347
Aex oxicola pilolcu ensis, a new genus and species o alse
skin bee les (Coleop e a, Biphyllidae) om he Chilean
Valdi ian ain o es
F ancisco Tello1, C is obal Tello-A iagada2
1 Depa amen o de Química y Biología, Facul ad de Ciencias Na u ales, Uni e sidad de A acama, Copiapó, Chile
2 P og ama de Magís e en Ecología Aplicada, Facul ad de Ciencias, Uni e sidad Aus al de Chile, Valdi ia, Chile
Co esponding au ho : F ancisco Tello ([email p o ec ed])
Copy igh :
© F ancisco Tello & C is obal Tello-A iagada.
This is an open access a icle dis ibu ed unde
e ms o he C ea i e Commons A ibu ion
License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
Abs ac
Aex oxicola Tello & Tello-A iagada, gen. no ., wi h i s ype species Aex oxicola pilolcu-
ensis Tello & Tello-A iagada, sp. no ., a new Sou h Ame ican genus and species o he
amily Biphyllidae, a e desc ibed om he Valdi ian ain o es in Chile. The mo pholog-
ical cha ac e s a e illus a ed and compa ed wi h mo phologically simila ex an and
ex inc gene a. Finally, a key o gene a o Biphyllidae eco ded in Chile is p o ided.
Key wo ds: Cucuji o mia, Neo opical auna, new species, species desc ip ion, axonomy
In oduc ion
The amily Biphyllidae LeCon e (Insec a, Coleop e a) is a small amily o
Cle oidea (sensu Robe son e al. 2015) ha comp ises a leas 190 de-
sc ibed species, dis ibu ed in eigh gene a wo ldwide, wo o which a e
known only om ossils (Cline and McHugh 2010; Cline and Shockley 2010;
Węg zynowicz 2015; Maka o and Pe ko sky 2020; Aleksee e al. 2023).
Adul s and la al s ages a e ypically ound in lea li e , ui ing bodies o
ungi (Ascomyco a and Py enomyce es), and/o decaying ee ba k; how-
e e , he ecology o mos species emains unknown o science (Good ich
and Sp inge 1992; Cline and Shockley 2010). Addi ionally, he in a amilial
phylogene ic ela ionships emain unknown, wi hou sub amilial o ibal
designa ions p o ided ye (Cline and Shockley 2010).
The cu en ly known Chilean auna o Biphyllidae comp ises only wo gen-
e a: Diplocoelus Gué in-Méne ille, wi h he species D. oblongus (Ge main)
and D. o eola us Rei e , and Ancho ius Casey, wi h A. dollyae Tello &
Tello-A iagada (Elgue a 2000; Węg zynowicz 2015; Tello and Tello-A iaga-
da 2024). Based on ecen ly collec ed specimens, we desc ibe he ein a new
genus and species o Biphyllidae om Chile.
Academic edi o :
John M. Lea engood, J .
Recei ed:
10 June 2025
Accep ed:
8 July 2025
Published:
8 Oc obe 2025
ZooBank: h ps://zoobank.
o g/0165FF49-18C1-4261-956A-
0E5BDD8CE4F6
Ci a ion: Tello F, Tello-A iagada C
(2025) Aex oxicola pilolcu ensis, a
new genus and species o alse skin
bee les (Coleop e a, Biphyllidae)
om he Chilean Valdi ian ain o es .
ZooKeys 1254: 347–354. h ps://doi.
o g/10.3897/zookeys.1254.161754
ZooKeys 1254: 347–354 (2025)
DOI: 10.3897/zookeys.1254.161754
348
ZooKeys 1254: 347–354 (2025), DOI: 10.3897/zookeys.1254.161754
F ancisco Tello & C is obal Tello-A iagada: Aex oxicola pilolcu ensis, a new genus and species o Biphyllidae
Ma e ials and me hods
The sys ema ic amewo k adop ed ollows he classi ica ions p oposed
by Cline and Shockley (2010) and Węg zynowicz (2015). Mo phological e -
minology is based on Cline and McHugh (2010) and Law ence e al. (2011).
Specimens we e ea ed wi h po assium hyd oxide (KOH) o so en is-
sues and clean body pa s. To emo e esidual deb is, an ul asonic cleane
(BAKU BK-3550) was addi ionally employed.
Mac oscopic images we e ob ained using a FLEXACAM C3 came a moun ed on
a Leica S9D s e eomic oscope, while mic os uc u es we e imaged wi h an Olym-
pus BX50 mic oscope. Image composi ions we e made using Adobe Pho oshop
CC 2025. Mo phome ic measu emen s we e aken wi h he Leica Applica ion
Sui e X (LASX) . 5.1.0 so wa e. Body measu emen s included: o al body leng h
(measu ed om he an e io ma gin o he lab um, wi h he head in na u al lexion,
o he apex o he ely a), head leng h and wid h, p ono al leng h and wid h, and el-
y al leng h and wid h. All measu emen s we e aken a hei maximum leng h and
wid h, espec ably. Dissec ed s uc u es we e moun ed on ca ds and associa ed
wi h hei espec i e specimens, wi h app op ia e elabeling unde aken.
Resul s
Sys ema ics
Family Biphyllidae LeCon e, 1861
Aex oxicola Tello & Tello-A iagada, gen. no .
h ps://zoobank.o g/8A32F862-1A15-4063-BFC5-91A7333C440F
Figs 1A–C, 2A–H
Type species. Aex oxicola pilolcu ensis Tello & Tello-A iagada, by mono ypy
and p esen designa ion.
E ymology. Aex oxicola de i es om he scien i ic name o he oli illo, Aex-
oxicon punc a um Ruiz & Pa . (Aex oxicaceae), which is he dominan ee spe-
cies in he ype locali y o he ype species. The gende is eminine.
Diagnosis. Aex oxicola gen. no . can be clea ly dis inguished om he ex an
gene a by he ollowing combina ion o cha ac e s: (1) small body size, less
han 2.5 mm; (2) 3-segmen ed an ennal club; (3) a some es I–IV educed, wi h
a some es II and III longe han I and IV; (4) i s en i e wi h wo pai s o pos -
me acoxal lines, la e al lines wi h sligh ly sepa a ed pa allel idges, o ming
b oad pos me acoxal la e al lines; medial pos me acoxal lines di e ging pos-
e io ly om a ounded in e coxal p ocess, o ming an in e ed “U” shape; idg-
es o ming pos me acoxal la e al lines sligh ly sepa a ed; (5) p ono al la e al
ca ina se a e; (6) a single, well-de ined longi udinal ca ina p esen la e ally on
each side o p ono um; (7) pos e io p ono al angles acu e and pos e io edge
o p ono um bisinua e; (8) body pubescence uni o m, wi h la ge, yellow se ae.
Desc ip ion. Male habi us (n = 1). Measu emen s: o al body leng h = 2.27 mm;
maximum body wid h = 0.88 mm; head leng h = 0.33 mm; head wid h = 0.56 mm;
p ono um leng h = 0.53 mm; p ono um maximum wid h = 0.83 mm. Body elon-
ga e in do sal iew, mode a ely con ex in la e al iew; in egumen eddish o
349
ZooKeys 1254: 347–354 (2025), DOI: 10.3897/zookeys.1254.161754
F ancisco Tello & C is obal Tello-A iagada: Aex oxicola pilolcu ensis, a new genus and species o Biphyllidae
b own; do sum ex u ed, wi h mode a ely sized punc a ion homogeneously dis-
ibu ed in ows; each punc u e wi h a yellow, long, semie ec se a (Fig. 1A–C).
Head. P ogna hous, ans e se, inse ed in o p o ho ax up o pos e io ex en
o eyes; empo a p esen ; se ae simila o hose on p ono um and ely a. F on-
oclypeal su u e absen . Genal pocke s p esen . P esence o deep suban en-
nal g oo es be ween he eye and mou h ca i y, wi h well-de eloped ca inae on
mou h ca i y sides. Eyes. Da k, la ge, ounded, mode a ely p o uding; in e ac-
e al se ae absen ; omma idia mode a ely la ge. An ennae. Capi a e, wi h small
se ae on he in egumen and a ew la ge se ae along he ma gins o each seg-
men ; scape cylind ical, b oade han pedicel; an ennal club comp ising an en-
nome es IX–XI, p ominen ly expanded la e ally wi h ounded can hus (Fig. 2A).
An ennal segmen s IX and X unca ed a he apex; segmen XI en i ely ounded.
Figu e 1. Male ype o Aex oxicola pilolcu ensis Tello & Tello-A iagada, gen. no . sp. no . A. Do sal habi us; B. Ven al
iew; C. La e al iew.
350
ZooKeys 1254: 347–354 (2025), DOI: 10.3897/zookeys.1254.161754
F ancisco Tello & C is obal Tello-A iagada: Aex oxicola pilolcu ensis, a new genus and species o Biphyllidae
Mou hpa s. Mandibles isible do sally and en ally, biden a e a he apex,
bea ing small den icles; mola egion on he mesal edge wi h se i e ous ma -
gins; mola se ae sho (Fig. 2D). Maxilla y palpi wi h ou palpome es (Fig. 2E);
maxilla y segmen I longe han wide ; segmen s II and III o simila size, na -
owe a he base and widened dis ally; segmen IV oblong, longe han wide,
apex unca ed, uncus absen ; galea and lacinia se ose a he apex (Fig. 2E).
Labial palpi wi h h ee palpome es, bea ing small longi udinally g ouped se ae;
segmen s I and II educed; segmen III widened and unca ed; men um iden-
a e, he cen al apex smalle ; ligule bilobed and pubescen (Fig. 2F).
P o ho ax. P ono um ans e se, wide han he head; p ono al disc con ex,
wi h sligh median dep ession along pos e io ma gin; an e io angles sub ec an-
gula ; pos e io angles acu e, sligh ly ex ended beyond he la e al ma gins; basal
ma gin o he p ono um bisinua e (Figs 1A, 2A); a single, well-de ined, longi udi-
nal subla e al ca ina p esen on each side o he do sal su ace o he p ono um;
edge o la e al p ono al ca ina se a e (Fig. 2A). Hypome al a ea o he p ono um
smoo h, wi hou appa en modi ica ions (Fig. 1B). P ocoxal ca i ies ec angula
and closed. P os e nal p ocess p ominen , sligh ly widened medially.
P e o ho ax. Scu ellum. Scu ella shield ans e se, small, app oxima ely 2.5
imes wide han long, wi h ounded ma gins. Ely a. Elonga e, wi h punc u es
a anged in longi udinal, s ia-like ows, each punc u e wi h long, yellow se a;
in e s iae smoo h; hume al angles no p o uding; ely al scu ella y s iole ab-
sen (Fig. 1A, C). Meso ho ax. Meso en al pla e p ominen , no ched an e io ly,
ex ending pos e io ly o me a en i e, o ming p ocoxal es s (Fig. 1B). Meso-
coxae ounded, sligh ly sepa a ed, dis ance be ween hem abou equal o ha
be ween p ocoxae. Me a ho ax. Me a en i e ans e se. Me a en al disc i-
men p esen bu incomple e, es ic ed o he pos e io hi d. Two pai s o pos -
mesocoxal lines p esen . Medial pos mesocoxal lines s a ing a medial edge
o coxal ca i ies as con inua ion o in e coxal p ocess, di ec ed owa ds ou e
co ne s o me acoxae, disappea ing in an e io hi d o me a en i e. La e al
pos mesocoxal lines sligh ly la ge han medial pos mesocoxal lines, s a ing
om he i s mid-sec ion o he mesocoxal idge and culmina ing a he la e al
edge o he me a en i e. Me acoxae ans e se, sepa a ed by he p ocess o
he i s abdominal en i e (Fig. 1B), no ex ending la e ally o mee ely a.
Legs. Slende wi h emo a ela i ely na ow. P o-, meso- and me a emo a
subequal. Tibiae s aigh , each bea ing a leas ou small apical spines. Ta sal
o mula 5-5-5; a some es I and IV educed; a some es II and III each 1.5 imes
longe han a some es I o IV; a some e V elonga ed, almos equal in leng h
o a some es I–IV combined; a some e III wi h conspicuous la e al lobe-like
modi ica ions. Ta sal claws simple (Fig. 2G).
Abdomen. Abdomen wi h i e ee en i es; i s en i e leng h (as mea-
su ed a he sho es dis ance behind he me acoxal ca i ies) subequal o
second en i e; i s en i e wi h wo pai s o pos me acoxal lines, each
pai consis ing o sligh ly sepa a ed pa allel idges, o ming b oad pos -
me acoxal lines; pos me acoxal medial lines di e ging om he edges o a
ounded in e coxal p ocess, o ming an in e ed “U” shape; he idges ha
o m he pos me acoxal la e al lines a e dis inc ly b oadened, o ming a
b oad pos me acoxal medial lines. Bo h medial and la e al lines connec ed
an e io ly and pos e io ly o he ma gins o he en i e (Fig. 2C). Remaining
en i es wi hou lines on he disc (Fig. 1B).
351
ZooKeys 1254: 347–354 (2025), DOI: 10.3897/zookeys.1254.161754
F ancisco Tello & C is obal Tello-A iagada: Aex oxicola pilolcu ensis, a new genus and species o Biphyllidae
Geni alia. Male geni alia absen in holo ype (p esumably b oken o and los . I
may ha e been damaged du ing collec ion o subsequen handling o he speci-
mens). Female geni alia wi h he baculum elonga e and well de ined; oblique bac-
ulum conspicuous, sub iangula ; s ylus wi h small, conspicuous punc u es, apex
ounded and bea ing sho se ae. Vul al lobes elonga e and subpa allel (Fig. 2H).
Rema ks. The unique combina ion o cha ac e s suppo s he ecogni ion
o Aex oxicola gen. no . as dis inc om he p e iously desc ibed, li ing and
ex inc gene a wi hin he amily. Fi s , in Aex oxicola, he ely al punc a ion and
pubescence a e o a single, consis en ype in e ms o size and shape, wi h
no ma ked a ia ion. In con as , mos Biphyllidae exhibi non-uni o m ely al
Figu e 2. Male ype and emale pa a ype o Aex oxicola pilolcu ensis Tello & Tello-A iagada, sp. no . (A, G. Holo ype;
B–F, H. Pa a ype). A. De ails o holo ype an ennae and p ono um; B. Female me a en i e; C. Fi s emale abdominal
en i e; D. Female mandible; E. Female maxilla; F. Female labium; G. Me a ibia and me a a sus; H. Female geni alia. LSC.
Longi udinal subla e al ca inae. PLLm. Pos mesocoxal la e al lines o he me a en i e. PMLm. Pos mesocoxal medial
lines o he me a en i e. PLL . Pos me acoxal la e al lines o he i s abdominal en i e. PML . Pos me acoxal medial
lines o he i s abdominal en i e.
352
ZooKeys 1254: 347–354 (2025), DOI: 10.3897/zookeys.1254.161754
F ancisco Tello & C is obal Tello-A iagada: Aex oxicola pilolcu ensis, a new genus and species o Biphyllidae
pubescence, o en wi h a mix u e o long and sho se ae, and punc u es o
a ying sizes o i egula dis ibu ion (e.g., Al haesia Pascoe, some Biphyllus De-
jean, Eude opus Sha p, Diplocoelus) (Maka o and Pe ko sky 2020). This ai
is sha ed wi h he ex inc genus Paleobiphyllus Maka o & Pe kos ky; howe e ,
in Aex oxicola he in egumen is smoo h. Secondly, he combina ion o only
one pai o longi udinal subla e al ca inae on he p ono um and wo pai s o
pos me acoxal lines on he i s abdominal en i e a e unique; mos ep esen-
a i es o Biphyllidae ha e 2–5 pai s o addi ional ca inae on he p ono al disc,
and one o wo pai s o pos me acoxal lines. The widening o he la e al pos -
me acoxal lines is no ewo hy and ep esen s ano he dis inc i e cha ac e o
his new genus. Thi d, he p ono al la e al ca inae a e dis inc ly se a e, while in
mos Biphyllidae he la e al ca inae a e c enula e o smoo h. The se a e ma -
gins a e obse ed in some species o Biphyllus (e.g., B. ma mo a us (Rei e );
see Pa k e al. 2012), howe e , his genus has a 2-segmen ed an ennal club.
Aex oxicola pilolcu ensis Tello & Tello-A iagada, sp. no .
h ps://zoobank.o g/EA34EA9B-288A-4C96-B7E5-D2AC82ED0337
Figs 1A–C, 2A–H
Type ma e ial. Holo ype • male, Chile, Los Ríos egion, Valdi ia p o ince,
Pilolcu a–Pun a Chungungo loca ion, 15.III.2024, leg. F. Tello. Coo dina es:
39°39'35.7"S, 73°21'40.1"W. Pi all ap; deposi ed a collec ion o F ancisco
Tello a ailable in Uni e sidad de A acama, Copiapó, Chile. Pa a ype ♀ (n = 1),
same da a as holo ype. Dissec ed o mo phological analysis.
E ymology. The speci ic epi he is an adjec i e de i ed om Pilolcu a, he
name o he ype locali y.
Desc ip ion. See genus desc ip ion.
Dis ibu ion. Known only om Pun a Chungungo, Pilolcu a, Los Ríos
egion, Chile.
Key o gene a o Biphyllidae known om Chile
1 P ono al disc do sally wi h 5 pai s o clea ly isible ca inae ........................
................................................................................... Ancho ius Casey, 1900
– P ono al disc do sally wi h wo o ewe pai s la e al ca inae ...................2
2 Ely al pubescence, o en wi h a mix u e o long o sho , yellow o b own
se ae; la e al p ono al ca ina smoo h o sligh ly c enula e; in e coxal p o-
cess o i s abdominal en i e dis inc ly acu e ...........................................
............................................................. Diplocoelus Gué in-Méne ille, 1843
– Ely al pubescence uni o m, wi h la ge, yellow se ae; la e al p ono al ca-
ina se a e; in e coxal p ocess o he i s abdominal en i e dis inc ly
ounded ...............................Aex oxicola Tello & Tello-A iagada, gen. no .
Discussion
The disco e y o Aex oxicola pilolcu ensis gen. no . sp. no . ep esen s a signi -
ican addi ion o he poo ly known Chilean auna o Biphyllidae and highligh s
he unde es ima ed di e si y o his g oup in sou he n Sou h Ame ica. The
353
ZooKeys 1254: 347–354 (2025), DOI: 10.3897/zookeys.1254.161754
F ancisco Tello & C is obal Tello-A iagada: Aex oxicola pilolcu ensis, a new genus and species o Biphyllidae
auna o Chile emains poo ly known om bo h axonomic and phylogene ic
pe spec i es. Indeed, he ype se ies o Diplocoelus o eola us is cu en ly miss-
ing (Tello, pe s. obs.); he e o e, i is alidi y is unce ain. Likewise, bo h D. o e-
ola us and D. oblongus equi e a ho ough e ision o assess he ex en o i s a -
ini y wi h he genus Diplocoelus. The e o e, u u e s udies combining molecula
da a, la al mo phology, and b oade geog aphic sampling will be essen ial o
cla i ying phylogene ic ela ionships wi hin Biphyllidae, including he placemen
o Aex oxicola and i s po en ial a ini y o o he Sou he n Hemisphe e lineages.
Acknowledgemen s
We hank ou anonymous e iewe s o hei ou s anding con ibu ion o he
imp o emen o his manusc ip .
Addi ional in o ma ion
Con lic o in e es
The au ho s ha e decla ed ha no compe ing in e es s exis .
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
No use o AI was epo ed.
Funding
G an ATA 22991, Minis e io de Educación.
Au ho con ibu ions
Concep ualiza ion: CSTA, FT. Da a cu a ion: FT. Fo mal analysis: FT. Funding acquisi ion:
FT. Me hodology: FT, CSTA. Visualiza ion: CSTA. W i ing - o iginal d a : FT. W i ing - e-
iew and edi ing: FT, CSTA.
Au ho ORCIDs
F ancisco Tello h ps://o cid.o g/0000-0003-0862-3475
C is obal Tello-A iagada h ps://o cid.o g/0000-0003-3920-9445
Da a a ailabili y
All o he da a ha suppo he indings o his s udy a e a ailable in he main ex .
Re e ences
Aleksee V, Bukejs A, Vi ali F (2023) New da a on Biphyllidae (Coleop e a: Cucuji o mia:
Cle oidea) om Bal ic ambe : mo e di e se assemblage han p e iously assumed
and i s clima ic in e p e a ion. His o ical Biology 36: 2520–2529. h ps://doi.o g/10.
1080/08912963.2023.2264858
Cline AR, McHugh JV (2010) New gene ic synonymy in Biphyllidae (Coleop e a: Cucu-
joidea), wi h a checklis o Ancho ius Casey, 1900. Coleop e is s Bulle in 64: 98–99.
h ps://doi.o g/10.1649/0010-065X-64.1.98
354
ZooKeys 1254: 347–354 (2025), DOI: 10.3897/zookeys.1254.161754
F ancisco Tello & C is obal Tello-A iagada: Aex oxicola pilolcu ensis, a new genus and species o Biphyllidae
Cline AR, Shockley FW (2010) 10.6. Biphyllidae LeCon e, 1861. In: Küken hal W, Leschen
R, Beu el R, Law ence J (Eds), Coleop e a, Bee les, Volume 2, Mo phology and Sys-
ema ics (Ela e oidea, Bos ichi o mia, Cucuji o mia pa im). De G uy e , Be lin,
306–311. h ps://doi.o g/10.1515/9783110911213.306
Elgue a M (2000) Es ado ac ual del conocimien o de los coleóp e os de Chile (Insec a:
Coleop e a). P IBES-2000. P oyec o pa a Ibe oamé ica de En omología Sis emá ica
17: 145–154.
Good ich MA, Sp inge CA (1992) A e ision o he amily Biphyllidae (Coleop e a) o
Ame ica, no h o Mexico. Coleop e is s Bulle in 46: 361–377.
Law ence JF, Ślipiński A, Seago AE, Thaye MK, New on AF, Ma aldi AE (2011) Phy-
logeny o he Coleop e a based on mo phological cha ac e s o adul s and la ae.
Annales Zoologici 61: 1–217. h ps://doi.o g/10.3161/000345411X576725
Maka o KV, Pe ko sky EE (2020) Smalles and oldes alse skin bee le: Paleobiphyllus
ponoma enkoi gen. e sp. no . (Coleop e a: Cle oidea: Biphyllidae) om San onian
Taimy ambe , no he n Russia. C e aceous Resea ch 106: 1–8. h ps://doi.
o g/10.1016/j.c e es.2019.104238
Pa k SJ, Ko ES, Ahn KJ (2012) Fi s eco d o he bee le amily Biphyllidae (Coleop e a)
in Ko ea. En omological Resea ch 42: 185–191. h ps://doi.o g/10.1111/j.1748-
5967.2012.00452.x
Robe son JA, Ślipiński A, Moul on M, Shockley FW, Gio gi A, Lo d NP, McKenna DD,
Tomaszewska W, Fo es e J, Mille KB, Whi ing MF, McHugh JV (2015) Phylogeny
and classi ica ion o Cucujoidea and he ecogni ion o a new supe amily Coccinel-
loidea (Coleop e a: Cucuji o mia). Sys ema ic En omology 40: 745–778. h ps://doi.
o g/10.1111/syen.12138
Tello F, Tello-A iagada C (2024) Fi s epo o he genus Ancho ius Casey, 1900
(Coleop e a: Biphyllidae) in Chile, wi h desc ip ion o a new species. Re is a Chilena
de En omologia 50: 625–631. h ps://doi.o g/10.35249/ che.50.4.24.04
Węg zynowicz P (2015) Ca alogue o Biphyllidae (Coleop e a) o he wo ld. Annales
Zoologici 65: 409–471. h ps://doi.o g/10.3161/00034541anz2015.65.3.002
