryeri 4
Chlorocyphinae GOMPHIDAE
Rhinocypha bisignatahisignaia 13 44 Gomphinae
EUPHAEIDAE Arigomphus maxwelli 1I
Euphaea amphicyana 1414 A. villosipes I1 I1
E. formosa 1313 DavidiusDavidius fujiamafujiama 2
E. 1313 13 yayeyamana D. m. moiwanusmom anus 2
b.h. brevicaudahrevicauda 4 44 BayaderaBayaderu D. nanus 2
EPIOPHLEBIIDAE Dromogomphus spinosus 5
I1 I1 EpiophlebiaEpiophlehiasuperslessuperstes ErpetogomphusErpelogomphus designatusdesignaius 4
AESHNIDAE Gomphurus fraternusfralernus 3
Aeshninae Gomphus australis 3
Aeshna canadensis I1 44 G. borealis 2
coerulea A. I1 G. brevis 2 2
A. constconstrictarida II1 1 342 D.F.J. Hilton
2 UV pattern type2 UV 2 pa.tenMype* P a«e™‘V^type Species'Species 1 Species 1 3 $ pattern^
G. cavillariscavil/aris l1 Sieboldius albardae 6
G. crassus 2 2 Lindeniin^e
G. descriptus 5 4 IctinogomphusIclinogomphus clavalusclavatus 5
G. diminutusdiminulus 2 I.1. pertinaxperlinax 6
G. exilis 4 4 PETALURIDAE
G. graslinellus 3 7 Petalurinae
G. hodgesi 10 10 TachopteryxTachopteryx thoreyi 2
G. hybridushybridas 2 Tanypteryginae
G. ivae I1 Tanypteryx pryeri 2
G. melaenops 6 CORDULEGASTR1DAE
G. minutusminutas 3 3 Chlorogomphinae
G. oklahomensis 6 ChlrogomphusChlorogomphus b. brunneus 13
G. ozarkensis 7 5 C. brunneus costaliscoslalis 4 7
G. pallidas I1 C. suzukii 5
G. plagiantusplagialus 11 Cordulegastrinae
G. potulentuspolulenlus 11 AnotogasterAnologaster sieboldii 5 7
G. pryeri 7 CordulegasterCordulegasler bidentatusbidenlalus 5
G. sandriussandrius 4 C. bilineata 4
G. septima 7 C. boltonibohoni 5
G. spicatus I1 I1 C. diastatopsdiastalops 5 6
G. townesi 11 C. erronea I1
G. vastus 3 C. maculatus 5 5
G. villosipes I1 C. obliquusobliquas 6
G. 1 G. vulgatissimusvulgalissimus 4 C. sayi I
Hylogomphus abbreviatusahhreviatus 2 CORDULIIDAE
H.H. geminatus I Corduliinae
! amurensis 4 Lanthusjamhas albistylus 3 33 Cordulia aenea amurensis
L.L fujiacus 2 C. shurtleffishunleffi I1 I1
L.L parvuluspan ulus 2 Dorocordulia libera 4 4
Nihonogomphus viridusviridus 4 Epicorduliaprinceps I1
Onychogomphusforcipatusforcipatus I1 EpithecaEpitheta canis 4 5
O. viridicostus 2 E.£ I1 I1 cynosuracynosura
Ophiogomphus carolus 4 E.£. marginata 5
SinogomphusSinvgomphusßavolimbalusflavolimbatus 2 2 E. princeps I1 I1
StylogomphusSlylogomphussuzukii 7 E.£. spinigera 4
Stylurus oculotusoculatus 3 Helocordulia selysia I1
TrigomphusTrigomphuscitimuscilimus tabeitahei 5 2 H. uhleri I1
T. interruptusimerruptus 2 HemicorduliaHemit ordulia okinawensisokinau ensis 5 2
T. melampus 4 4 Neurocordulia alabamensis I1
T. ogumai 2 NA.. xanthosoma 4 4
Gomphoidinae N. yamaskanensis 5 4
AphyllaAphvlla williamsoni 4 Rialla villosa 4
ProgomphusProgumphus belleihellei 1 SomatochloraSomaiochlora alpestrisalpestris 5
S. 4 P. obscurusohscurus 1 S. arcticaarctica
Hageniinac S. clavata 4
HageniusHaxenius brevistylushrevistylus 6 6 S. elongata I1 Ultraviolet in patterns Odonata 343
UV 2 UV 2 pattern typetype’ patternpattem typetype^ Species 11 Species' $<3 92 $<5 2$
S51. ßavomaculalaflavomaculata 4 LL. hudsonica iI I1
S. forcipata 4 L.L intactaIntacta Ii I1
S. franklini 7 LL. proximaproximo Ii I1
S. graeseri aureola 4 I1 L.L rubicundarubicunda I1
S. hudsonica 4 4 Libellulinae
linearis S. 5 UbellulaLibellula angelina iI
S. S. metallicamelallica 4 4 L.L axilena i
S. minor S. minor I1 1 LL. comanche iI
S. semicircularis 4 5 L.L fulva iI
S. tenebrosalenebrosa I1 4 L.L julia Ii I1
S. uchidai 11 L.L luctuosa I1
S. viridiaenea 4 4 LL. lydiaIvdia iI I1
costaliscoslalis TetragoneuriaTeiragoneuria 1 L. pulchella iI I1
T T. spinosa I L.L quadrimaculataquadrimaculala iI I1
Gomphomacromiinae LL. asahinai I q. i I1
Gomphomacromia Lokia coryndoni iI I
paradoxa 4 2 Lyriothemis elegantissimaeleganlissima Ii I1
MACROMIIDAE L.L pachygastra 2 2
Didymops floridensis I1 OrthemisOrlhemis ferruginae 4
D. transversa I1 I1 OrthetrumOnhelrum albistylumalbisn lum
EpophthalmiaEpophlhalmiaelegans 4 speciosum I1 4
Macromia a. amphigenaamphigena 6 O. cancellatumcancellalum I1 I1
M. georgina I1 O. coerulescens 4
illinoiensis M. 1 O. j.j.japonicumjaponicum 44 4
SYNTHEMISTIDAE O. luzonicum 66
euslralacla Synthemis eustralacta I1 4 O. pruinosum neglectum I1
LIBELLULIDAELIBELLUL1DAE O. s. sabinasahina I1
Brachydiplacinae O. triangularetrianguläre melania 5 5
Brachydiplax chalybeachalyhea Sympetrinae
flavobittaflavohitta 5 AcisomaAcisuma I p. panorpoides 1
1 1 ChalcostephiaChalcoslephiaflavifrons Brachythemis contaminatacontanunata 9 4
Micrathyria hypodidyma I Crocothemis serviliaservi/ia I1 4
Nannophyapygmaea II II Deielia phaon 4
Uracis fastigialfastigiataa 5 I1 Diplacodes trivialis I1 I1
Leucorrhiniinae ErythemisErylhemis simplicicollis I1 4
Cannacria batesihatesi 4 ErythrodiplaxErylhrodiplaxfunereafunerea I1
CelithemisCelilhemis berthaherlha I1 E.E fusca I1
C. elisa I1 E.£. umbrata I1 4
C. C. eponina 13 13 Lepthemis vesiculosa I1
C. •verna I1 verna Neurothemis r. ramburiramhuri I1 I1
Leuœrrhinia Leucorrhinia albifronsalhifrons I1 I1 Pachydiplax longipennis 4 4
borealishurealis LL. I1 I1 SympetrumSympetrwn bacchahaccha
L.L dubiaduhia I1 I1 matutinum I1 I1
L. d. orientalis 1 S. 1 S. coscostiferumliferum 4 4
LL. frigidafrigida I1 2 S. croceolumcroceotum II II
L. glacialis I1 I1 S. danae I1 I1 344 D.F.J. Hilton
2 UV UV pattern type 2 _ patternpattern type , „ SpeciesSpecies'' Spec.es'Species' <5 9 <3 9$
S. darwinianum iI I1 S.5. uniforme 5
S.S. depressiusculumdepressiusculum 4 I1 S. vicinum 5 44
S. 44 I1 S.S. e. eroticumerolicum I1 2 S. vulgavulgaiumtum
S. flaveolum II Trithemistinae
S.S. frequens I1 I1 Dythemis veloxvelo.x I I1
S. gracile I1 I1 Macrothemi.sMacrothemis imitansimitons 4
S. infuscatum 11 4 Pseudothemis zonata 4 4
S. internum 4 4 Trithemis aurora I1
S. kunckeli I1 2 Palpopleurinae
S. maculatummaculalum 4 4 Palpopleura lucia portia I1 I1
S. obtrusumohlrusum I1 I1 Perithemis domitiaJomilia 13
S. occidentale fasciatumfascialum IlII II P. seminole 13 I1
S. pallipes I1 I1 Trameinae
S. parvulum 2 2 Miathyria marcella 44 4
S. pedemontanumpedemomanum I1 I1 PantalaPanlala flavescens I 1 4
4 I1 I1 S. p. elatumelalum 5 Rhyothemis fuliginosa
risi R. 13 13 S. r. 5 R. variegatavariegata imperatriximperatrix 13 13
rubicundulumruhk undulum 12 12 S. I1 TholymisTho/ymis Itillargaillarya
S. sanguineum I1 Urothemistinae
S. semicinctum IlII II MacrodiplaxMacrodipla.x balteatahalleala I1 I1
S. 4 I S. s. speciosumspedosum 1 Zygonychinae
S. striolatum I1 I1 ZigonyxZygonyx takasagotakasago I1
S. s.s. imitoidesimiioides 5 I1
most Coenagrionidae), whereas
species like Pachydiplax longi- pennis and Gomphus hodgesi
exhibit sexual dimorphism in
body UV-reflectance and wing
UV-absorbanee, respectively.
One would expect such species
to utilizeU V patterns in aspects
of their mating behavior, just
as ROBEY (1975) has shown
for Pachydiplax longipennis.
Figs 13-14. Daylight (D) and ultra-
violet (U) patterns of: (13) Calopteryx
amatum (F); (14) Euphaea am-
phicyana (M). Ultraviolet patterns in Odonata 345
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