Mülle , Schneide , Ikemeye & B ocha d Desc ip ion o las ins a la a o Co dulegas e kalkmani
108
In e na ional Jou nal o Odona ology │ Volume 26 │ pp. 108–113
In e na ional Jou nal o Odona ology
2023, Vol. 26, pp. 108–113
doi:10.48156/1388.2023.1917208
Resea ch A icle
OPEN ACCESS
This a icle is dis ibu ed
unde he e ms o he
C ea i e Commons
A ibu ion License,
which pe mi s un es ic ed use,
dis ibu ion, and ep oduc ion in
any medium, p o ided he o iginal
au ho and sou ce a e c edi ed.
Published: 4 Sep embe 2023
Recei ed: 11 Ap il 2023
Accep ed: 9 Augus 2023
Ci a ion:
Mülle , Schneide , Ikemeye &
B ocha d (2023):
Desc ip ion o las ins a la a o
Co dulegas e kalkmani (Odona a:
Co dulegas idae).
In e na ional Jou nal o
Odona ology, 26,
108–113
doi:10.48156/1388.2023.1917208
Da a A ailabili y S a emen :
All ele an da a a e
wi hin he pape .
Desc ip ion o las ins a la a o
Co dulegas e kalkmani
(Odona a: Co dulegas idae)
Ole Mülle a, Thomas Schneide b, Die ma Ikemeye c & Ch is ophe B ocha dd
a Independen Resea che , Bi kenweg 6d, 15306 Libbenichen, Ge many;
Email: muelle [email p o ec ed]om
b Independen Resea che , A nold-Knoblauch-Ring 76, 14109 Be lin-Wannsee, Ge many;
Email: homas. [email protected]
c Independen Resea che , Bille becke S . 6, 48329 Ha ixbeck, Ge many;
Email: DKJIkemey[email p o ec ed]
d Independen Resea che , Lening adweg 14, 9752 TP G oningen, The Ne he lands;
Email: in o@cb ocha d.com
Co esponding au ho : Ch is ophe B ocha d
Abs ac . The inal ins a la a o Co dulegas e kalkmani Schneide , Vie s ae e, Mülle ,
an Pel , Caspe s, Ikemeye , Snego aya & Dumon 2021 is desc ibed and illus a ed based
on one emale exu ia collec ed in Alakabük, Bi lis p o ince, Tu key, in July 2022. The exu-
ia o Co dulegas e kalkmani is compa ed wi h he exu iae o membe s o he “bol onii
g oup” in he eas e n pa o he Wes e n Palaea c ic: C. anb inkae, C. pic a, and C. he os.
The exu ia o C. kalkmani shows no clea dis inc ion in compa ison o he o he membe s
o he eas e n “bol onii g oup”.
Key wo ds. Exu iae, habi a , Tu key
In oduc ion
Co dulegas e kalkmani was ecen ly desc ibed as a Wes e n Palaea c ic Co dule-
gas e species (Schneide e al., 2021). The main mo phological cha ac e s we e
con i med by 16 addi ional males ecen ly collec ed in Eas e n Tu key (Schneide
e al., 2022). The special ea u e o he in e io appendage, which is b oade han
long so ha he dis al poin ed lobes a e isible om abo e, is unique in he genus.
This makes i easy o iden i y male imagines o his species. C. kalkmani belongs
o he ”bol onii g oup” o Co dulegas e species o he Wes e n Palaea c ic. This
”bol onii g oup” o he Wes e n Palaea c ic includes h ee wes e n dis ibu ed spe-
cies: C. bol onii, C. inac iae, and C. p inceps, whe eas C. he os, C. pic a, C. kalk-
mani, and C. anb inkae a e dis ibu ed in he eas . The exu iae o he ”bol onii
g oup” a e all e y simila (Holuša, 2022; Seidenbusch e al., 2015; Ve schu en,
1989). The la al s ages o C. kalkmani a e no known so a . In his pape he
exu ia o C. kalkmani based on a specimen collec ed in Eas e n Tu key in Bi lis
p o ince is desc ibed. In his egion no o he membe o he ”bol onii g oup” oc-
cu s (Schneide e al., 2022). We compa e i wi h he cha ac e is ics o he exu iae
o o he ela ed eas e n species o he g oup om di e en geog aphical egions
(C. he os, C. pic a, and C. anb inkae).
Me hods
Th oughou he pape he e m ‘exu iae’ is used o e e o he inal ins a exu iae
and no o any ea lie ins a .
Mülle , Schneide , Ikemeye & B ocha d Desc ip ion o las ins a la a o Co dulegas e kalkmani
109
In e na ional Jou nal o Odona ology │ Volume 26 │ pp. 108–113
Examined ma e ial
One exu ia was ound du ing a ield ip in Eas e n Tu -
key which ook place om 14. ii. o 26. ii.2022 (Schnei-
de e al., 2022). The exu ia was ound on a b ook
(38°21’47.9’’ N, 42°45’36.7’’ E, 1,988 m a.s.l.) on July
25 h, 1 km SE o Alakabük, Bi lis p o ince.
We did no obse e any ha ching imagines. We as-
signed he exu ia o C. kalkmani based on he ac ha
he only adul s o Co dulegas e lying along he b ook
belonged o his species. Se e al pa olling males o
C. kalkmani we e caugh and de e mined. One emale
was obse ed du ing o iposi ion unde he shel e o a
bush o Salix spec.
Sampling
The exu ia was collec ed on a oo o a willow (Salix
spec.) abou 40 cm abo e he wa e line o a li le pool
whe e he cu en eloci y was lowe . See below o
mo e in o ma ion on he habi a .
Measu emen s
Fo he desc ip ion and he compa ison o he spe-
cies di e en measu emen s we e ca ied ou . As all
measu emen s we e ca ied ou on one single exu ia
o a emale la a, we used measu emen s o sec ions
o in ac body pa s and hose ha we e only sligh -
ly de o med. These included he wid h and leng h o
he labium and he abdomen. The absolu e leng h o
he body and he wid h o he head, howe e , can-
no be de e mined accu a ely om his exu ia. Mea-
su emen s we e done wi h a sliding gauge (1/100 mm
accu acy). Mic oscopic pho og aphs we e aken o
measu emen s o he labium by using a CMEX 5000
(Eu omex) came a. The measu emen dis ances in he
pho os we e measu ed wi h ImageFo o 3 (1/100 mm
accu acy).
Desc ip ion o he la a
Gene al. The la a o C. kalkmani has he ypical ap-
pea ance o co dulegas id la ae (Fig. 1). The o al
leng h o he emale exu ia is abou 47 mm, he colo
o he body is b own, wi h no dis inc colo pa e ns,
he en i e body is co e ed wi h ine hai s, be ween he
hai s, sedimen pa icles om he habi a adhe e o he
body. Longe hai s and se ae a e p esen a he pos e-
io edge o ho ax, abdominal e gi es, an e io bo de
o s e ni es and legs.
Head. Nea ly ec angula in do sal iew (Fig. 2A), maxi-
mum wid h abou 8 mm, clypeus b own in colo , an e i-
o ma gin o ons wi h a ow o sho hai s, compound
eyes la ge, o ming an e o-la e al angles, in e io pa s
o compound eyes da k b own, segmen s 5–7 o bo h
an ennae a e missing in he specimen, leng h o an en-
nae segmen s (AS1 = 0.31; AS2 = 0.69; AS3 = 0.33; AS4 =
0.38 mm).
The labium has he gene al shape o a co dulegas-
id (Figs 2B, C, E). P emen um elonga ed. Leng h :
wid h a io o p emen um 6.23 : 5.93 = 1.05. An e io
ma gin o ligula wi h a ow o dense and e y sho se-
ae, in medial iew wi h wo humped p o ube ances
sligh ly di e en in shape (Fig. 2D). The la e al ma gins
o p emen um wi h a ow o sho , egula ly a anged
se ae. The mesial ma gins o palpal lobes wi h p omi-
nen i egula ly poin ed ee h (Figs 2B, C, E). Tips o all
ee h s ongly scle o ized, indica ed by a s ong b own
colo . Mo eable hooks long and s ou wi h da k b own
apex. P emen um in do sal iew wi h i e long se ae
a he igh side, and ou sho se ae medially placed,
pa s o he le side a e missing (Fig. 2E, g ay-shaded
a ea). Palpal lobes wi h i e palpal se ae, ou e ma -
gins o palpal lobes wi h wo ows o dense se ae (de-
ails Fig. 2E).
Tho ax. Wing cases nea ly pa allel, o ewings 10.3 mm,
hindwings 9.3 mm, spa sely se osed. The an e io ma -
gin o o ewings ex ending up o he pos e io ma gin
o S3, ma gin o hindwings almos ex ends o he an-
e io ma gin o S4 (Fig. 1). All legs wi h ows o se ae
in e up ed by ba e a eas. Hindlegs ex ending o 7 h ab-
dominal segmen , all a si wi h long cu ed claws, a sal
o mula: 3–3–3 (Figs 1, 3B).
Abdomen. The abdomen is enla ged in he middle
(Figs 3A, D), maximum wid h on S5; all e gi es wi h
e y sho se ae beside some single longe se ae, la -
e al ma gins o abdominal segmen s wi h ows o some
longe se ae, no do sal spines a all (Fig. 3F), pos e o-
la e al spines a S9 (Figs 3D, E), only a hump-shaped
la e al p o ube ance on S8, hidden be ween se ae o
segmen ’s ma gin.
The caudal appendages o m a cone shaped py a-
mid in do sal iew (Figs 3A, C). Epip oc (2.49 mm)
app oxima ely as long as pa ap oc s (2.57 mm), ce ci
(0.89 mm) a e less han hal as long as epi- and pa a-
p oc s. In la e al iew, he pa ap oc s cu e sligh ly en-
ally (Fig. 3F).
The o iposi o a S9 cu ed in a c escen shape does
no each S10, maximum leng h 2.36 mm (Figs 3D, F).
Habi a
In Bi lis p o ince C. kalkmani was ound a h ee locali-
ies (Schneide e al., 2022). In he moun ainous egion
sou h o he highway D 300 (Van Bi lis Yolu) be ween
he Kuskunki an unnel a he Bi lis-Van bo de and
wes wa d o Gün epe he e a e a numbe o b ooks.
These b ooks eed a s eam which uns pa allel o he
highway and lows nea Gün epe in o he Kesan Ri e .
The b ook, whe e he exu ia was ound (Fig. 4), is 70–
110 cm b oad and abou 10–40 cm deep wi h a s ony
s eambed wi hou subme ged ege a ion bu pa ially
Mülle , Schneide , Ikemeye & B ocha d Desc ip ion o las ins a la a o Co dulegas e kalkmani
110
In e na ional Jou nal o Odona ology │ Volume 26 │ pp. 108–113
Figu e 1. Recons uc ion o he las ins a o a emale la a o
Co dulegas e kalkmani. D awing OM
Mülle , Schneide , Ikemeye & B ocha d Desc ip ion o las ins a la a o Co dulegas e kalkmani
111
In e na ional Jou nal o Odona ology │ Volume 26 │ pp. 108–113
shaded by bushes and ees (Salix spec.). I s low eloc-
i y was abou 2–3 km/h. Sec ions wi h ipples al e na -
ed wi h low- low calm wa e wi h sandy sedimen s o
de i us. Pools caused by la ge s ones c ea ed a mosaic
wi h a iable low condi ions and subs a e pa ches.
The habi a is shown in Figu e 4.
Table 1. Compa ison o measu emen s o di e en Co dulegas e species (*Holuša 2022).
species Co dulegas e
kalkmani
Co dulegas e
he os*
Co dulegas e
pic a*
Co dulegas e
anb inkae*
N 1 30 30 30
Leng h o spine on segmen 8 0 0.3–0.5 0–0.25 0
Leng h o spine on segmen 9 0,23 0.4–0.6 0.15–0.35 0.05–0.3
Ra io la e al spine 9/segmen 9 0,10 0.20–0.32 0.12–0.22 0.03–0.15
Numbe o palpal se ae 5555
Numbe o long p emen al se ae 5 5 6 ( a ely 7) 5 ( a ely 6)
Numbe o sho p emen al se ae 4 2–3 3–4 1–3
Figu e 2. Head o la a o Co dulegas e kalkmani. A – head, do sal iew (do ed lines indica e missing an ennae segmen s);
B – head, on al iew; C – labium, en al iew; D – an e io ma gin o ligula in en al iew; E – labium, do sal iew (missing
s uc u es indica ed wi h g ay-shaded a ea). D awing OM
Mülle , Schneide , Ikemeye & B ocha d Desc ip ion o las ins a la a o Co dulegas e kalkmani
112
In e na ional Jou nal o Odona ology │ Volume 26 │ pp. 108–113
Figu e 3. De ails o he abdomen and he ho ax o he la a o Co dulegas e kalkmani. A – abdomen, do sal iew; B – a sus,
hindleg; C – anal py amid, do sal iew; D – abdomen, en al iew; E – de ails o la e al spines o p o ube ances, la e al iew;
F – abdomen la e al iew. D awing OM
Figu e 4. Habi a o Co dulegas e kalkmani whe e he exu ia was collec ed. Two sec ions o he same b ook a e shown,
38°21’47.9’’ N, 42°45’36.7’’ E, 1,988 m a.s.l., Eas Tu key, Bi lis p o ince.
Mülle , Schneide , Ikemeye & B ocha d Desc ip ion o las ins a la a o Co dulegas e kalkmani
113
In e na ional Jou nal o Odona ology │ Volume 26 │ pp. 108–113
Discussion and conclusions
Co dulegas e kalkmani is a ecen ly desc ibed Wes -
e n Palaea c ic Co dulegas e species (Schneide e
al., 2021, 2022). I belongs o he ”bol onii g oup” and
is he only membe o his g oup in he Wes e n A me-
nian Highlands (Schneide e al., 2022). The exu iae o
C. pic a (pic us) and C. he os we e al eady desc ibed
by Ve schu en (1989), he exu ia o C. anb inkae was
i s desc ibed by Seidenbusch e al. (2015) and hen
again oge he wi h he la ae by Holuša (2022). The
exu ia o C. kalkmani is desc ibed he e o he i s
ime and is compa ed o he o he eas e n membe s
o he Wes Palaea c ic “bol onii goup”. In he egion
whe e he exu ia was ound no o he membe o he
“bol onii g oup” occu s (Schneide e al., 2021, 2022).
The a he slende and elonga ed p emen um, and
i e o ewe p emen al se ae is in line wi h he eas -
e n membe s o he bol onii clade-(g oup) al eady
ecognized by Ve schu en (1989). Seidenbusch e al.
(2015) wo ked ou se e al di e en ial c i e ia in he
mo phology o exu iae be ween C. anb inkae and
C. pic a, o example, sho e epip oc and pa ap oc s
in compa ison wi h leng h o men um, leng h o su-
p aanalis and leng h o e gi e 10 and s e ni e 10.
Co dulegas e kalkmani clea ly i s wi hin he p opo -
ions o he o he h ee species o his g oup. All he
species show he same numbe o palpal se ae: i e.
On he p emen um C. pic a mos ly has six long p e-
men al se ae while he o he species ha e i e ( a ely
six in C. anb inkae). The numbe o sho p emen al
se ae a ies among di e en species and wi hin he
same species. Co dulegas e he os and C. anb inkae
ha e no mo e han h ee sho p emen al se ae,
C. pic a a y be ween h ee and ou and C. kalk-
mani has ou . On he abdomen, C. he os has la e al
spines on S8, C. pic a shows a ia ion in his c i e ion
whe eas C. anb inkae has no la e al spine on S8. The
exu ia o C. kalkmani doesn’ ha e la e al spines on
segmen 8. Co dulegas e he os has s onge la e al
spines on segmen 9 han C. kalkmani, C. pic a, and
C. anb inkae.
Holuša (2022) compa es he leng h o he ce ci in
ela ion o he leng h o S10. His da a sugges ha
ce ci a e longe han S10 in all compa ed species.
Howe e , examina ion o his igu e o C. anb inkae,
does no suppo his iew. Fu he mo e, ou own
da a o exu iae o C. anb inkae and he published
da a by Seidenbusch e al. (2015) do no suppo ei-
he his assump ion. Indeed, he ce ci o exu iae o
C. anb inkae a e sho e han S10.
Using ou mo phological da a plus c edible pub-
lished cha ac e s a i s a emp o a dicho omic key
ha summa izes he main di e ences be ween hese
ou species is made. As al eady s essed, wi h only
one exu ia o C. kalkmani i is no possible o con i m
in aspeci ic mo phological di e ences.
Ne e heless, a s a has been made:
1 Numbe o sho p emen al se ae ≥ 4 .................... 2
............................................ (C. pic a & C. kalkmani)
1’ Numbe o sho p emen al se ae < 4 .................... 3
.......................... (C. he os, C. anb inkae & C. pic a)
2 Numbe o long p emen al se ae 6 o 7 ....... C. pic a
2’ Numbe o long p emen al se ae 5 ....... C. kalkmani
3 Leng h o spine on segmen 9 ≥ 0.40 mm ... C. he os
3’ Leng h o spine on segmen 9 < 0.40 mm ................
......................................... (C. anb inkae & C. pic a)
4 Anal py amid sho ; leng h o e gi e 10 : leng h
o epip oc = 1 : < 2.1 ...............................................
o leng h o s e ni e 10 : leng h o pa ap oc s =
1 : < 1.4 ............................................... C. anb inkae
4’ Anal py amid long; leng h o e gi e 10 : leng h
o epip oc = 1 : > 2.1 ...............................................
o leng h o s e ni e 10 : leng h o pa ap oc s =
1 : ≥ 1.5 ......................................................... C. pic a
Acknowledgemen s
We hank Richa d Seidenbusch † o his i eless and p ecise mo -
phological wo k on Co dulegas e species. We also hank Ca in
Mülle , Da id Chelmick, Rodol o No elo-Gu ié ez, and F ank Suh-
ling o hei commen s on he manusc ip .
Re e ences
Schneide , T., Vie s ae e, A., Mülle , O., an Pel , G. J., Caspe s, M.,
Ikemeye , D., Snego aya, N. & Dumon , H. J. (2021). Taxonomic
e ision o eas e n pa o Wes e n Palaea c ic Co dulegas e us-
ing molecula phylogeny and mo phology, wi h he desc ip ion
o wo new species (Odona a: Anisop e a: Co dulegas idae). Di-
e si y, 13, 667. h ps://doi.o g/10.3390/d13120667
Schneide , T., Ikemeye , D., Mülle , O. & an Pel , G. J. (2022). No es
on Co dulegas e kalkmani in Eas Tu key (Odona a: Co dulegas-
idae). Odona ologica 51(3/4), 289–300. h ps://doi:10.5281/
odon. 51i3-4.a
Seidenbusch, R., Ikemeye , D. & Schneide , T. (2015). Desc ip ion o
he las ins a exu iae o Co dulegas e anb inkae om I an and
compa ison wi h he las ins a exu iae o Co dulegas e insig-
nis nobilis (Odona a: Co dulegas idae). Sulzbach-Rosenbe ge
Libellen undb ie e, 18, 1–25. [p i a ely published]
Holuša, O. (2022). Desc ip ion o he las ins a la a o Co dule-
gas e anb inkae and eme gence place om no he n I an
(Odona a: Co dulegas idae). In e na ional Jou nal o Odona ol-
ogy, 25(2),72–79. h ps://doi:10.48156/1388.2022.1917175
Ve schu en, D. (1989). Re ision o he la ae o Wes e n-Palaea c ic
Co dulegas e Leach, 1815 (Odona a, Co dulegas idae) wi h a
key o he conside ed axa and a discussion on hei a ini y. Bul-
le in e Annales de la Socié é Royale Belge d‘En omologie, 125,
5–35.
