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h ps://doi.o g/10.1007/s11756-021-00997-2
ORIGINAL ARTICLE
New eco ds o Xenanoe us species (As igma a: His ios oma idae)
inWes e n Eu ope, and hei associa ion wi hpig ca casses
MedjedlineHani1 · Ma aI.Saloña‑Bo das2 · M.Alejand aPe o i1
Recei ed: 11 Sep embe 2021 / Accep ed: 15 Decembe 2021
© The Au ho (s) 2022
Abs ac
The genus Xenanoe us Mahunka, 1969, only desc ibed by he hypopial s age, is cha ac e ised by an enla ged gna hosoma, he
p esence o double claws in legs I, II and III in addi ion o ema kably b oadened emu and genu o legs I and II, pa icula ly
isible in la e al iew. Only i e species om his genus ha e been desc ibed and hey a e all om he no he n hemisphe e.
Hypopi a e gene ally ound as pho e ic on Sphae oce idae lies and Hyd ophilidae bee les. Un il p esen , hypopi o he spe-
cies Xenanoe us g andiceps ha e only been epo ed in Mongolia, desc ibed by Mahunka in 1973. Xenanoe us g andiceps is
cha ac e ised by he p esence o a pai o a ches poin ing an e io ly in he pos e io pa o he gna hosoma. He e we epo
o he i s ime X. g andiceps in Spain, and o he second ime X. es igialis. Bo h species we e ound in associa ion wi h
ca cass emains, as pho e ic on lesse dung lies (Sphae oce idae), which we e collec ed in a mixed deciduous o es nea he
A lan ic coas . Addi ional new in o ma ion on mo phological cha ac e is ics o hese Xenanoe us species, as well as habi a
equi emen s and biology a e p esen ed in his epo .
Keywo ds Myianoe inae· Ca ion· Pho esy· Re ision· Xenanoe us es igialis· Xenanoe us g andiceps
In oduc ion
Xenanoe us is a genus o Myanoe inae (His ios oma idae,
Sa cop i o mes), a sub amily o as igma id mi es, s ic ly
associa ed wi h dung lies and bee les. Unde s ess ul con-
di ions, His ios oma idae mi es moul in o a modi ied deu-
onymph known as hypopus, an adap a ion o pho esy on
lying insec s o secu ing anspo a ion o mo e a ou able
en i onmen al condi ions (Hughes 1976; B aig and Pe o i
2009). Hypopi p esen s uc u es adjus ed o pho esy such as
en al sucke s ha a e modi ied o enable hei a achmen
o hei ca ie s, equen ly insec s (Hughes 1976; B aig and
Pe o i 2009). Once de ached om he ca ie , hey moul
in o he nex ins a and con inue hei li e cycle unde op i-
mal en i onmen al condi ions. The lack o knowledge on
adul o ms o many membe s o he His ios oma idae has
p omp ed he use o he pho e ic ins a o species iden i ica-
ion (Hughes 1976; Fain 1984; Samšiňák 1989).
This is he case o Xenanoe us hypopi, whose species
ha e only been desc ibed based on he he e omo phic deu-
onymph. Xenanoe us hypopi ha e double claws as Myone i-
nae and can easily be ecognised by hei b oaden gna ho-
some (= palposome) and hei enla ged emu and genu o
legs I and II (Mahunka 1969; Fain 1984; Samšiňák 1989).
Un il p esen , only i e species o his genus a e known and
only by he pho e ic hypopus. Samšiňák (1989) included
his species in he e iew o mi es associa ed wi h Sphae -
oce idae lies. In Eu ope, only one species om Hunga y
has been desc ibed, which is X. es igialis Mahunka, 1969.
The i e known species a e pho e ic on dung lies
(Sphae oce idae, Dip e a) and dung bee les (Hyd ophi-
lidae, Coleop e a). Sphae oce idae a e small lies consid-
e ed o be cop ophilous (Samšiňák 1989). Hyd ophilidae
a e aqua ic bee les ha ha e been epo ed o be associa ed
wi h subme ged ca casses (Hobischak 1997; Cazo la Pe e i
and Mo ales Mo eno 2019). In ecen yea s, he inc easing
numbe o esea ch p ojec s add essing mammal decomposi-
ion, using pig ca cass emains, o en epo ed he p esence
o Sphae oce idae lies and Hyd ophilidae bee les as compo-
nen s o he sca enging e es ial auna (Cas illo Mi albés
* M. Alejand a Pe o i
[email p o ec ed]
1 Ecology andE olu iona y Biology, School o Biological
Sciences, Uni e si y o Reading, Reading, UK
2 Depa men o Zoology andAnimal Cell Biology, Uni e si y
o heBasque Coun y, UPV/EHU, Bilbao, Spain
/ Published online: 7 Feb ua y 2022
Biologia (2022) 77:731–737
1 3
2002; Ga cía-Rojo 2004; Tabo e al. 2005; Ma inez e al.
2007; Ebe ha d and Ellio 2008; Ma uszewski e al. 2008;
Al-Mesbah e al. 2012; Ho ens ein e al. 2012; A naldos
e al. 2014; Diaz Ma ín and Saloña-Bo das 2015; among
o he s), and his also includes he Basque Coun y (No h o
Spain), whe e sampling ook place. Thei p esence on a ca -
cass en i onmen may be acciden al bu highligh s he lack
o knowledge on he mic o-sa cosap ophagous auna. The
pho e ic mi es anspo ed by his o e looked insec auna
a e e en less known (Pe o i and B aig 2009).
A e he desc ip ion o X. es igialis Mahunka, 1969
om Hunga y, new epo s eco ded he mi es associa ed
wi h Sphae oce idae, which we e collec ed in a wa e ap
in England (Os oja-S a zewski 1999) and in a sub-dese
en i onmen in he no h o Spain (Ped ochi Renaul 1998;
Blasco Zume a and Melic 1999; I aola 1999); un o una ely,
hese ouche specimens we e no a ailable o s udy. The
second species included in his e iew, X. g andiceps, has
no ye been eco ded since i s o iginal desc ip ion om
Mongolian s eppes (Mahunka 1973). In bo h species, new
mo phological cha ac e is ics and habi a condi ions we e
obse ed and p esen ed he e.
Ma e ial andme hods
Ten pig ca casses (Sus sc o a L., 1758) we e deposi ed on
he soil o a mixed deciduous o es wi h pine ees. The ca -
casses we e p o ec ed by a me allic cage and obse ed o e
a pe iod o 12weeks; i e o he ca casses we e posi ioned
in he ea ly summe o 2009, and he o he i e eplica es
we e s udied in he summe o he consecu i e yea . Me h-
odology and insec s collec ed ha e been de ailed by Diaz
Ma ín and Saloña-Bo das (2015). Collec ed insec s, o his
wo k: lesse dung lies (Dip e a, Sphae oce idae) eco ded as
“Dip” we e ca e ully inspec ed unde a s e eomic oscope o
check o he p esence o pho e ic mi es a ached o he su -
ace. Mi es we e de ached om he ca ie wi h a mic o dis-
sec ion needle, hypopi we e clea ed in lac ic acid (50% / )
and pe manen ly moun ed on glass slides in Hoye ’s medium
o u he mic oscopic analysis (K an z 1978). Mo phologi-
cal nomencla u e ollows Hughes (1976), Mahunka (1969,
1973), and Fain (1984).
Resul s
Fi e hypopi we e isola ed om lesse dung lies (Sphae -
oce idae) eco ded as Dip 20, Dip 32, and Dip 51. Sphae -
oce idae lies we e a ac ed o pig ca casses on day 3 (Dip
51) and day 13 (Dip 20, Dip 32) a e deceased; he e o e,
he ca casses we e a he esh s age o Dip 51 and ad anced
s age o decomposi ion o Dip 20 and Dip 32. Hypopi we e
compa ed wi h pa a ype HMNH- 1448 and HMNH-1459
o X. g andiceps and X. es igialis espec i ely, ype se ies;
ollowing Mahunka’s e e ence collec ion deposi ed in he
Hunga ian Na u al His o y Museum, Budapes , Hunga y. An
illus a ion o he sucke pla es in bo h species is p esen ed
in Fig.1a–d. Bo h pa a ypes and he specimens om No h
Spain show di e ences be ween each o he as well as wi h
he o iginal desc ip ions. Compa a i e da a a e summa ised
in Online Resou ce: TableS1.
Xenanoe us g andiceps is cha ac e ised by a p opodo-
soma ab up ly na owed a he basis, inc easing he slope
owa ds he on , ollowing an S-shape (Fig.2a, b), and a
pai o a ches an e io ly poin ed in he gna hosoma (Fig.2c,
d). Basis o hese a ches un pa allel and on op o hem
appea s a scle o ised iangula s uc u e p esen in speci-
mens om he Basque coun y only (Online Resou ce:
Fig.S1). The gna hosoma is longe han b oad (L:W
1.23–1.25) and be ween i s e minal hai s, eme ging cheli-
ce ae a e obse ed (Online Resou ce: Fig.S2). The su ace
o he body and he coxis e nal egion is co e ed wi h ine
do s (see Online Resou ce: Fig.S3 o X. es igialis, which
also displays his pa e n).
In con as o he o iginal desc ip ion, a la ge suc o-
ial pla e (SP) is obse ed in specimens o X. g andiceps
collec ed om he Basque coun y (Fig.1a), bea ing wo
la ge, se a ed, unc ional suc o ial discs and 6 well de el-
oped conoides, wi h he 4 medial ones being s aigh and
an e io ly di ec ed, while he 2 la e al ones a e sligh ly
cu ed (Figs.1a and 2e). In pa a ype HMNH- 1448 only 4
conoides a e p esen and imme ge om a ∪ -shaped g oo e
(Figs.1b and 2 ). Addi ional dissimila i ies we e obse ed
in he posi ioning o se ae ound in genu I. Al hough smalle ,
some specimens a e in he ange o he ype se ies, being
210–227µm in leng h and 143–146µm in wid h.
Xenanoe us es igialis om he Basque coun y is bigge
han he pa a ype HMNH-1459 (Online Resou ce: TableS1)
and has a b oade gna hosoma. In all specimens he gna-
hosoma is ully co e ed by he p opodosoma (Fig.3a-g,
b-h, c-i). La e al ma gins o he gna hosoma un sligh ly
di e gen , bo h in he pa a ypes and in he Basque speci-
mens (Fig.3d, e, ) and no pa allel, as ou lined by Samšiňák
(1989). The imminen geni al sli (Fig.1c, d) is la ge and
he egion is pi i o m wi h one pai o pla es o ming each
al e (Fig.3m); om each al e eme ge wha appea s o
be a pai o spines (Fig.3m). The suc o ial egion (pla e)
is en olded an e io ly by lanking pla es (Figs.1c, d and
3m) and la e ally, by ligamen s eme ging om o using o
he edges o he ∪ -shaped g oo e, unning adjacen ly o he
discs. The ends o bo h s uc u es appea o be using o he
pos e io egion o he coxae o he ou h pai o legs. These
obse a ions may lead o he assump ion ha he suc o ial
egion, enclosing he discs and he conoides can be con-
olled by he mo emen o legs IV. Simila cha ac e is ics,
732 Biologia (2022) 77:731–737
1 3
al hough less clea , we e obse ed in he suc o ial pla es
o X. g andiceps specimens. The suc o ial pla e (SP) he e-
o e, is delimi ed by discon inuous so o pla es; a pai o
suc o ial discs is loca ed in he an e io egion o he suc o-
ial pla e and hei bo de s a e se a ed. Mo eo e , be ween
he discs, an incipien po en ial anal ape u e is pe cei ed
(Online Resou ce: Fig.S4), and 4 conoides eme ging om
a ∪ -shaped g oo e in he Basque specimens (Figs.1c and 3k,
l) and, again 4 conoides a e seen in he pa a ypes (Figs.1d
and 3j). In he o iginal desc ip ion only 8 suc o ial disks
we e men ioned (a anged 2min + 2 la ge + 4 medium sized)
(Online Resou ce: TableS1).
Addi ional di e ences a e obse ed in he ela i e posi-
ion o he p opodosomal se ae. Sce se ae a e no aligned
and longe in Basque specimens (Fig.3h, i) in compa ison
o he pa a ype (Fig.3g). Se ae sci a e di e en in shape o
he o iginal desc ip ion and o he s udied pa a ypes om
Hunga y, ins ead o se i o m, hey appea o be smoo h
and ape ing (Fig.3h). The dis ance be ween sce and sci
se ae is la ge in he pa a ypes han in he Basque popula-
ion (Fig.3g, h, i). In addi ion, wo po es o al eoli we e
obse ed a he basis o he p opodosoma nea he do sose-
jugal g oo e in one specimen (Online Resou ce: Fig.S3).
These po es migh ha e become isible as a esul o excess
o p essu e pe o med du ing he moun ing p ocess, as hey
we e no no ed by Mahunka in he o iginal desc ip ion.
Mi es ound in he Basque coun y a i ed ea ly on he
ca ie o he decomposing ca cass a a esh s age o decom-
posi ion, on day 3 a e dea h.
Discussion
Xenanoe us species a e known o ha e es ablished a close
associa ion wi h lesse dung lies (Samšiňák 1989). Bo h
mi e species we e ound on Sphae oce idae om he esea ch
Fig. 1 Xenanoe us spp.
Compa ison o suc o ial pla es
o Xenanoe us g andiceps
Mahunka, 1973: a om he
Basque Coun y and b pa a ype;
and o Xenanoe us es igi-
alis Mahunka, 1969: c om
he Basque Coun y and d
pa a ype. DI -suc o ial discs.
CO -conoides. GO- incipien
geni al opening. AO -eme gen
anal opening. LG -ligamen s.
GRO -g oo e. FP - lanking
pla es. Scale ba 20µm
733Biologia (2022) 77:731–737
1 3
conduc ed in Aiako Ha ia (Gipuzkoa, Spain). Simila ly,
in England, he mi e has been ound on Cop oica acu an-
gula (Ze e s ed , 1847) (Dip e a, Sphae oce idae) (Os oja-
S a zewski 1999). Ne e heless, bo h species desc ibed by
Mahunka we e collec ed om dung associa ed a h opods
o he han dung lies; he bee le Sphae idium (Coleop e a,
Hyd ophilidae) and he insec Hyd oe ia meleag is Du .
cap u ed by D . P. Somogyi in Ulan-Baa o (Mongolia). Fo
his wo k, i was no possible o con i m he iden i y o his
species, H. meleag is, and we assume ha he e is a possible
misspelling o synonymy, which we could no ind. A e i-
sion o he po en ial iden i y o his elusi e hos sugges s
ha he au ho migh ha e p oposed a mo h, Hyd oecia, a
synonymous o Go yna (Noc uidae, Lepidop e a) a pes in
cul u es (c FAO). Howe e , no species o Hyd oecia is con-
nec ed o he au ho Du .
Basque specimens show di e ences wi h he o iginal
desc ip ion including he shape and loca ion o he p opo-
dosomal se ae, as well as la ge suc o ial pla es (SP) in X.
es igialis, and a a iable numbe o conoids, a scle o ized
Fig. 2 Xenanoe us g andiceps
Mahunka, 1973, compa a i e
ana omy o pa a ype 1448
and Basque specimens. a
Basque specimen do sal iew;
b pa a ype en al iew; c
Basque specimen gna hosoma;
d pa a ype 1448 gna ho-
soma; e Basque specimen
suc o ial pla e and incipien
geni al opening; pa a ype
1448 suc o ial pla e and
incipien geni al opening. Scale
ba s 100µm(b), 20µm(d, )
734 Biologia (2022) 77:731–737
1 3
egion on he gna osoma in addi ion o incipien chelice ae
a e obse ed be ween he e minal se ae o he gna osoma in
X. g andiceps. In bo h species, lanking pla es and ligamen s
we e obse ed a ound he discs and used o he la e al sides
o he g oo e on one end, and o he pos e io egion o
coxae IV on he o he end; enclosing he suc o ial egion.
The lanking pla es and ligamen s migh be used o he
unc ional mo emen o discs and conoides. The mi es a e
possibly able o con ol hem by mo ing hei ou h pai o
legs. An addi ional incipien ape u e was obse ed be ween
he discs, which migh de elop la e in o he anal opening.
Due o hei minu e size as well as he lack o esea ch on
pho on s and hei ca ie s, hese mi es and hei hypopi a e
equen ly o e looked. Fo many species o mi es associa ed
wi h animal decomposi ion, only he pho e ic ins a s ha e
been desc ibed and he o he s ages emain unknown, as
is he case o Xenanoe us species (Mahunka 1969, 1973;
Samšiňák 1989). Mo eo e , habi a equi emen s a e a ely
de ailed in he desc ip ion o new species. He e we expand
he habi a cha ac e is ic o bo h mi e species, ha can be
ound ei he in a id (Mongolian and Moneg os s eppes) o in
highly humid en i onmen s (humid o es s in Hunga y and
Spain and a wa e ap in England).
Is he e some hing in common in such dispa a e en i on-
men s? Sphae oce idae lies a e conside ed cop ophilous and
can be ound in la ge numbe s du ing he p ocess o decom-
posi ion. These lies migh be associa ed wi h cop ophagous
animals ha isi he ca casses using sa cosap ohagous
a h opods as pho on s (Ma shall 2012). La ae de elop in
humid en i onmen s such as dung, ca ion and o he decay-
ing emains (Ma shall 2012). These epheme ous en i on-
men s a e common in decomposing ca casses and in ponds
whe e decaying ma e accumula es on he su ace leading
o he a ac ion o cop ophilous insec s.
Flies a e amongs he i s o a i e du ing decompo-
si ion whe eas, Coleop e a may be acciden al hos s o
Xenano eus ha uses he bee les o depa and lea e he
ca cass possibly looking o a new sou ce o ood and
es ablishing new colonies, once he cycle is comple ed.
Only i e hypopi we e collec ed om h ee ca ie s a wo
pa icula occasions du ing he decomposi ion p ocess,
a esh s age, pe haps b ough by he lies hemsel es,
Fig. 3 Xenanoe us es igialis Mahunka, 1969, compa a i e ana omy
o pa a ype 1459 and wo Basque specimens 1 and 2. a Pa a ype 1459
en al iew; b Basque specimen 1 do sal iew; c Basque specimen
2 do sal iew; d pa a ype 1459 gna hosoma; e Basque specimen 1
gna hosoma; Basque specimen 2 gna hosoma; g pa a ype 1459
p opodosoma wi h de ails o se ae sci and sce; h Basque specimen 1
p opodosoma wi h de ails o se ae sci and sce; i Basque specimen 2
p opodosoma wi h de ails o se ae sci and sce; j pa a ype 1459 suc o-
ial pla e and incipien geni al opening; k Basque specimen 1 suc o-
ial pla e and incipien geni al opening; l Basque specimen 2 suc o ial
pla e; m pa a ype 1459 suc o ial pla e and incipien geni al opening.
Scale ba s 100µm(a), 20µm(d, g, j, m)
735Biologia (2022) 77:731–737
1 3
and a ad ance decay, possibly lies wi h mi es, ca ied by
dung bee les -a case o hype -pho esy. The e o e, a mo e
in dep h esea ch is equi ed o gain knowledge on bo h
mi es and hei ca ie s, li e cycles, o be e unde s and
he habi a equi emen s o bo h he insec s and hei asso-
cia ed pho e ic mi es.
Ou indings con i m he p e e ence o bo h he lies and
hei associa ed mi es o humid en i onmen s, ich wi h
decaying ca ion. Whe eas X. es igialis was collec ed a
an ea ly esh s age o decomposi ion on D3, X. g andiceps
was no collec ed un il day 13 when he ca casses eached
he ad anced s age o decomposi ion close o skele oni-
za ion. The mi es e iewed he e, epo new addi ional
in o ma ion o he o iginal desc ip ions done by Mahunka
in 1969 and 1973. Xenano eus g andiceps emained un e-
co ded since i s i s desc ip ion in 1973 by Mahunka.
Bo h species o Xenanoe us we e ound on Sphae oce i-
dae, which con i ms he in ima e ela ionship be ween hese
mi es and hei ca ie s. All he species om his genus a e
closely associa ed wi h dung lies (Sphae oce idae, Dip-
e a). These small lies a e conside ed o be cop ophilous
(Samšiňák 1989). In ecen yea s, an inc easing numbe o
esea ch was conduc ed using ca cass emains and equen ly
epo ed he p esence o Sphae oce idae (Cas illo Mi albés
2002; Ga cía-Rojo 2004; Tabo e al. 2005; Ma inez e al.
2007; Ma uszewski e al. 2008; Ebe ha d and Ellio 2008;
Al-Mesbah e al. 2012; Ho ens ein e al. 2012; A naldos
e al. 2014; Diaz Ma ín and Saloña-Bo das 2015; among
o he s). Thei p esence du ing he decomposi ion p ocess
o ca casses may be acciden al bu due o hei small size,
hese lies can be o e looked when collec ing en omological
e idence; explaining he limi ed knowledge and in o ma ion
on sa cosap ophagous auna, and he e o e hei associa ed
mi es. Conclusions and assump ions on hei biology and
dis ibu ion should be aken cau iously.
This e iew con ibu es wi h wo new eco ds o Xena-
noe us hypopi associa ed wi h dung lies, which we e
collec ed om decomposing ca casses. The i s eco d
e e s o X. es igialis ecen ly epo ed om Moneg os,
a sub-dese en i onmen conside ed a emnan o he
Asian s eppes (Ped ochi Renaul 1998). The new obse -
a ion epo s he p esence o X. es igialis in a empe a e
and humid o es close o he coas , in he No h o Spain,
ai ly di e en o he sub-dese en i onmen , whe e he
mi e was i s collec ed. Aiako Ha ia na u al pa k ain all
eco ds exceed 2000mm pe yea (c . Basque Go e n-
men 2019), whe eas p ecipi a ion eco ds in Moneg os
oscilla es be ween 350 and 451mm (c . Cen o de Desa -
ollo Moneg os 2012) and in Mongolian s eppes be ween
100–350mm pe yea (B i annica Encyclopaedia 2021).
Supplemen a y In o ma ion The online e sion con ains supplemen-
a y ma e ial a ailable a h ps:// doi. o g/ 10. 1007/ s11756- 021- 00997-2.
Acknowledgemen s We hank D . Csaba om he Hunga ian Ins i u e
o Fo ensic Sciences and D . Esz e Lazanyi om he Hunga ian Na u-
al His o y Museum o acili a ing access and s udy o he pa a ypes
deposi ed in he Mahunka’s collec ion (HNHM), in Budapes (Hun-
ga y). We also hank D . Diaz Ma ín (Sociedad de Ciencias A anzadi),
who ca ied ou ieldwo k and collec ion o samples; unded by he
Basque Go e nmen and he Uni e si y o he Basque Coun y UPV/
EHU PhD p og ams.
Au ho s' con ibu ions D . Saloña-Bo das did he li e a u e e iew
needed o he p ope iden i ica ion o he mi es. D . Hani selec ed he
axonomic cha ac e s and ook he pic u es ha illus a e hem. All
au ho s con ibu ed in he edesc ip ion o bo h species.
Funding D . Saloña-Bo das was unded by he Spanish Go e nmen ,
P og ama de Mo ilidad de In es igado es “Sal ado de Mada iaga”,
Minis e io de Educación, Cul u a y Depo e.
O he au ho s decla e ha no suppo was ecei ed.
A ailabili y o da a and ma e ial Ma e ial is a ailable o ex e nal
s udy. All da a ga he ed is p o ided in his s udy.
Code a ailabili y Specimens a e p ese ed in slides moun ed in Hoy-
e ’s luid and s o ed wi h he codes X e2009Dip51 o Xenanoe us
es igialis, and Xg 09Dip32, Xg 10Dip20 o Xenanoe us g andiceps.
Decla a ions
E hical app o al Does no apply.
In o med consen Does no apply.
Con lic s o in e es /Compe ing in e es s Au ho s decla e no con lic
o in e es .
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737Biologia (2022) 77:731–737
