42 Ameiican Lnromologisr Sjring 200S

D
uiing liglr, insecr vings Lenu anu rvisr
uiamarically, anu rle insranraneous, rliee-
uimensional slae o vings may inluence
many asecrs o liglr eioimance. Insecrs lave
lirrle conriol ovei rlis Lenuing anu rvisringving
ueoimarions aie laigely assive anu aie conriolleu
iimaiily Ly rle aiclirecruie anu mareiial ioei-
ries o rle ving. Hovevei, oui unueisranuing o
lov insecr ving uesign aecrs lexiLiliry anu as-
sive ving ueoimarion iemains limireu. Heie, ve
uiscuss lov insecr ving venarion aecrs oveiall
Lenuing sriness, lov sriness vaiies rliouglour
vings, anu lov rlese earuies o ving uesign aecr
assive Lenuing.
1le arrein o suoiring veins in insecr vings
vaiies viuely among insecr oiueis anu amilies.
Given rle laige lylogeneric clanges in ving vena-
rion arrein (Iig. 1), one miglr execr insecr vings
ro uislay laige meclanical uieiences rlar voulu
aecr rleii ueoimaLiliry uuiing liglr.
We examineu rle ielarionsli Lerveen insecr
ving lexiLiliry anu venarion Ly measuiing lexuial
sriness (I) anu quanriying venarion arrein in 16
insecr secies iom six oiueis. Ilexuial sriness is a
comosire measuie o rle oveiall Lenuing sriness o
a ving, comLining rle mareiial ioeiries o rle ving
(, Young`s mouulus) anu rle geomeriic uisriiLurion
o rlis mareiial (I, seconu momenr o aiea).
We measuieu oveiall I o vings in rle sanvise
uiiecrion (iom Lase ro ri) anu rle cloiuvise uiiec-
rion (iom leauing ro riailing euge) Ly eioiming
sraric Lenuing resrs. We arracleu eacl ving ar rle Lase
(oi leauing euge), alieu a knovn oice ar rle ri
(oi riailing euge), anu measuieu rle uislacemenr o
rle ving. We rlen calculareu I virl a simle Leam
equarion (see ComLes anu Daniel 2003a). We also
uigirizeu rle ving venarion o eacl secies anu ue-
iiveu hve measuies o venarion arrein (ComLes anu
Daniel 2003a). 1o iemove rle eecrs o lylogeny,
ve calculareu sranuaiuizeu inueenuenr conriasrs o
venarion anu sriness measuiemenrs anu examineu
rle coiielarions Lerveen rlese conriasrs.
Oui measuiemenrs slov rlar I is sriongly
coiielareu virl ving size (Iig. 2), Lur rle uerails
o venarion arrein uo nor aeai ro aecr oveiall
lexuial sriness (no signihcanr coiielarions veie
ounu Lerveen conriasrs o ving venarion arrein
anu I). 1le measuiemenrs also ieveal a laige
anisorioy, oi uieience, Lerveen sanvise anu
cloiuvise lexuial sriness, sanvise I is 1 ro
2 oiueis o magniruue giearei rlan cloiuvise I
in all secies resreu (Iig. 2).
1o uereimine lov ving sriucruie may conriiL-
ure ro rlis arrein o sriness anisorioy, ve cie-
areu a simliheu hnire-elemenr mouel o a ManJuca
(lavkmorl) ving. 1o cieare rlis mouel, a uigirizeu
Ilexural Stiffness in Insect Wings:
Effects of Wing Venation and Stiffness
Distribution on Passive Bending
S. A. Ccnbes anJ T. I. Daniel
Fig. 1. Drawings of forewings from insects used in this study, arranged on the
phylogenetic tree used to calculate independent contrasts. Veins are drawn at
actual thickness; wings are not shown to scale. Genus and species names (when
known) are shown under each wing, and orders are listed at their branching
points (from Combes and Daniel 2003a).
43 Ameiican Lnromologisr Vclune S1, Nunber 1
image o rle ving vas uiscierizeu, oi Lioken uovn
inro numeious elemenrs (squaies oi riiangles),
eacl o vlicl vas given a ceirain rlickness anu
Young`s mouulus (see Iig. 4A, ler). 1lis comu-
rarional moueling reclnique allovs us ro solve a
comlex marlemarical ioLlem (lov rle enriie
ving iesonus ro oices) Ly Lieaking ir uovn inro
a numLei o simlei ioLlems (lov eacl elemenr
inreiacrs virl irs neiglLois). Oui goal vas nor ro
ieiouuce rle Lelavioi o a ieal ManJuca ving,
Lur iarlei ro cieare a geneial mouel ro exloie lov
auuing oi sriengrlening veins in ceirain iegions o
rle ving aecrs oveiall lexuial sriness.
We incieaseu rle Young`s mouulus o ceirain
veins in rle mouel, alieu a sraric oice, iecoiueu
rle iesulring uislacemenr, anu rlen calculareu
oveiall I o rle viirual ving. We ounu rlar
auuing oi sriengrlening veins in rle leauing euge
o rle ving uiamarically incieases sanvise lex-
uial sriness, geneiaring rle sanvisecloiuvise
anisorioy measuieu in ieal vings. 1le acr rlar
clusreieu oi rlickeneu veins aie ounu in rle leau-
ing euge o neaily all insecr vings suggesrs rlar rlis
sriness anisorioy may Le a common uncrional
earuie among insecrs.
Alrlougl oui measuiemenrs o oveiall I io-
viue inoimarion aLour rle scaling anu anisorioy o
Lenuing sriness, rle insranraneous slae o a la-
ing ving is inluenceu nor only Ly oveiall sriness,
Lur also Ly rle uisriiLurion o lexiLiliry rliouglour
rle ving. 1o uereimine lov ving lexiLiliry vaiies
sarially, ve measuieu vaiiarion in lexuial sriness
along rvo axes o rle ving (sanvise anu cloiu-
vise) in rvo seciesrle lavkmorl ManJuca sexta
anu rle uiagonly Aeslna nulticclcr.
We hisr measuieu conrinuous uislacemenr Ly
eioiming sraric Lenuing resrs as Leoie, Lur also
illuminareu vings virl lines o lasei liglr along rle
Lenuing axis anu lorogialeu vings Leoie anu
arei loauing virl a oinr oice. 1le uieience in
rle osirion o rle lasei line in rlese lorogials
ioviueu us virl rle conrinuous uislacemenr o
rle ving. 1o calculare I along rlis axis, ve useu
a conrinuous veision o rle Leam equarion (see
ComLes anu Daniel 2003L) anu ieaiiangeu rle
equarion so rlar ve veie solving oi uislacemenr
(along rle ving) as a uncrion o rle I uisriiLu-
rion. We oseu vaiious alreinarives oi lov lexuial
sriness miglr vaiy along rle ving (ieiesenreu
Ly simle marlemarical uncrions), ieuicreu rle
arreins o uislacemenr rlar voulu iesulr iom
loauing, anu rlen uereimineu vlicl I uisriiLurion
iouuceu a arrein o uislacemenr mosr similai
ro rlar measuieu in a ieal ving.
Oui measuiemenrs slov rlar lexuial sriness
ueclines exonenrially iom ving Lase ro ri anu
iom rle leauing ro rle riailing euge in vings o
M. sexta (Iig. 3A). In auuirion, ve ounu rlar
cloiuvise lexuial sriness ueclines ai moie
slaily in male morls rlan in emale morls, anu
rle vings o male anu emale morls uislay a laige
uoisal/venrial uieience in I. Neasuiemenrs on
A. nulticclcr slov rlar sriness also ueclines ex-
onenrially in rlis secies (Iig. 3B), uesire laige
uieiences in ving slae anu venarion arrein Le-
rveen lavkmorls anu uiagonlies. 1lis similaiiry
Fig. 2. Flexural stiffness
versus span/chord length
in 16 insect species.
ndividuals of each
species are plotted in the
same color. Axes are on
a logarithmic scale. A.
Spanwise fexural stiffness
versus wing span; for log-
log transformed data, y =
2.97x + 0.08, r
2
= 0.95. B.
Chordwise fexural stiffness
versus chord length; for
log-log transformed data
y = 2.08x - 1.73, r
2
= 0.91
(from Combes and Daniel
2003a).
Fig. 3. Flexural stiffness distribution in wings of Manduca sexta and Aeshna multicolor,
and in fnite element models of M. sexta wings. n each graph, spanwise fexural
stiffness is shown above (longer lines) and chordwise fexural stiffness below (both
on a log scale); dorsal measurements are in black and ventral measurements in
gray. Each line within these groups represents the fexural stiffness distribution of a
different individual (from Combes and Daniel 2003b).
0.005 0.01 0.1 0.002 0.01 0.04
A B
Wing span (m) Chord length (m)
S
n
a
p
w
s
i
e
I
E
(

N
m

2
)
Odonata
Hymenoptera
Diptera
Lepidoptera
Isoptera
Neuroptera
Aeshna
Pachydiplax
Lestes
Ischnura
Zootermopsis
Hemerobius
Pepsis
Sceliphron
Bombus
Tipula
Villa
Eristalis
Calliphora
Manduca
Ochlodes
Pieris
1s10
3
1s10
4
1s10
5
1s10
6
1s10
7
1s10
8
d
r
o
h
C
w
s
i
e
I
E
(

N
m
2
)
1s10
3
1s10
4
1s10
5
1s10
6
1s10
7
1s10
8
44 Ameiican Lnromologisr Sjring 200S
in oveiall sriness uisriiLurion Lerveen rvo veiy
uieienr secies suggesrs rlar a slai uecline in I
rovaiu rle ri anu riailing euge may Le a common
earuie o many insecr vings.
1o uereimine lov rlese sriness arreins aecr
assive ving Lenuing, ve cieareu rvo simliheu
hnire elemenr mouels virl rle same geomeriy, Lur
virl uiamarically uieienr sarial arreins o lex-
uial sriness. In one mouel, values o Young`s mouu-
lus uecline uiagonally rovaius rle ri, iouucing an
exonenrially ueclining I uisriiLurion similai ro rle
arrein measuieu in ieal vings (mouel 2, Iigs. 3A,
4A). In a simlei mouel virl hxeu values o Young`s
mouulus oi veins anu memLianes, rle sarial ar-
rein o I is veiy uieienr iom rlar measuieu in
vings (mouel 1, Iigs. 3A, 4A).
We comaieu arreins o Lenuing in rle rvo
mouels in iesonse ro a sraric oinr loau alieu
ar rle ri, sraric iessuie oice alieu ro rle
enriie suiace, oi uynamic iorarion aiounu rle
ving Lase. In rle mouel ving virl exonenrially
ueclining lexuial sriness, Lorl sraric anu uy-
namic Lenuing aie concenriareu neai rle ri anu
riailing euge o rle ving, vleieas Lenuing in rle
simlei mouel occuis along rle enriie san (Iig 4,
oi movies o uynamic Lenuing, see lrr://aculry.
vaslingron.euu/uanielr/movies).
Fig. 4. Finite-element models
(FEMs) based on Manduca sexta
wings. A. FEMs 1 and 2 with colors
representing different values of
Young's modulus. Model 1 has fxed
values of Young's modulus for vein
and membrane elements; whereas
Model 2 has values of Young's
modulus that decline towards the
tip and trailing edge. B. Results
of static loading with a point force
(green arrow). Displacement from
original position (black outline)
is indicated by the color bar. C.
Results of static loading with a
pressure force applied to the lower
surface (green arrows), equivalent
to the approximate pressure on
a wing during steady fight (from
Combes and Daniel 2003b).
1lese iesulrs suggesr rlar rle slaily ueclining
lexuial sriness measuieu in ieal vings lels main-
rain iigiuiry neai rle ving Lase, vlile localizing
Lenuing ro rle ri anu riailing eugeiegions o
airiculai imoirance in conriolling aeiouynamic
oice iouucrion. 1le uruie inregiarion o assive
lexiLiliry anu sarial arreins o lexuial sriness
inro mouels o insecr liglr vill Le key in uereimin-
ing rle uncrional signihcance o ving sriucruie anu
uynamic Lenuing ro insecr liglr eioimance.
References Cited
Combes, S. A., and J. L. Daniel. 2003a. Ilexuial sriness
in insecr vings. I. Scaling anu rle inluence o ving
venarion. |. Lx. Biol. 206: 29929S.
Combes, S. A., and J. L. Daniel. 2003b. Ilexuial sriness
in insecr vings. II. Sarial uisriiLurion anu uynamic
ving Lenuing. |. Lx. Biol. 206: 29S9299.
S. A. Combes is a Nillei Reseaicl Iellov in rle
Der. o Inregiarive Biology, Univ. o Calioinia,
Beikeley, CA 9420, scomLes@Leikeley.euu. Sle
is cuiienrly sruuying liglr maneuveiaLiliry anu
sraLiliry in insecr anu lummingLiius. Tom. L. DanieI
is a ioessoi anu claii o rle Der. o Biology, Univ.
o Waslingron, Box 351S00, Searrle, WA 9S195,
uanielr@u.vaslingron.euu. 7