233
Mo phological and molecula e ision o he sub amily
He e olepisma inae (Zygen oma, Lepisma idae), wi h desc ip ions
o wo new gene a om he A acama Dese , Chile
Ra ael Mole o-Bal anás1* , Ál a o Zúñiga-Reinoso2* , Miquel Gaju-Rica 1, Reinha d P edel2
1 Depa men o Zoology, Uni e si y o Có doba, C-1 Campus de Rabanales, 14071 Có doba, Spain
2 Ins i u e o Zoology, Uni e si y o Cologne, Zülpiche S . 47b, 50674 Köln, Ge many
Co esponding au ho s: Ra ael Mole o-Bal anás ([email p o ec ed]); Ál a o Zúñiga-Reinoso ([email p o ec ed])
Copy igh : © Ra ael Mole o-Bal anás e al.
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
This wo k aims o upda e he sys ema ics o sil e ish o he sub amily He e olepisma i-
nae (Zygen oma, Lepisma idae) based on mo phological and molecula da a o new
axa om he A acama Dese (Chile) wi h a compa ison wi h specimens om A gen-
ina, om whe e species o iginally desc ibed as belonging o he genus He e olepisma
Esche ich, 1905 come. This compa ison also includes COI gene sequences o A gen ine
specimens a ibu ed o H. andinum (Sil es i, 1902), and o Aus alian He e olepisma i-
nae p e iously classi ied as He e olepisma o Visma Smi h e al., 2021. As a esul , he
gene a Lapidisma gen. no . and Cac isma gen. no . a e desc ibed om Chile, and new
mo phological cha ac e s diagnos ic o H. andinum a e p o ided. The species Lapidisma
paposanum sp. no . and Cac isma camanchaca sp. no . a e associa ed wi h di e en
habi a s. The species He e olepisma annec ens (Sil es i, 1924) is en a i ely a ibu ed
o he i s genus (Lapidisma annec ens comb. no .). The molecula da a place he h ee
Sou h Ame ican gene a in sepa a e lineages, bu clus e hem as a sis e g oup o he
Aus alian He e olepisma + Visma. This suppo s he p oposal o he genus Sil es isma
gen. no . o include Aus alian species p e iously assigned o He e olepisma. Addi ion-
ally, he new name Vis olepisma nom. no . is p oposed o sil e ish species p e iously
assigned o he genus Visma, since his gene ic name is p e-occupied. Wi h his e-a -
angemen , he sub amily He e olepisma inae consis s o six gene a, pending u he
s udies in o he geog aphic egions whe e poo ly s udied species occu ha we e (and
a e p o isionally) assigned o He e olepisma.
Key wo ds: Cac isma, He e olepisma, Lapidisma, sil e ish, Sou h Ame ica, Visma, Vis-
olepisma
In oduc ion
He e olepisma inae Mendes, 1991 is a sub amily o sil e ish (O de Zygen-
oma, amily Lepisma idae) widesp ead in he Sou he n Hemisphe e, p oba-
bly wi h a Gondwanan o igin (Mole o-Bal anás e al. 2024). I was de ined by
Mendes (1991) and he mos ema kable cha ac e s ha he p o ided o he
diagnosis (no necessa ily synapomo phic cha ac e s) o his sub amily a e he
Academic edi o : Pa el S oe
Recei ed:
4 Ma ch 2025
Accep ed:
28 Augus 2025
Published:
20 No embe 2025
ZooBank: h ps://zoobank.
o g/3A8CC4DB-048F-4A43-9568-
FE283DAB257E
Ci a ion: Mole o-Bal anás R,
Zúñiga-Reinoso Á, Gaju-Rica M,
P edel R (2025) Mo phological and
molecula e ision o he sub amily
He e olepisma inae (Zygen oma,
Lepisma idae), wi h desc ip ions o
wo new gene a om he A acama
Dese , Chile. ZooKeys 1260:
233–278. h ps://doi.o g/10.3897/
zookeys.1260.151902
ZooKeys 1260: 233–278 (2025)
DOI: 10.3897/zookeys.1260.151902
* These au ho s con ibu ed equally o his wo k.
234
ZooKeys 1260: 233–278 (2025), DOI: 10.3897/zookeys.1260.151902
Ra ael Mole o-Bal anás e al.: Re ision o he sub amily He e olepisma inae (Zygen oma)
smoo h mac ochae ae, apically bi id o i id; apical a icle o maxilla y palp
wi h ew cylind ical sensilla ( he p onged apical sensilla o Mole o-Bal anás
e al. 2024), in some species wi h sexual dimo phism; apical a icle o labial
palp wi h 3+2 papillae a anged in wo ows; an e io ma gin o p ono um wi h
se al colla ; ho acic s e ni es well de eloped; pa ame es esiculi o m, lacking
glandula a ea. Mo e ecen ly, Mendes (2018) conside ed a g ea e a iabili y o
he pa ame es o his sub amily (also subcylind ical and pseudo-a icula ed).
All species belonging o He e olepisma inae had been included in he genus
He e olepisma Esche ich, 1905, un il Smi h and Mi chell (2019) and Smi h e
al. (2021) dis inguished he gene a Ma i isma Smi h & Mi chell, 2019 and Vis-
ma Smi h, Mi chell & Mole o-Bal anás, 2021, espec i ely; bo h wi h ep esen-
a i es only in Aus alia and neighbou ing Paci ic islands, wi h he excep ion o
Ma i isma dispa (Yosii, 1939), desc ibed om Japan as He e olepisma dispa .
The gene ic name Visma is p e-occupied o a gas opod belonging o he ami-
ly Py amidellidae and he sil e ish o me ly known by his name ecei es a new
name he e. The e o e, Visma e e ing o Lepisma idae is only used be o e he
Resul s sec ion. Excluding he species o he abo e-men ioned wo new gen-
e a, he genus He e olepisma cu en ly consis s o 27 species (see i s column
in Table 1).
The ype species o He e olepisma is H. pampeanum (Sil es i, 1902), de-
sc ibed om se e al locali ies in A gen ina as Lepisma pampeana Sil es i,
1902 and ans e ed o he genus He e olepisma when i was e ec ed (Esch-
e ich 1905). Ano he A gen ine species belonging o his genus is H. andinum
(Sil es i, 1902). F om con inen al Chile he e a e no eco ds o He e olepisma
species, bu one species, H. annec ens (Sil es i, 1924), was desc ibed om
he a chipelago o Juan Fe nández (Sil es i 1924). Some au ho s ans e ed
all He e olepisma o Isolepisma Esche ich, 1905, a genus c ea ed o I. ise o-
sum Esche ich, 1905, bu Sil es i conside ed bo h gene a as synonyms and,
consequen ly, hey ha e been ea ed by mos au ho s du ing he las 60 yea s
as He e olepisma. Acco ding o Mendes (2011), I. ise osum should be ea ed
as species inqui enda.
A comp ehensi e su ey o insec s ac oss he A acama Dese as pa o
he p ojec “Ea h and E olu ion a he d y limi ” (h ps://s b1211.uni-koeln.de/)
e ealed se e al undesc ibed axa o Zygen oma ( o Maind onia Bou ie , 1897,
see Zúñiga-Reinoso and P edel 2019). Based on molecula and mo phological
da a, we he e desc ibe wo new gene a o he poo ly s udied sub amily He e o-
lepisma inae and discuss he ela ionships o he gene a cu en ly included in
his sub amily.
Ma e ials and me hods
Sampling o he ype se ies and gene ically simila specimens om neighbou ing
a eas ook place be ween Augus – No embe 2023 along he coas al sec ion o
he A acama Dese in An o agas a P o ince (Chile) a ound Paposo and Cues a
el Cob e (sampling pe mi s CONAF N° 016/2021, 000319°N). Addi ionally, spec-
imens assigned o He e olepisma we e collec ed nea Sanogas a, La Rioja (A -
gen ina; 29°21.03'S, 67°46.93'W, sampling pe mi o he Sec e a ía de Ambien e:
244/22) and Talcahuano in Sou he n Chile (36°42.52'S, 73°7.05'W). Specimens
we e collec ed using an aspi a o and di ec ly ans e ed in o 96% e hanol.
235
ZooKeys 1260: 233–278 (2025), DOI: 10.3897/zookeys.1260.151902
Ra ael Mole o-Bal anás e al.: Re ision o he sub amily He e olepisma inae (Zygen oma)
DNA ex ac ion, ampli ica ion, and sequencing
F om he ype specimens, DNA was ex ac ed om legs o emaining issues wi h-
ou gu a e he dissec ion and p epa a ion o he exoskele on o mo phological
analyses (ma ked wi h “*” in he examined ma e ial sec ion). Addi ionally, DNA
Table 1. Lis o sil e ish species p e iously asc ibed o he genus He e olepisma. Species ha we e al eady ans e ed
o he gene a Ma i isma and Vis olepisma ( o me ly Visma) p io o ou s udy a e no conside ed he e. He e olepisma
ise osum is no included as i is conside ed a species inqui enda (see Discussion); species highligh ed in bold equi e
e ision o con i m gene ic assignmen ; * indica es insu icien o iginal desc ip ion wi hou upda ed edesc ip ion.
Species, au ho (s), and yea o desc ip ion Dis ibu ion Rema ks (o iginal names, synonyms, p oposed
gene ic s a us, e c.)
H. andinum (Sil es i, 1902)* A gen ina Desc ibed as Lepisma andina
H. annec ens (Sil es i, 1924)* J. Fe nández Islands P o isionally ans e ed in his wo k o Lapidisma
annec ens comb. no .
H. bise osum (Ca pen e , 1916)* Seychelles, Somalia Redesc ip ion by Mendes (1988) did no p o ide
enough in o ma ion o upda ing gene ic s a us
H. bun ono um Smi h, 2016 Tasmania (Aus alia) Sil es isma bun ono um comb. no .
H. cooloola Smi h, Mi chell, Lee & Espinasa, 2019 Aus alia (QLD) Sil es isma cooloola comb. no .
H. coo ongooba Smi h, Mi chell, Lee & Espinasa, 2019 Aus alia (NSW) Sil es isma coo ongooba comb. no .
H. exac um (Sil es i, 1918)* Zanzíba (Tanzania) Desc ibed as Isolepisma exac a
H. he aldense Smi h & Mi chell, 2019 Aus alia (Co al Sea T.) Sil es isma he aldense comb. no .
H. highlandi Smi h, 2014 Aus alia (NSW) Sil es isma highlandi comb. no .
H. ho ni S ach, 1933* Ecuado (Ca acas Bay)
Ca ibbean
S ach (1933) p o ided an iden i ica ion key o he
species known a ha ime
H. howense Wome sley, 1942 Lo d Howe Island
(Aus alia)
Sil es isma howense comb. no ., Redesc ip ion by
Smi h and Mi chell (2019)
H. insula e (Banks, 1901)* Galapagos Isl.
(Ecuado )
Desc ibed as Lepisma insula is
= H. in e medium Folsom, 1924
H. japonicum (Uchida, 1968) Japan Desc ibed as Isolepisma japonica
H. k aepelini Sil es i, 1908* Aus alia (SA, WA) (Sil es isma k aepelini comb. no .)
H. michaelseni Sil es i, 1908* Aus alia (SA, WA) (Sil es isma michaelseni comb. no .)
H. milledgei Smi h & Mi chell, 2019 Lo d Howe Island
(Aus alia)
(Sil es isma milledgei comb. no .)
H. mossambicense Mendes, 1993 Mozambique Desc ibed as H. mossambicensis
H. mum o di (Sil es i, 1935)* Ma quesas Isl. (F .
Polynesia)
Desc ibed as Isolepisma mum o di
H. pampeanum (Sil es i, 1902)* A gen ina Type species o he genus. Wygodzinsky (1948) ga e
some cha ac e s, bu wi hou comple ing he desc ip ion
H. pa um Smi h, 2013 Ba ow Island
(Aus alia)
Sil es isma pa um comb. no ., desc ibed as
He e olepisma pa a
H. p ima um (Sil es i, 1949) Angola Desc ibed as Isolepisma p ima a
H. py amidum Smi h & Mi chell, 2019 Lo d Howe Island
(Aus .)
Sil es isma py amidum comb. no .
H. ouxi (Sil es i, 1915)* New Caledonia Desc ibed as Isolepisma ouxi
H. scle ophyllum Smi h, 2014 Aus alia (NSW) Sil es isma scle ophyllum comb. no ., desc ibed as
He e olepisma scle ophylla
H. se anoi Mendes, 2011 B azil (Pa aiba)
H. onga Mendes, 2012 Tonga
H. zelandicum (Tillya d, 1924) New Zealand A e he e y incomple e o iginal desc ip ion as
No olepisma zelandica, Wygodzinsky (1961) p o ided a
e-desc ip ion bu i does no p o ide su icien de ails
o an upda ed gene ic desc ip ion
236
ZooKeys 1260: 233–278 (2025), DOI: 10.3897/zookeys.1260.151902
Ra ael Mole o-Bal anás e al.: Re ision o he sub amily He e olepisma inae (Zygen oma)
was ex ac ed om he head and p ono um o u he specimens. DNA ex ac ion
was pe o med using a modi ied p o ocol o he EZNA Insec DNA Ki (Omega
Bio- ek, Inc., No c oss, U.S.A). A agmen o he mi ochond ial gene cy och ome
oxidase I (COI) was ampli ied using he p ime s LCO1490COL* (WYTCDACWAAY-
CRYAARGAYATYGG) and C1-N-2329COL* (ACTGTAAATATRTGATGWGCTCA).
The he mal polyme ase chain eac ion (PCR) s eps we e 94 °C o 2 min, ollowed
by 36 cycles a 94 °C o 30 s, 53 °C o 45 s, and 72 °C o 1 min, wi h a inal ex en-
sion a 72 °C o 2 min. The PCR p oduc s we e pu i ied using peqGOLD Cycle-Pu e
ki (Peqlab Bio echnologie GmbH, E langen, Ge many), and bo h o wa d and e-
e se p oduc s we e sequenced a Eu o ins Genomics GmbH (Ge many) using he
Sange sequencing me hod. The sequences we e edi ed in BIOEDIT 7.0.5.3 (Hall
1999) and checked wi h BLAST o con i m ha he sequences belong o Zygen-
oma. COI sequences o o he Zygen oma (He e olepisma inae: Visma, He e o-
lepisma; C enolepisma inae: The mobia Be g o h, 1890, C enolepisma Esche ich,
1905, Hemi elsella Smi h, 2016, Qan elsella Smi h, 2015; Lepisma inae; Neoas -
e olepisma Mendes, 1988, Lepisma Linnaeus, 1758; Maind oniidae: Maind onia)
a ailable in GenBank we e selec ed o cons uc a mo e comp ehensi e ma ix o
a phylogene ic analysis. DNA sequences we e e iewed, o hologous sequences
we e aligned using he Clus al W (Thompson e al. 1994) algo i hm implemen ed
in BIOEDIT 7.0.5.3, and hen manually checked o inconsis encies. The alignmen
is a ailable in Suppl. ma e ial 1. Subsequen ly, we calcula ed he gene ic dis ance
unde he model p-dis ance o all axa in MEGA 7 . 26 (Kuma e al. 2016).
Phylogene ic econs uc ion
The aligned ma ix was used o he phylogene ic econs uc ions using Bayes-
ian in e ence (BI) algo i hms. The sequence o Maind onia (undesc ibed spe-
cies belonging o clade A in Zúñiga-Reinoso and P edel 2019; GenBank acces-
sion numbe : MN218531) was used o oo he phylogene ic ee. Selec ion o
subs i u ion models o he phylogene ic analysis was done using Bayesian in-
o ma ion c i e ia wi h he so wa e Model inde . 2.0 (Kalyaanamoo hy e al.
2017) implemen ed on IQTREE web se e (T i inopoulos e al. 2016). Selec ed
in o ma ion was used as a p io se ing in a M . Bayes block. The BI was pe -
o med wi h he p og am M . Bayes 3.2.6 (Ronquis e al. 2012) implemen ed in
he se e CIPRES Science Ga eway 3.3 (Mille e al. 2010). Fou sepa a e uns
we e conduc ed using ou chains. Each un ini ia ed wi h a andomly gene -
a ed ee, o 10 million gene a ions, and sampled e e y 1000 ees. The ini ial
25% o he ees gene a ed we e excluded as bu n-in. Once he con e gence
o hese ou independen uns was e i ied based on me ics like he a e age
s anda d de ia ion o spli equencies and he po en ial scale educ ion ac o ,
he ou comes om hese uns we e combined, esul ing in a o al o 90,004
ees. Las ly, a consensus ee was c ea ed by applying a 50% majo i y ule,
wi h node assessmen s de e mined using pos e io p obabili y (pp).
Mo phological s udy
Insec s p ese ed in e hanol (96%) we e dissec ed, moun ed on slides using Ten-
dei o Medium and examined using an Axioskop 2 plus mic oscope (Ca l Zeiss
GmbH, Gö ingen, Ge many), equipped wi h a Zeiss AxioCam MRm came a. Fig-
237
ZooKeys 1260: 233–278 (2025), DOI: 10.3897/zookeys.1260.151902
Ra ael Mole o-Bal anás e al.: Re ision o he sub amily He e olepisma inae (Zygen oma)
u es o mo phological cha ac e s we e d awn based on hese pho og aphs, wi h
he help o GIMP2 so wa e. Fo he desc ip ion o mo phological cha ac e s,
we used he e minology o Smi h (2013) and Mole o-Bal anás e al. (2024); o
example, he e m ‘comb’ (o mac ochae ae) is used in he sense indica ed in
hese wo ks, i.e., “ ows o mac ochae ae ha a e mo e o less pe pendicula o
he la e al ma gin. They can be composed o ew (2 o 3) o a highe numbe o
mac ochae ae each” and “in some cases, a comb may be educed o a single
mac ochae a bu i s subma ginal posi ion makes i clea ha his ep esen s a
educed numbe o mac ochae a in a no mally posi ioned comb and i is coun -
ed as a comb.” Fo he desc ip ion o he posi ion o ichobo h ia on ho ac-
ic no a and o he ypes o an ennal sensilla, he sugges ions o Smi h e al.
(2021) and he e minology o Adel (1984) we e adop ed, espec i ely. Fo cylin-
d ical sensilla o he apical a icle o maxilla y palps (Mendes 1991), in sil e ish
axonomy also e e ed o as p onged sensilla (see Mole o-Bal anás e al. 2024),
we use he e m s yloconic sensilla; hey consis o se e al apical senso y cones
(sho and apically subacu e o longe and inge -shaped) ha inse in o a cy-
lind ical base (s yle). De ails o some specimens we e examined wi h a Zeiss
Sigma 300-VP scanning elec onic mic oscope (SEM). P e ious o he SEM ex-
amina ion, insec s s o ed in 96% e hanol we e dehyd a ed as ollow: 2h absolu e
e hanol ollowed by 3 × 2h hexame hyldisilazane (HMDS), inally e apo a ion
o HMDS a oom empe a u e. D ied samples we e di ec ly placed on ca bon
adhesi e discs (Plano, We zla , Ge many) on he specimen holde and subse-
quen ly spu e -coa ed wi h gold laye s. P io o dehyd a ion, some specimens
we e ul asonically cleaned, which gene ally esul ed in a loss o mos scales
and mac ochae ae. The ype ma e ial is deposi ed in he ollowing ins i u ions:
MNNC Museo Nacional de His o ia Na u al, San iago (Chile);
MZUC Museo de Zoología de la Uni e sidad de Concepción, Concepción
(Chile);
UCO Depa amen o de Zoología, Uni e sidad de Có doba, Có doba (Spain).
Resul s
Taxonomic accoun
Lapidisma gen. no .
h ps://zoobank.o g/E9D457F2-A04C-4FA5-A107-855277955B0C
Type species. Lapidisma paposanum sp. no .
Diagnosis. Medium-sized sil e ish (adul s eaching 7.0–8.5 mm). Body
shape usi o m, wi h ho ax sligh ly wide han he abdomen base. An ennae
as long as body o a li le longe ; caudal ilamen s sligh ly sho e han body.
Epide mal pigmen p esen , do sal scales pigmen ed. Mul i- adia e o bicula
scales, ounded o subquad angula , co e ing he body do sally and en ally,
also p esen on coxae and clypeus. Scales o emo a and ibiae o oid, wide,
smalle han hose o coxae and body, wi h hei base no o bicula (no su -
ounding he socke ). Sub iangula apically unca e scales p esen on scape
and pedicel o an ennae and basal a icles o maxilla y palps; some scales
showing his shape a e also p esen on emo a. Scales absen om lab um,
238
ZooKeys 1260: 233–278 (2025), DOI: 10.3897/zookeys.1260.151902
Ra ael Mole o-Bal anás e al.: Re ision o he sub amily He e olepisma inae (Zygen oma)
apical a icles o maxilla y palps, labial palps, a si, and abdominal s yli; caudal
ilamen s wi h some na ow lanceola e scales. Mac ochae ae smoo h, apically
bi u ca e. Compound eyes wi h 12 omma idia. F on al chae o axy concen a ed
on la e al pa s, wi h a wide gap in he middle. Clypeus almos e ical, wi h 1+1
la e al u s o mac ochae ae and ounded scales. Lab um olded backwa ds,
wi h a ans e se g oup o mac ochae ae no o ming clea la e al u s. An en-
nae wi h ichobo h ia and also wi h chae ic, ichoid, coeloconic and se e al
ypes o basiconic sensilla. Galea wi h a small apical peg. Maxilla y palp wi h
he apical a icle bea ing basiconic sensilla ypes B and C, and a longi udinal
ow o h ee s yloconic sensilla. Apical a icle o he labial palp wi h i e ypical
papillae a anged in a jagged line, no clea ly o ming wo lines, no a s aigh
one; he ou e la e al a ea wi h basiconic sensilla ypes B and C. An e io ma -
gin o p ono um wi h se al colla . La e al ma gins o ho acic no a wi h se e al
mac ochae ae, mos o hem isola ed, in some places o ming implied combs
o wo mac ochae ae; pos e io ma gin de oid o se ae. T ichobo h ial a eas
open, Type 1 acco ding o classi ica ion o Mendes (1986). Tho acic s e ni es
pa abolic, wi h con ex o sligh ly unca ed hind ma gin. Coxae wi h a ow o
mac ochae ae on hei an e io ma gin and on he pos e io ma gin. Me a ibiae
bea ing a long an e io ichobo h ium. Ta si wi h ou a icles; p e a sal claws
ypical, wi h a sho s ia ed medial empodial claw. U o e gi es I−VIII wi h 3+3
combs o mac ochae ae; hose a submedian posi ion can consis o 1–2+1–2
mac ochae ae (when wo a e p esen , hey a e inse ed e y close oge he and
a anged obliquely o he pos e io ma gin o he e gi e). U o e gi e IX de oid
o se ae. U o e gi e X sho apezoidal, wi h hind ma gin almos s aigh o wi h
i s median pa sligh ly con ex, wi h se e al se ae on hei la e al ma gins, lack-
ing combs. U os e ni e I wi hou se ae. U os e ni es II−VIII in male and II−VII in
emale wi h 1+1 single mac ochae ae; each coxi e VIII o emales also wi h only
one isola ed mac ochae a. Fou pai s o abdominal s yli in bo h sexes. Pa am-
e es o oid o sho subcylind ical, wi h a di ision delimi ing an apical pa wi h
some glandula se ae. Females wi h a hin o iposi o o he p ima y ype.
Rema ks. The combina ion o cha ac e s indica ed in he diagnosis o his
genus is mo e simila o hose sha ed by he Aus alian species included in he
genus Visma han o hose sha ed by Aus alian axa included in He e olepis-
ma. As he name Visma is p e-occupied (see In oduc ion), he new name as-
signed he e o his genus o Lepisma idae is Vis olepisma nom. no .; he p e-
ix ‘Vis o’- e e s o is um, a name o a cephalic ‘ iso ’ consis ing o on al
scales, which inspi ed Smi h e al. (2021) o use he name Visma o his genus.
Compa ison wi h Sou h Ame ican species assigned o He e olepisma is
mo e di icul since specimens in good condi ions belonging o hese Ame ican
axa a e no a ailable, wi h he excep ion o a ecen ly collec ed He e olepisma
om A gen ina ha is ea ed he e as H. andinum. Simila i ies and di e ences
wi h o he axa o He e olepisma inae a e p esen ed in Tables 2, 3. We em-
phasise ha , like Vis olepisma, Lapidisma gen. no . has bi id mac ochae ae
on lab um, scales on scapus, h ee slende s yloconic sensilla in he apical a -
icle o maxilla y palp, wide scales on emo a and ibiae, and sho u o e gi e
X, bu i di e s om Vis olepisma in lacking 1+1 pos e io mac ochae ae on
he hind ma gins o ho acic no a, he con ex shape o ho acic s e ni es ( un-
ca e in Vis olepisma), a highe numbe o mac ochae ae on in ala e al combs,
he shape o clypeal scales and he p esence o 3+3 combs on u o e gi e VIII
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Table 2. Compa ison o cephalic and ho acic mo phological cha ac e is ics o he new gene a o He e olepisma inae
om Chile wi h Sil es isma gen. no ., Ma i isma, and Vis olepisma nom. no . om Aus alia. Fo compa ison wi h
He e olepisma inae om o he con inen s (all o hem a ibu ed so a o he genus He e olepisma s. s .), see Table 4.
Cha ac e Lapidisma gen. no . Cac isma gen. no . Sil es isma gen. no . Vis olepisma nom. no . Ma i isma
Median gap on
on al inge
Yes Yes Va iable (some species
no gap), o he s wi h
1+1 mac ochae ae in
he middle o he gap)
Yes No
Clypeal scales P esen , ounded
(de ec ed wi h
SEM)
No de ec ed (SEM
used)
No de ec ed o
lanceola e
De ec ed in some
species, iangula o
lanceola e
No de ec ed
(SEM no used)
Bi id mac o-
chae ae on lab um
Yes Yes No, only small se ae Yes Yes
Scales on scapus Wide unca e,
sub iangula (SEM)
No de ec ed (e en
wi h SEM)
Absen . Pe haps
in some species
lanceola e
T iangula o lanceola e
(in his case, wi h an
apical inden a ion)
No de ec ed
(SEM no used)
Scales on pedicel T iangula , unca e,
o e en bi id (SEM)
No de ec ed, e en
wi h SEM
Absen (o pe haps
lanceola e in some
species)
Some imes, iangula Absen
S yloconic sensilla
on maxilla y palp
3, slende s yles
wi h se e al sho
cones
(2-)3, slende s yles
wi h se e al sho
cones
Va iable, bu mos ly
3 sho s yles wi h
se e al long cones
3, slende s yles wi h
se e al long cones
No de ec ed,
appa en ly only
wi h a basiconic
sensillum C
Scales on
maxilla y palp
Na ow lanceola e
o sub iangula
and unca e, on
basal a icle
No de ec ed, e en
wi h SEM
Absen o no de ec ed Some imes obse ed,
iangula
Absen o no
de ec ed
Papillae on labial
palp
5 compac , o ming
wo ows. Apical
a icle wide.
5 compac , o ming
an o al shape. Apical
a icle na owe .
5 usually compac ,
o ming wo mo e o
less de ined ows
5 compac , a iable
a angemen bu
ending o o m only
one ow
5 no compac ,
o ming wo
ows in a e y
wide a icle
Mac ochae ae on
pos e io ma gins
o ho acic no a
0 0 1–2 1–2 1
Scales on emo a
and ibiae
Wide, ounded;
absen in ou e
side o ibiae, wide
(SEM)
Wide, sub iangula
na owe (SEM)
Absen o lanceola e T iangula o ounded,
wide
Absen
Tho acic s e ni es Pa abolic, wi h
con ex o sligh ly
unca e hind
ma gin
Pa abolic, wi h
con ex hind ma gin
T iangula o pa abolic,
usually wi h con ex
hind ma gin
T apezoidal, wi h
unca e hind ma gin
Sub-pa abolic,
wi h con ex
hind ma gin
(2+2 in Vis olepisma in his posi ion), al hough he submedian combs o his
u o e gi e can be educed o one o wo mac ochae ae. Looking o a ini ies
wi h o he Sou h Ame ican species p e iously desc ibed and compa ing he
new axon om A acama Dese wi h he species geog aphically ound in he
Sou he n Cone, which is he opo ypic a ea o He e olepisma sensu s ic o (s.
s .), such as H. andinum and H. pampeanum, we can clea ly dis inguish he new
axon om bo h species because i has a highe numbe o s yli (one pai in
H. andinum and wo pai s in H. pampeanum). Mo eo e , H. andinum is di e en
om Lapidisma gen. no . because he la e has only 1+1 la e al mac ochae ae
on u os e ni es and H. andinum bea s 1+1 la e al combs o se e al mac ochae-
ae, 4–6 acco ding o S ach (1933); H. pampeanum is simila in his cha ac e
o he new species om A acama Dese . O he cha ac e s ha ha e p o en
o be ele an o gene ic diagnosis, such as ichobo h ial a eas, dis ibu ion,
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Table 3. Compa ison o abdominal mo phological cha ac e is ics o he new gene a o He e olepisma inae om Chile
wi h Sil es isma gen. no ., Ma i isma and Vis olepisma nom. no . om Aus alia. Fo compa ison wi h He e olepisma i-
nae om o he con inen s (all o hem a ibu ed so a o he genus He e olepisma s. s .), see Table 5.
Cha ac e Lapidisma gen. no . Cac isma gen. no . Sil es isma gen. no . Vis olepisma nom. no . Ma i isma
U o e gi e I
chae o axy
3+3 combs 2+2 combs 2+2 o 3+3 combs 2+2 o 3+3 combs 3+3 combs
A: 6 A: 2 A: 1–3 A: 1–3 A: 1
B: 4–5 B: 1 B: 1–3 B: 1–2 B: 1
C: 1 o 2 C: 0 C: 0–1 C: 0–1 C: 1
U o e gi es II–
VII chae o axy
3+3 combs 3+3 combs 3+3 combs 3+3 combs 3+3 combs
A: 7–8 A: 3–5 A: 2–5 A: 1–6 A: 1–3
B: 5 B: 2–3 B: 1–4 B: 1–4 B: 1–3
C: 1–2 C: 1–2 C: 1–2 C: 1 C: 1
U o e gi e VIII
chae o axy
3+3 combs 2+2 combs 2+2 o 3+3 combs 2+2 combs 2+2 combs
A: 8 A: 2 A: 3 A: 2–5 A: 1–2
B: 5 C: 1–2 B: C: 1 C: 1
C: 1–2 C: 1–2
U o e gi e IX
chae o axy
Wi hou se ae Wi hou se ae o
wi h ew cilia
Va iable. Ba e o ew
in a-la e al se ae
Wi h in ala e al
g oups o cilia
Wi h 3+3
in ala e al se ulae
U o e gi e X
hind ma gin
and shape
Sub apezoidal sho
o sub iangula ,
sligh ly acu e.
T unca e. Sho
apezoidal shape
Rela i ely long, ound, o
some imes app oaching
apezoidal
Usually con ex sho Sligh ly con ex
and e y sho
Scales on
basal di isions
o ce ci
Some lanceola e long,
na ow acu e scales
No de ec ed, e en
wi h SEM
Absen o na ow
lanceola e and acu e
Lanceola e, na ow,
and acu e; iangula in
some species
Absen
U os e ni e I
chae o axy
Wi hou se ae? Wi hou se ae? Va iable (wi h o wi hou
a median comb)
Wi hou se ae Wi h one median
comb
U os e ni es
II–VII
chae o axy
Wi h 1+1 single
mac ochae ae
Wi h 1+1 single
mac ochae ae
Va iable (1+1
mac ochae ae o 1+1
combs)
Usually wi h 1+1 single
mac ochae ae
Wi h 1+1
combs o 4–12
mac ochae ae
U os e ni e /
Coxi e VIII
1+1 single
mac ochae ae
Wi h 2+2
mac ochae ae
Va iable (1+1
mac ochae ae o 1+1
combs)
Wi h 1+1 single
mac ochae ae
Wi h 1–2 + 1–2
mac ochae ae
Numbe o
pai s o s yli
4 pai s in bo h sexes Only one pai 1–3 pai s; in some
species, di e ences
be ween sexes
3–6 pai s; in some
species di e ences
be ween sexes and
be ween gene ically
simila popula ions
Only one pai
Scales on s yli No isible in SEM
images
No de ec ed wi h
SEM
Absen T iangula Absen
Pa ame es La ge, o oid, as long
as he nin h coxi e and
wi h an apical di ision
No males collec ed Small, conical, mo e o
less long, smalle han
he nin h coxi e
Small, conical, mo e o
less long, smalle han
he nin h coxi e
Rela i ely la ge,
conical, almos as
long as he nin h
coxi e
and shape o scales on appendages, shape and chae o axy o ho acic s e n-
i es, cephalic chae o axy, e c., we e no men ioned in he o iginal desc ip ions
o hese species o in la e published manusc ip s ha lis ed some addi ional
cha ac e s o he A gen ine species (S ach 1933; Wygodzinsky 1948). How-
e e , examina ion o he specimen collec ed in A gen ina, which co esponds
o H. andinum in he numbe o s yli and abdominal chae o axy, sugges s ha
his species p obably belongs o a di e en genus, as i di e s om Lapidisma
gen. no . in he shape and dis ibu ion o scales on appendages, cephalic and
en al chae o axy, among o he hings (see Tables 2, 3). I is likely ha mos
cha ac e s o H. andinum a e sha ed wi h H. pampeanum, he ype species o
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Ra ael Mole o-Bal anás e al.: Re ision o he sub amily He e olepisma inae (Zygen oma)
he genus He e olepisma, al hough his equi es examining he ypes o his
species o , a leas , some ma e ial iden i ied by expe s (such as P. Wygodzin-
sky) as his species. This could con i m ou hypo hesis ha He e olepisma s.
s . is endemic o he eas e n side o he Andean moun ain ange, and ha he
Andean biogeog aphic ba ie sepa a es He e olepisma s. s . om he Chilean
gene a desc ibed in his wo k. A e-examina ion o H. annec ens om Juan
Fe nandez Islands may cla i y whe he his species should be assigned o La-
pidisma gen. no ., since some sha ed cha ac e s, such as he numbe o s yli,
he la ge size o pa ame es, and he absence o chae o axy on he hind ma gins
o ho acic no a, suppo his hypo hesis. We p opose ha his species can
be conside ed as Lapidisma annec ens comb. no . Compa ed o L. annec ens,
L. paposanum sp. no . has a highe numbe o mac ochae ae on in ala e al
combs, bu o he di e ences canno be es ablished because he incomple e
desc ip ion o Sil es i (1902).
E ymology. This gene ic name is a usion o he wo ds lapidum and lepisma,
his la e used o mos gene a o sil e ish belonging o he amily Lepisma i-
dae. Lapidum is he geni i e plu al o m o lapis, a La in wo d ha means s one,
and e e s o he usual habi a o his insec , which is ound associa ed wi h
s ones, usually unde hem. Thus, he li e al ansla ion o he wo d would be
‘ he sil e ish o s ones’. The g amma ical gende o his genus is neu e ollow-
ing he opinion o ICZN (2018) o all gene a de i ed om Lepisma.
Lapidisma paposanum sp. no .
h ps://zoobank.o g//77D018FD-4D45-4283-8776-623035A201F7
Figs 1–9
Type ma e ial. Holo ype: Chile • An o agas a Region, Paposo, La Rinconada,
unde s ones; 24°56.644'S, 70°29.70'W; 60 m a.s.l.; 01.IX.2023. Leg. A. Zúñiga.
1♀* moun ed on slide [MNNC]. Pa a ypes: Same collec ion da a as o he holo-
ype. Two specimens moun ed on slide: 1♂* [MZUC], 1♂ [UCO], wo specimens
moun ed on disc and gold coa ed: 1♂ [MZUC], 1♂ [MNNC] and wo specimens
p ese ed in e hanol 70%: 2♀ [UCO]. Chile • An o agas a Region, Paposo, El
Gaucho, unde s ones; 24°57.589'S, 70°28.489'W; 65 m a.s.l.; 29.IX.2024. Leg.
A. Zúñiga. Fou specimens, p ese ed in e hanol 70%: 1♂* [MNNC], 1♂/1♀
[MZUC], 1♂ [UCO].
Diagnosis. As indica ed o he genus. In ala e al combs wi h 6–8 mac o-
chae ae (L. annec ens has 5 mac ochae ae on each in ala e al comb). O iposi-
o su passing he apex o coxi es IX by ~3.6× hei leng h (~5× in L. annec ens).
Desc ip ion. Habi us as in Fig. 1. Body leng h o holo ype: 8 mm. Maximum
body leng h obse ed in ype se ies: 8.5 mm. Maximum p ese ed leng h o an-
ennae in ype specimens is 5 mm, bu in li e specimens an ennae a e as long
as he body o sligh ly longe . Epide mal pigmen p esen , b ownish o b own-
ish- iole , mo e in ense on head, an ennae (whe e i is uni o m), legs (especially
on ibiae, i s a some e and apex o emo a), la e al ma gins o do sal pla es,
hind ma gins o u o e gi es, da ke on he pos e io segmen s and caudal ila-
men s; en ally, he pigmen is mo e in ense on he ou e ma gins o coxi es IX
in bo h sexes and on he apex o pa ame es in males. Mac ochae ae smoo h,
bi id, hyaline o yellowish. Body co e ed wi h o bicula scales (i.e., wi h la e al
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Ra ael Mole o-Bal anás e al.: Re ision o he sub amily He e olepisma inae (Zygen oma)
wo). One ichobo h ium, appa en ly no associa ed wi h a comb, on he la e al
ma gin a ~0.6 o i s leng h, and ano he one o ano he specimen a a mo e
pos e io posi ion, associa ed wi h he penul ima e la e al comb, a ~0.8 o he
leng h o he mesono um (Fig. 6C).
The me ano um (Fig. 6D) has 10+10 combs o one o h ee mac ochae ae,
he numbe o combs wi h h ee mac ochae ae is lowe han on he mesono-
um and hese a e p esen in he an e io pa o he la e al ma gin. A la e al
ichobo h ium on he pos e io pa o he la e al ma gin a ~0.8 o he leng h
o he me ano um; he posi ion o he an e io ichobo h ium has no been
clea ly disce ned.
P es e num o p o ho ax wi h wo ans e se ows o mac ochae ae. P os-
e num subquad angula o co di o m, sligh ly longe han i s maximum wid h
( a io L/W ~1.1), wi h 1+1 an edis al combs o wo o h ee mac ochae ae, i s
hind ma gin unca e o sligh ly ounded (Fig. 6E, F); la e al ma gins wi h se e al
hin and long se ae. Mesos e num wi h a simila a io L/W and shape; he 1+1
an edis al combs wi h h ee o ou mac ochae ae, and wi h a lowe numbe o
hin and long se ae in i s la e al ma gin (Fig. 6G). Me as e num wide han long;
a io L/W ~0.81; 1+1 combs o wo o h ee mac ochae ae; hin and long se ae
less abundan and limi ed o he pos e io pa o he la e al ma gin. The hind
ma gin o his s e ni e widely ounded (Fig. 6H).
An e io ma gin o coxae wi h a dense ow o mac ochae ae, usually a -
anged in oblique combs o wo o h ee mac ochae ae; he pos e io ma gin
wi h one ow o ew mac ochae ae, mainly inse ed on he dis al pa o he
a icle (Fig. 6I).
Femo a co e ed wi h scales, mo e nume ous on he inne ( en al) side,
whe e hey a e o al o ounded, smalle han hose co e ing he body (~60–
70 µm long), wi h he apical ma gin unca ed, wi h some shallow inden a ions
(Fig. 7A); on he ou e (do sal) side, scales a e limi ed o he an e o-apical a ea
and hei shape is mo e he e ogeneous, some o hem a e na owe and/o a e
bi u ca ed on hei apical ma gin (Fig. 7B). The inne side o ibiae is co e ed
wi h ounded o o al scales, simila in shape o hose o emo a, bu smalle
(mos o hem a e <50 µm long; see Fig. 7A); he ou e side o his a icle is
appa en ly de oid o scales, only co e ed wi h se ae (Fig. 7B). Mos se ae o
emo a and ibiae de ached and only hei inse ions a e isible; some inse -
ions a e la ge , sugges ing ha hey co espond o small mac ochae ae. When
p ese ed, mac ochae ae a e sho e han he diame e o ibiae; on p o ibiae
one mac ochae a is p esen on he median pa o he an e io (do sal) ma -
gin and h ee a e isible on he hind ma gin (one basal, one median and one
subapical; see Fig. 7C); on me a ibiae wo inse ions a e isible on he an e io
ma gin and he numbe o mac ochae ae is highe a he pos e io ma gin, bu
a iable in he a ailable specimens; in SEM images inse ions o se ae wi h
an in e media e size a e isible, smalle han inse ions o mac ochae ae and
la ge han hose o no mal se ae (Fig. 7A). P o ibiae ~3.4–3.9× longe han
wide; me a ibiae ~4.1–4.7× longe han wide and ~1.25× longe han p o ibiae.
Ta some es lacking scales; a some e 1 o he hi d leg ~0.57× he leng h o he
me a ibiae, 2.25× longe han he a some e 2, ~4× longe han he a some e
3 and 2.2× longe han he ou h a some e; hese p opo ions in he i s leg
a e 0.55, 2.4, 3, and 2, espec i ely. P e a sal claws smoo h, as in Fig. 7C; he
medial empodial claw ~1/2 he leng h o la e al claws.
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Figu e 7. Lapidisma paposanum sp. no ., SEM images o legs. A. Ven al iew o he hi d leg showing scales wi h special
shape on emu and ibia; B. Ou e side o he hi d leg ( he side obse ed do sally, excep o coxa and ochan e ), scales
wi h special shape can be obse ed on he emu ; C. Ven al iew on apical pa o p o ibia wi h ibial spu and a sus wi h
p e a sal claws. Abb e ia ions: Cx: coxa, E: empodial claw, Fe: emu , L c: la e al claws, M : me as e num, Mulc: la e al
comb o u os e ni e III consis ing o wo mac ochae ae o di e en leng h, T: ochan e , Ti: ibia, T 1: i s a some e, T 4:
ou h a some e, UI/II: i s /second abdominal s e ni e (u os e ni e I/II).
U o e gi es I−VIII wi h 3+3 combs o mac ochae ae. The in ala e al combs
bea 6–8 mac ochae ae and he la e al combs 3–5. A a submedian posi ion
he e a e one o wo mac ochae ae on each side (Fig. 8A, E) which a e in e -
p e ed as educed submedian combs. When he submedian comb consis s o
wo mac ochae ae, hey a e ei he o equal size o he mo e pos e io one is
smalle (Fig. 8A, D, E). In se e al cases ( o example, on he u o e gi e VII o
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Figu e 8. Lapidisma paposanum sp. no ., abdomen, do sal. A. U o e gi es I and II wi h inse ions o mac ochae ae
(combs); B. Mic og aph showing in de ail he combs on u o e gi e I ma ked in A; C. U o e gi e V; D. U o e gi e VI; E. SEM
image o he pos e io u o e gi es IV–X (ma ked as U4–U10), Lm: mac ochae ae o la e al comb, Sm: mac ochae ae o
submedian comb; F. Hind ma gin o u o e gi e X and posi ion o epip oc and pa ap oc s; G. Mic og aph showing he
a ea ma ked in F. Scale ba s: 0.1 mm. Abb e ia ions: Im: mac ochae ae o in ala e al comb, Lm: mac ochae ae o la e al
comb, Sm: mac ochae ae o submedian comb.
he holo ype), he smalle mac ochae a is absen (Fig. 8C) o e en he en i e
comb disappea s ( o example, he holo ype has an asymme ic chae o axy on
u o e gi e VI, wi h 2+3 combs, lacking he submedian one on one side and, on
he o he side, he e is only one mac ochae a). Some small se ae a e inse ed
pos e io ly o he combs, close o he hind ma gin o he e gi e.
U o e gi e IX educed in leng h and lacking se ae. U o e gi e X sho , wi h
hind ma gin con ex, almos s aigh o ounded, in some specimens o ming an
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Figu e 9. Lapidisma paposanum sp. no ., abdomen, en al. A. U os e ni e IV wi h inse ions o la e al mac ochae ae,
he mic og aph showing he de ails o one inse ion, su ounded by addi ional inse ions o smalle se ae; B. U os e ni e
VIII wi h inse ions o la e al mac ochae ae and s yli on he pos e io ma gin; C. SEM image o he pos e io abdominal
segmen s o a male specimen, he s aigh ma gin o u os e ni e VIII is clea ly isible, while ha o he u os e ni e VII is
mos ly co e ed by scales; D. Coxi es VIII and IX o he emale holo ype wi h o iposi o , s yli IX a e de ached and only hei
inse ions a e isible; E. De ail o he le s ylus on coxi e VIII; F. U os e ni e VI wi h inse ions o s yli; G. U os e ni e VII
wi h a single s ylus p ese ed; H. Coxi es IX o a male showing he penis and pa ame es; I. De ail o pa ame e nex o he
inne p ocess o he coxi e IX o ano he male specimen; J. Apex o gonapophysis VIII; K. De ail o some ce cal di isions
showing inse ions o di e se ypes o se ae, ichobo h ia and na ow lanceola e scales. Abb e ia ions: C: ce cus, Cs:
ce cal scale, Cx9: coxi e IX (pa ame es and penis ha dly isible below i s inne ma gin), I: inse ions o s yli, In 7: in a-
la e al comb o mac ochae ae o u o e gi e VII (pa o he do sal e gi e ben en ally), Lm6/8: la e al mac ochae ae on
pos e io ma gin o u os e ni es VI/VIII, Us7/8: discs o u os e ni es VII/VIII.
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ill-de ined ob use sub iangula a ea in he middle (Fig. 8E) and in some o he s
sligh ly conca e (Fig. 8F); i has only some se ae on hei la e al ma gins bu
lacks combs o mac ochae ae.
U os e ni e I de oid o se ae, i s median pa is b oken in he holo ype (bu
isible in ano he specimen; see Fig. 7A). U os e ni es II−VIII wi h 1+1 isola ed
la e al mac ochae ae (Figs 7A, 9A). Each o hese mac ochae ae is accompa-
nied by some smalle se ae; equen ly, one o wo o hese can be iden i ied
as ichoid sensilla su ounding he la ge mac ochae ae pos e io ly. In males,
he u os e ni es VIII en i e (no di ided in o wo la e al coxi es), wi h a s aigh
hind ma gin be ween he s yli (Fig. 9B, C); in emales, each coxi e VIII has, apa
om he isola ed mac ochae a inse ed on he hind ma gin ou wa ds o he
s ylus, ano he mac ochae a inse ed inwa ds, accompanied by wo smalle
se ae (Fig. 9D).
Fou pai s o abdominal s yli in bo h sexes, inse ed on segmen s VI−IX
(Fig. 9). Coxi es IX o males as in Fig. 9H, I, wi h he inne p ocess ~1.15× lon-
ge han wide a i s base and 4× longe han he ou e p ocess. In emales, he
inne p ocess o he nin h coxi e is ~1.4–1.5× longe han wide a i s base and
3.5× longe han he ou e p ocess (Fig. 9D). The ma gins o he p ocesses
o coxi es IX a e co e ed by nume ous long and hin se ae. Males bea o oid
pa ame es, di ided in o a basal po ion and an apical segmen , his one bea ing
wo small g oups o glandula se ae (Fig. 9I); he size o pa ame es is small o
medium, no eaching he apex o inne p ocess o coxi e IX ( he la ge ones
only sligh ly sho e in la ge specimens, as in Fig. 9I, some o he s only wi h a
leng h ~1/2 he leng h o coxi e IX, as in Fig. 9H). The o iposi o has ~34 o 35
di isions and su passes he apex o coxi es IX by ~3.6× hei leng h (Fig. 9D).
Apex o gonapophyses VIII as in Fig. 9J. Epip oc well de eloped and pigmen -
ed, co e ing he base o he pa ace cus and almos as long o e en longe han
he u o e gi e X, when his e gi e has a s aigh o sligh ly conca e hind ma gin
(Fig. 8F, G), lanked by wo simila ly la ge pa ap oc s. Te minal ilamen s pig-
men ed, wi h a leas one ing o mac ochae ae on each di ision; in addi ion o
chae ic and ichoid sensilla, ichobo h ia, and na ow lanceola e scales wi h
acu e o sligh ly unca ed apex (Fig. 9K; one can be seen in Fig. 9C).
Habi a and beha iou . The habi a is on a coas al e ace close o he beach
domina ed by cac i o he species Copiapoa gigan ea Backeb. The sandy clay
soil is co e ed by many medium o small s ones (Fig. 10). Sca ce pe ennial
ege a ion is suppo ed by he s ong ogs ypical o he a ea (camanchaca),
in addi ion he a ea ecei es i egula ain all, occasionally esul ing in a sho -
li ed he baceous laye o annuals. Lapidisma paposanum sp. no . has been
ound unde s ones, clinging like The mobia o he aised s ones (Fig. 1).
E ymology. The speci ic name e e s o he illage o Paposo, whe e he new
sil e ish species has been ound.
Cac isma gen. no .
h ps://zoobank.o g/51FF634A-64E1-45FE-A258-B4782731A736
Type species. Cac isma camanchaca sp. no .
Diagnosis. Medium-sized sil e ish (adul s eaching 6.5–8.5 mm). Body
shape slende , usi o m. An ennae as long o sligh ly sho e han body; cau-
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Figu e 10. Habi a ypical o Lapidisma paposanum sp. no . in La Rinconada, Paposo
(A acama Dese , Chile), specimens we e collec ed unde he medium sized ocks.
dal ilamen s almos as long as body leng h. Epide mal pigmen sca ce, do sal
scales pigmen ed. Scales co e ing he body do sally and en ally (including
coxae) ounded o o al, o bicula , wi h mo e o less de eloped p ocess su -
ounding socke s, mul i- adia e and o di e se sizes. Scales absen on clypeus,
lab um, an ennae, maxilla y and labial palps, abdominal s yli, o iposi o , and
caudal ilamen s. Scales o emo a and ibiae wi h modi ied shape; smalle ,
sub iangula , na ow, wi h apical ma gin unca e o bi id, wi h a median in-
den a ion. Mac ochae ae smoo h, apically bi u ca e. Compound eyes wi h 12
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Ra ael Mole o-Bal anás e al.: Re ision o he sub amily He e olepisma inae (Zygen oma)
omma idia, some imes only 10 o 11 omma idia de eloped. F on al chae o axy
concen a ed on la e al pa s, wi h a wide gap in he middle showing only one
ow o se ae. Clypeus almos e ical, wi h a con inuous ans e se ield o
mac ochae ae. Lab um wi h se e al i egula ly a anged bi id se ae. An ennae
wi h ichobo h ia and also wi h chae ic, ichoid, and basiconic sensilla. Maxil-
la y palps wi h he apical a icle bea ing wo o h ee s yloconic sensilla, se e al
basiconic sensilla ype B and a single basiconic sensillum ype C. Apical a icle
o he labial palp wi h i e papillae a anged in an o al shape, ou e la e al pa
wi h se e al basiconic sensilla ype B and a single basiconic sensillum ype C.
An e io ma gin o p ono um wi h se al colla . La e al ma gins o ho acic no a
wi h se e al mac ochae ae, mos o hem isola ed, in some places o ming a
small comb o wo mac ochae ae; he e a e 1+1 pos e io combs o wo mac o-
chae ae in a e y la e al posi ion, lea ing he pos e io ma gin ba e. T ichobo h-
ial a eas open (Type 1 acco ding o Mendes’ classi ica ion). Tho acic s e ni es
pa abolic, wi h con ex o sligh ly unca ed hind ma gins. Coxae wi h ows o
mac ochae ae on hei an e io ma gin ha a e a anged in indis inc combs;
he pos e io ma gins wi h a single ow o ew long and hin mac ochae ae.
Ta si consis ing o ou a icles; p e a sus wi h wo smoo h claws and a me-
dial empodial claw. U o e gi es I−VIII wi h 3+3 combs o mac ochae ae; he
submedian combs some imes consis ing o only one mac ochae a. U o e gi e
IX wi hou se ae. U o e gi e X sho apezoidal, wi h s aigh hind ma gin, se -
e al se ae on i s la e al ma gins and pos e ola e al edges; lacking clea combs.
U os e ni e I de oid o se ae. U os e ni es II−VII wi h 1+1 single mac ochae ae;
coxi e VIII o emales wi h one o wo mac ochae ae. One pai o abdominal
s yli. Females wi h a slende o iposi o o he p ima y ype. Male unknown.
Rema ks. This new genus is di e en o Lapidisma gen. no . om Chile and
o he o he gene a o He e olepisma inae by he cha ac e s lis ed in Tables 2,
3. I is clea ly di e en o Aus alian He e olepisma due o he p esence o bi-
id mac ochae ae on lab um (only small acu e se ae in Aus alian axa), he
absence o pos e io combs o ho acic no a (p esen in Aus alian species),
sho e u o e gi e X and di e en shape o scales co e ing appendages; o
Ma i isma due o he same di e ences excep o he shape o u o e gi e X,
and addi ional cha ac e s such as he di e en u os e nal chae o axy and labi-
al palp cha ac e s. Cac isma gen. no . is di e en o Vis olepisma (p e iously
Visma) because o he absence o pos e io combs on ho acic no a, di e en
shape o ho acic s e ni es (widely unca e in Vis olepisma), sho e u o e gi e
X and lowe numbe o abdominal s yli. Di e ences o Lapidisma gen. no . in-
clude di e en shape o scales o appendages, lowe numbe o mac ochae ae
on in ala e al u o e gal combs, and lowe numbe o pai s o s yli. P oblems
a ise when compa ing Cac isma gen. no . wi h some incomple ely desc ibed
species o He e olepisma om egions o he han Aus alia, as in o ma ion
on dis ibu ion and shape o scales on appendages o chae o axy o lab um
is missing. Conside ing he cha ac e s a ailable, Cac isma gen. no . is di e -
en o o he p e iously desc ibed species assigned o He e olepisma due o
he absence o mac ochae ae on he hind ma gin o ho acic no a and sho e
u o e gi e X. Apa om Lapidisma gen. no . (and H. annec ens, now assigned
o Lapidisma), he only species sha ing bo h cha ac e s, a e he He e olepis-
ma desc ibed om A gen ina, i.e., H. andinum and he ype species o he ge-
nus, H. pampeanum. A specimen assigned o H. andinum was a ailable o ou
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Ra ael Mole o-Bal anás e al.: Re ision o he sub amily He e olepisma inae (Zygen oma)
s udy and di e ences conce ning he shape o scales in appendages (lance-
ola e scales in legs, o example) and he on al chae o axy indica e ha he
A gen ine species indeed belong o a di e en genus, so He e olepisma s. s .
co esponds only o A gen ine species (and, pe haps, o o he species o his
genus desc ibed om o he geog aphic a eas, bu no om Chile so a ); see
Tables 4, 5, and addi ional commen s in he Discussion sec ion.
E ymology. This gene ic name is a usion o he wo ds ‘cac us’ and ‘lepisma’,
he la e used o mos gene a o Lepisma idae. Cac us de i es h ough La in
om he ancien G eek wo d κάκτος used o an inde e mined spiny plan and
cu en ly used o designa ing plan s belonging o he amily Cac aceae; decay-
ing specimens o hese plan s p o ide he habi a whe e his new sil e ish has
been ound. The g amma ical gende o his genus is neu e .
Table 4. Compa ison o cephalic mo phological cha ac e is ics be ween he new gene a om Chile and he non-Aus a-
lian species o He e olepisma. Cha ac e s obse ed in H. andinum om A gen ina a e included in a sepa a e column.
Remaining species o He e olepisma a e e y he e ogeneous and hei o iginal desc ip ions o en lack cha ac e s use ul
o compa ison (see no es in he co esponding column).
Cha ac e Lapidisma gen. no . Cac isma gen. no . He e olepisma
andinum
Remaining He e olepisma (excluding Aus alian
species)
Median gap on
on al inge
Yes Yes No Yes, bu no clea ly desc ibed o se e al species.
The desc ip ion o H. pampeanum sugges s he
absence o gap
Clypeal scales P esen , ounded
(de ec ed wi h SEM)
No de ec ed (SEM
used)
No de ec ed No de ec ed in mos species
Bi id mac ochae-
ae on lab um
Yes Yes Yes No desc ibed in mos species, appa en ly p esen
in H. bise osum
Scales on scapus Wide unca e,
sub iangula
No de ec ed (SEM
used)
Lanceola e wi h
an apical
No desc ibed in mos species
Scales on pedicel T iangula , unca e
o e en bi id
No de ec ed,
(SEM used)
No de ec ed No desc ibed in any species
S yloconic
sensilla on
maxilla y palp
3, slende s yles
wi h se e al sho
cones
3, slende s yles
wi h se e al sho
cones
3, slende s yles
wi h se e al
sho cones
When desc ibed, he sensilla a e simila o hose o
Aus alian species, sho s yles wi h se e al long
cones ( lowe -shaped in H. bise osum, H. japonicum,
H. se anoi and H. onga), less complex in emales
Scales on
maxilla y palp
Na ow lanceola e
o sub- iangula
and unca e, on
basal a icle
No de ec ed,
(SEM used)
No de ec ed No desc ibed in any species
Papillae on labial
palp
5 compac , o ming
wo ows. Apical
a icle wide.
5 compac ,
o ming an o al
shape. Apical
a icle na owe .
No desc ibed
and no isible
in he a ailable
specimen
When desc ibed, 5 compac (3+2 in wo ows).
Shape a iable
Mac ochae ae on
pos e io ma gins
o ho acic no a
0 0 0 H. annec ens lacks hese mac ochae ae.
H. pampeanum also lacks pos e ola e al
mac ochae ae. The emaining species bea 1+1 o
2+2 mac ochae ae
Scales on emo a
and ibiae
Wide, ounded;
absen on ou e side
o ibiae
Wide, sub iangula
(na owe han in
Lapidisma)
Lance shaped,
wi h apical
inden a ion o
acu e
In some species lance -shaped (usually acu e,
wi hou apical inden a ion), obse ed in H.
bise osum, H. ho ni, H. mossambicense, H. se anoi
and H. onga. No desc ibed o some species
Tho acic s e ni es Pa abolic, wi h
con ex o sligh ly
unca e hind ma gin
Pa abolic, wi h
con ex hind
ma gin
Pa abolic, wi h
con ex hind
ma gin
Co di o m wi h con ex hind ma gin in mos
species, only in some he me as e num is
unca ed pos e io ly. H. pampeanum is he mos
simila o he new gene a and o H. andinum. No
desc ibed in some species
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Table 5. Compa ison o abdominal mo phological cha ac e is ics be ween he new gene a om Chile and he non-Aus-
alian species o He e olepisma. Cha ac e s obse ed in H. andinum om A gen ina a e included in a sepa a e column.
Remaining species o He e olepisma a e e y he e ogeneous and hei o iginal desc ip ions o en lack cha ac e s use ul
o compa ison (see no es in he co esponding column). *, acco ding o he o iginal desc ip ion and/o wi h he key o
S ach (1933).
Cha ac e Lapidisma gen. no . Cac isma gen. no . He e olepisma
andinum
Remaining He e olepisma (excluding Aus alian
species)
U o e gi e I
chae o axy
3+3 combs 2+2 combs 3+3 combs Fo some species 2+2 o 3+3 combs desc ibed,
o o he species da a missing
A:6 A: 2 A: 4
B: 4–5 B: 1 B: 4
C: 1 o 2 C: 0 C: 1 o 2
U o e gi es II–VII
chae o axy
3+3 combs 3+3 combs 3+3 combs 3+3 combs wi h a iable numbe o
mac ochae ae, usually less han in Lapidisma
gen. no .
A: 7–8 A: 3–5 A: 4–5
B: 5 B: 2–3 B: 4
C: 1–2 C: 1–2 C: 1–2
U o e gi e VIII
chae o axy
3+3 combs 2+2 combs 2+2 combs Some species wi h 2+2 combs, some o he s
wi h 3+3 combs.
A: 8 A: 2
B: 5, C: 1–2
C: 1–2
U o e gi e IX
chae o axy
Wi hou se ae Wi hou se ae o
wi h ew cilia
Wi hou se ae? Some species lack se ae and some o he s wi h
1+1 in ala e al wi h one o ew educed se ae
U o e gi e X hind
ma gin and shape
Sub apezoidal sho
o sub iangula ,
sligh ly acu e.
T unca e. Sho
apezoidal shape
Sho and sligh ly
con ex.
Long apezoidal in mos species, s aigh o
sligh ly con ex hind ma gin
Scales on basal
di isions o ce ci
Some lanceola e long,
na ow, and acu e
scales
No de ec ed,
e en wi h SEM
No de ec ed No obse ed in mos species
U os e ni e I
chae o axy
Wi hou se ae? Wi hou se ae? ? (damaged) Se e al species wi h a median comb, bu
wi hou se ae in some o he s. Inadequa ely
epo ed in some species, including he ype
species H. pampeanum
U os e ni es II–VII
chae o axy
Wi h 1+1 single
mac ochae ae
Wi h 1+1 single
mac ochae ae
1+1 combs wi h
se e al (3 o mo e
mac o-chae ae)*
Fo some species 1+1 single mac ochae ae,
o o he s 1+1 combs (p obably including
H. pampeanum) desc ibed, o o he species
da a missing
U os e ni e /
Coxi e VIII
1+1 single
mac ochae ae
Wi h 2+2
mac ochae ae
1+1 combs wi h 3
mac ochae ae
Mos species wi h 1+1 o 2+2 mac ochae ae
Numbe o pai s
o s yli
4 pai s in bo h sexes Only one pai Only one pai Va iable. Usually he numbe is 1–3 pai s o
s yli, wi h sexual dimo phism in some species
Scales on s yli No isible in SEM
images
No de ec ed in
SEM images
No obse ed No obse ed
Pa ame es La ge, o oid, as long
as he coxi e IX and
wi h an apical di ision
No males
collec ed
Males no
obse ed
Usually e y small, only la ge o oid in
H. annec ens and H. insula e
Cac isma camanchaca sp. no .
h ps://zoobank.o g/E232770E-672A-4FD7-BD29-FDAC0A1E7FFD
Figs 11–17
Type ma e ial. Holo ype: Chile • An o agas a Region, Cale a el Cob e, Cues a
el Cob e, unde d ied cac i o he species Copiapoa (Pilocopiapoa) sola is (F.
Ri e ) F. Ri e ; 24°17.83'S, 70°29.6'W; 750 m a.s.l.; 04.IX.2024. Leg. A. Zúñi-
ga. 1♀* moun ed on slide [MNNC]. Pa a ypes: Same collec ion da a as o ho-
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Figu e 11. Cac isma camanchaca sp. no ., li ing specimen om he ype locali y a Cues a el Cob e, A acama Dese ,
Chile. (Pho o: P. Pin o).
lo ype. One specimen moun ed on slide: 1♀ ju enile [UCO]. Two specimens
moun ed on disc and gold coa ed: 1♀ [MZUC], 1♀ [MNNC] and wo specimens
p ese ed in e hanol 70%: 1♀ adul /1♀ ju enile [UCO].
Diagnosis. As indica ed o he new genus Cac isma gen. no .
Desc ip ion. Habi us as in Fig. 11; body shape subcylind ical, mo e slende
han Lapidisma paposanum sp. no . Body leng h o holo ype: 7.5 mm. Maxi-
mum body leng h obse ed in ype se ies: 8.5 mm. Maximum p ese ed leng h
an ennae is 4.8 mm, bu in li e specimens an ennae a e as long as he body
o sligh ly sho e . Epide mic pigmen p esen , ligh b ownish, only in ense on
ma gins o head, an ennae, palps, u o e gi e X, s yli, and caudal ilamen s. Mac-
ochae ae smoo h, bi id, hyaline, o pale yellowish. Body co e ed wi h o al o
ounded scales (Fig. 12A); hose wi h o al shape a e he la ges , eaching 180 ×
110 µm, while he usual size o ounded do sal and en al scales is ~100 × 100
µm; he apical ma gin is ounded and he basal ma gin su ounds he socke
o ming mo e o less de eloped expansions (o bicula scales, Fig. 12B); all o
hem ha e e y dense pa allel ibs. Scales a e also p esen on coxae, emo a
and ibiae and appa en ly absen om he emaining appendages; he shape o
emo al and ibial scales is modi ied (see below).
Cephalic capsule wi h scales, excep on clypeus and lab um. F on al chae o-
axy ex ends om he pe iocula a eas o he on o-la e al co ne ; in he medi-
an pa o he ons is an a ea wi h only a single ow o mac ochae ae close o
he on o-clypeal su u e. The on o-la e al inge o mac ochae ae has h ee
o ou se ies o mac ochae ae, excep o a la e al pa an e io o he com-
pound eyes whe e mac ochae ae ex end o he inne pa o he ons o ming
a sub iangula u (Fig. 12C). The clypeus is mos ly co e ed by a con inuous
b oad inge o mac ochae ae consis ing o six o eigh i egula ly a anged
ows, including a ow close o he ons; an a ea wi hou se ae on he ma gin
close o he lab um. Lab um wi h a con inuous inge o densely packed se ae
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Figu e 17. Cac isma camanchaca sp. no ., abdomen, en al. A. U os e ni es IV–VII; B. Inse ion o a single mac ochae a
and su ounding small se ae on u os e ni e V; C. Pos e io abdominal s e ni es including base o o iposi o ; D. La e al
o e iew o o iposi o ; E. Apical pa s o gonapophyses VIII and pa o a ce cus in he uppe hal ; F. Mic og aph o
epip oc and pa ap oc s obse ed behind he u o e gi e X. A, B, D. Schema ic d awings; C, E. SEM images.
U os e ni e I wi hou se ae. U os e ni es II−VII wi h 1+1 isola ed mac ochae-
ae in la e al posi ions (Fig. 17A), in he holo ype one o he la e al mac ochae-
ae is missing on u os e ni e VII. The isola ed mac ochae a on each side is usu-
ally accompanied by se e al small se ae (Fig. 17B). Coxi es VIII wi h a comb
o wo se ae on hei pos e io ma gin (Fig. 17C). Coxi e IX only wi h ma ginal
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Figu e 18. Habi a ypical o Cac isma camanchaca sp. no . a Cues a el Cob e (A aca-
ma Dese , Chile) wi h d ied pa s o Copiapoa (Pilocopiapoa) sola is (F. Ri e ) F. Ri e .
se ae on hei inne and ou e p ocesses; he single mac ochae a on he apex
o he inne p ocess canno be conside ed as ma ginal. In emales, inne p o-
cess o he coxi e IX ~1.6× longe han wide a he base and ~4–5× longe han
he ou e p ocess (Fig. 17D). Only one pai o s yli on he nin h segmen ; hese
s yli a e la ge, wi h da k pigmen , and 2.2–2.3× longe han he inne p ocess
o he coxi e IX. O iposi o o p ima y ype, e y long (Fig. 17D), i s apex as in
Fig. 17E, su passing he leng h o s yli IX by 2.7–3.1× and he apex o coxi es
IX by ~7.5–8×. Gonapophyses wi h 43–45 di isions. Pa ap oc s and epip oc
la ge, he apex o he epip oc eaching o e en su passing he le el o he hind
ma gin o he u o e gi e X, wi h a median inden a ion (Fig. 17F). Caudal ila-
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Ra ael Mole o-Bal anás e al.: Re ision o he sub amily He e olepisma inae (Zygen oma)
men s bea ing bi id mac ochae ae, ichobo h ia, chae ic and ichoid sensilla;
appa en ly lacking scales (Fig. 17E). Male unknown.
Habi a and beha iou . Habi a simila o ha desc ibed abo e o L. paposa-
num, bu he ype locali y a highe al i ude on he Coas al Co dille a, ecei -
ing mo e humidi y om egula og (camanchaca). The e y sca ce pe ennial
ege a ion is domina ed by he cac us species Copiapoa (Pilocopiapoa) sola is
(Fig. 18). Adul s and ju eniles we e ound oge he unde d ied and al eady o -
ing cac i (Fig. 18), and hey escaped apidly when dis u bed.
E ymology. The speci ic name camanchaca e e s o he local name gi en
o he og ha ises om he Paci ic Ocean and egula ly p o ides wa e o he
coas al egion o he A acama Dese .
He e olepisma andinum (Sil es i, 1902)
Figs 19–20
Lepisma andina Sil es i, 1902: 230. Type locali y: A gen ina, Mendoza, Cacheu a.
He e olepisma andina (Sil es i).—Esche ich 1905: 64.
Isolepisma andina (Sil es i).—Wygodzinsky 1948: 219.
Examined ma e ial. A gen ina • La Rioja P o ince, Sanogas a, Cues a de Mi-
anda, unde s ones; 29°21.05'S, 67°46.94'W; 2000 m a.s.l.; 13. II. 2022. Leg. A.
Zúñiga. 1♀* moun ed on slide [MZUC].
Desc ip i e ema ks. The specimen used o mo phological s udies, collec -
ed in an a ea o A gen ine Andes close o he bo de wi h Chile, ma ches he
combina ion o cha ac e s a ibu ed o H. andinum in he key o S ach (1933)
and he ew use ul ai s indica ed in he obsole e o iginal desc ip ion p o ided
by Sil es i (1902). I has one pai o s yli, he u o e gi e X is sho , he pos e-
io ma gins o u os e ni es II−VI bea mo e han 1+1 mac ochae ae ( ou o
i e mac ochae ae), and he submedian combs o u o e gi es a e composed o
only one mac ochae a (a second smalle se a is inse ed obliquely o he la ge
mac ochae a, which depending on he c i e ion, could be also conside ed as
he p esence o a comb o wo mac ochae ae di e en in size). This insec is
en a i ely ea ed he e as H. andinum.
O he cha ac e is ics o his A gen ine species a e de ailed below.
F on al inge o mac ochae ae con inuous, wi hou a clea median gap, ap-
p oxima ely h ee o ou ows o mac ochae ae in he middle, wi h a wide
iangula a ea la e ally be ween ons and compound eyes (Fig. 19A). Clypeus
wi h 1+1 la e al u s o mac ochae ae (Fig. 19A); scales no isible. Lab um
wi h se e al bi id mac ochae ae i egula ly a anged in a ans e se inge.
Scales on scapus lanceola e, wi h a small inden a ion a hei apex (Fig. 19B);
scales on pedicel no de ec ed. Se e al ypes o sensilla ha e been obse ed
on he lagellum (chae ic, ichoid and ichobo h ia). Apical a icle o he max-
illa y palp bea ing h ee small s yloconic sensilla wi h ew sho cones; scales
no obse ed on maxilla y palp. P ono al colla con inuous, wi h abundan mac-
ochae ae a anged in wo o i e ows (Fig. 19C). La e al ma gins o ho ac-
ic no a wi h se e al mac ochae ae ha in some cases a e a anged in small
combs o wo o h ee mac ochae ae and in some o he s hey do no o m
clea ly de ined combs; in some posi ions isola ed mac ochae ae, which can
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be in e p e ed as educed la e al combs; a he pos e ola e al co ne he e a e
1+1 o such educed combs o mac ochae ae ha consis s o one o h ee
mac ochae ae and some small ichoid sensilla. Pos e io ma gins o ho acic
Figu e 19. He e olepisma andinum, schema ic d awings o head and ho ax. A. Chae o axy o ons, clypeus and lab um;
B. Scales o scapus; C. P ono um wi h p ese ed chae o axy on ma gins; D. Mesono um wi h p ese ed chae o axy on
ma gins; E. Pos e o-la e al igh ma gin o me ano um, showing he posi ion o an e io and pos e io ichobo h ia;
F. Hind ma gin o mesos e num; G. Hind ma gin o me as e num.
AB
C
D
E
F
G
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no a wi hou se ae (Fig. 19C, D). Two pai s o ichobo h ia on each no um;
on he p ono um, only a single ichobo h ium de ec ed a 0.69 o he la e -
al ma gin; his is p obably he pos e io ichobo h ium. On he mesono um,
he an e io ichobo h ium a 0.59 o he la e al ma gins and he pos e io
ichobo h ium a 0.75 (Fig. 19D). On he me ano um, an e io and pos e io
ichobo h ia a e obse ed a 0.65 and 0.79 o he la e al ma gins, espec i ely
(Fig. 19E). Tho acic s e ni es pa abolic, wi h 1+1 an edis al combs o h ee o
ou mac ochae ae each (Fig. 19F, G); hese s e ni es a e b oken, and he p os-
e num is also ben on he slide, so hei a io leng h/wid h is no indica ed.
Scales o emo a and ibiae lanceola e, wi h a small inden a ion a he apex
(Fig. 20A, B); ibial scales sligh ly smalle han emo al ones. U o e gi es I−IV
wi h 3+3 combs, he in ala e al comb wi h ou o i e mac ochae ae, he la e -
al combs wi h h ee o ou mac ochae ae, and he submedian comb is educed
o one o wo mac ochae ae (Fig. 20C). U o e gi es V−VII damaged, u o e gi e
VIII wi h 2+2 combs ( he submedian comb is missing). U o e gi e IX appa en -
ly lacking se ae; u o e gi e X sho , con ex, sub iangula o unca e, wi h he
pos e io ma gin almos s aigh (Fig. 20D), wi h some ma ginal mac ochae ae
ha do no o m isible combs. U os e ni e I damaged; u os e ni es II−VI wi h
1+1 combs o h ee o ou mac ochae ae (Fig. 20E). Chae o axy o abdominal
segmen VII no clea ly obse ed. Coxi es VIII wi h a small comb o h ee mac-
ochae ae, appa en ly lacking s yli, so he e is only one pai o s yli on segmen
IX. Inne p ocess o he coxi e IX ~1.35× longe han wide and 3.6× longe han
he ou e p ocess (Fig. 20F). The s yli a e ~2.3× longe han he inne p ocess
o he coxi e IX. O iposi o o he p ima y ype, sho , wi h ~32 o mo e di i-
sions, su passing he apex o s yli only by 0.5× and su passing he apex o he
inne p ocess o he coxi e IX by 2.2× (Fig. 20F); he a ailable specimen seems
o be subadul (sugges ed by he sho di isions o gonapophyses) so i is likely
ha he o iposi o is longe in adul emales. Epip oc densely co e ed by mac-
ochae ae. Caudal ilamen s b oken, hei basal di isions appa en ly wi hou
scales (Fig. 20D). Male specimens no a ailable.
Rea angemen o he sub amily He e olepisma inae
Acco ding o he molecula da a, he Aus alian He e olepisma and Vis olepis-
ma ( o me ly Visma) seem o be sis e o he Sou h Ame ican He e olepisma i-
nae (see below), and his is cong uen wi h he mo phological se o cha ac e s
analysed o he same clades. The e o e, he cu en ly ecognised genus He -
e olepisma p obably does no ep esen a na u al g oup. The p ima y e e ence
o de ining he cha ac e is ics o He e olepisma s. s . a e A gen ine species,
since he ype species o he genus, H. pampeanum, was desc ibed om A gen-
ina. Since no ecen ly collec ed specimens o H. pampeanum we e a ailable,
we ha e used specimens ha can be a ibu ed o H. andinum, ano he species
desc ibed om A gen ina oge he wi h H. pampeanum. We can conclude ha
he Aus alian He e olepisma species a e di e en om he a ailable He e o-
lepisma om A gen ina (see ele an mo phological di e ences compa ing
Tables 2, 3 wi h Tables 4, 5) and he name Sil es isma gen. no . is p oposed
o Aus alian species. The species Sil es isma michaelseni (Sil es i, 1908),
o iginally desc ibed as He e olepisma michaelseni, is p oposed as he ype spe-
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Figu e 20. He e olepisma andinum, schema ic d awings o legs and abdomen. A. Me a emu and me a ibia wi h posi-
ion o mac ochae ae and long se ae (small se ae a e no ep esen ed), and ew scales; B. De ail (mic og aph) o he
subapical a ea o he emu ma ked in A., showing se e al emo al scales; C. Hind ma gin o he igh la e al-in ala e al
pa o u o e gi e III wi h inse ions o mac ochae ae a anged in combs ( he one on submedian posi ion educed o one
mac ochae a, su ounded by smalle se ae close o he pos e io ma gin); D. U o e gi e X and base o caudal ilamen s
(b oken); E. U os e ni e IV wi h inse ions o mac ochae ae a anged in 1+1 la e al combs; F. Geni al a ea o he emale
specimen examined, showing coxi es IX, one s ylus inse ed on one o hese coxi es, and i s o iposi o . Di isions o go-
napophyses a e indica ed only o a eas whe e hey a e clea ly isible. The pos e io ma gins o coxi es VIII a e ou lined
o show ha hey lack inse ions o s yli.
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cies o he new genus Sil es isma, since i is he i s desc ibed species o
his g oup (Sil es i 1908). The diagnos ic cha ac e s de ining Sil es isma gen.
no . a e he ollowing: on al chae o axy wi h a median gap wi hou se ae o
wi h 1+1 median mac ochae ae; al e na i ely wi h a con inuous inge connec -
ing la e al a eas o mac ochae ae. Lanceola e scales o en p esen a leas on
emo a, equen ly on ibiae and scape (excep o some species), appendages
wi hou iangula scales and only wi h ounded scales on coxae. S yloconic
sensilla o he las a icle o maxilla y palp usually lowe -like (i.e., sho , wide
and wi h se e al b anches), al hough i is slende in emales o some species
whe e he s yloconic sensilla show sexual dimo phism. Papillae o he las a -
icle o he labial palp compac , 3+2 usually a anged in a cu ed clus e ( wo
cu ed ows). P ono al colla comple e. Pos e io ma gin o ho acic no a wi h
1–2 +1–2 mac ochae ae. La e al combs o ho acic no a wi h 0–3 mac ochae-
ae. Tho acic s e ni es usually iangula o co di o m, wi h con ex hind ma gin
(some imes pa abolic and/o hind ma gin o me as e num sligh ly unca e).
U o e gi e I wi h 2+2 o 3+3 combs o mac ochae ae (when he e a e 2+2, hose
on submedian posi ion a e missing). U o e gi es II–VII wi h 3+3 combs o mac-
ochae ae; submedian combs small, wi h one o wo mac ochae ae. U o e gi e
VIII wi h 2+2 o 3+3 combs o mac ochae ae (when he e a e 2+2, hose on
la e al posi ion a e missing). U o e gi e IX wi hou se ae o wi h ew in ala -
e al small se ae (acu e se ae o educed mac ochae ae). U o e gi e X long,
pa abolic o apezoidal depending on whe he i s hind ma gin is con ex o
s aigh . U os e ni e I glab ous o wi h one small median comb o mac ochae-
ae. U os e ni es II−VIII o males and II−VII o emales wi h 1+1 isola ed la e al
mac ochae ae o 1+1 small combs (usually wi h 2–5 mac ochae ae packed
oge he ). One o wo pai s o abdominal s yli, some imes one addi ional pai in
emales o some species. Pa ame es small, usually less han 1/2 he leng h o
he inne p ocess o coxi e IX.
Di e ences be ween Sil es isma gen. no . and he emaining lineages o
he sub amily He e olepisma inae a e p esen ed in Tables 2–5 and commen -
ed upon in he Discussion. Table 1 includes he species ha can be assigned
o he genus Sil es isma gen. no . (all p e iously desc ibed om Aus alia),
and hose ha emain as belonging o He e olepisma. The species He e olepis-
ma howense Wome sley, 1942 om Lo d Howe Island can be assigned o his
genus because hei cha ac e s, de ailed by Smi h and Mi chell (2019), clea ly
ma ch wi h he diagnosis abo e. The possible a ini ies o he emaining spe-
cies a e commen ed upon in he Discussion.
Molecula analysis
The ma ix o COI gene sequences conside s 16 species o Lepisma idae
(L. paposanum / C. camanchaca wi h sequences o 2/3 specimens, esp.) and
Maind onia sp. as ou g oup and has a leng h o 614 bp, o which 281 a e poly-
mo phic si es and 250 a e pa simony in o ma i e. The subs i u ion model se-
lec ed by Bayesian In o ma ion C i e ia was TIM2+F+G4.
The ee ob ained (Fig. 21) shows Lepisma inae (Lepisma saccha ina Linnae-
us, 1758 + Neoas e olepisma sp.) as sis e o he emaining axa (pp = 1), he la -
e wi h C enolepisma inae and He e olepisma inae as sis e g oups (pp = 0.95).
C enolepisma inae is di ided (pp = 1) in o a clade wi h The mobia domes ica
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(Packa d, 1873) and (C enolepisma longicauda um Esche ich, 1905 + C enolepis-
ma cal um (Ri e , 1910)) (pp = 1) and a clade including he Aus alian Hemi elsel-
la and Qan elsella (pp = 1). He e olepisma inae is di ided in o a clade wi h he
Aus alian membe s as sis e o he Sou h Ame ican axa (pp = 0.99). Wi hin he
Aus alian clade, Sil es isma gen. no . (i.e., S. cooloola (Smi h, Mi chell, Lee & Es-
pinasa, 2019) + S. coo ongooba (Smi h, Mi chell, Lee & Espinasa, 2019) is sis e
o he genus Vis olepisma (pp = 1). Wi hin he Sou h Ame ican He e olepisma i-
nae, H. andinum is eco e ed as sis e o he es (pp = 1). The la e shows, wi h
low node suppo (pp = 0.6), C. camanchaca as sis e o an undesc ibed He e o-
lepisma inae om sou he n Chile and L. paposanum (pp = 1). Gene ic dis ance
be ween gene a o Sou h Ame ican He e olepisma inae is ≥ 22%, while in aspe-
ci ic dis ance o he ma e ial s udied is ≤ 2% (see Suppl. ma e ial 2).
Discussion
This s udy is a i s s ep owa ds a be e unde s anding o he di e si y o he
He e olepisma inae o he wo ld; ou desc ip ion o new gene a also allows a
mo e de ailed compa ison wi h he ai ly well-s udied auna o Aus alia. The
Figu e 21. Bayesian phylogene ic ee based on COI gene sequences o Lepisma idae. Pos e io p obabili y alues a e
gi en a he nodes. Sequences ob ained om GenBank ha e he accession numbe added a e he name. An as e isk
(*) deno es he holo ype specimen.
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p esen ed phylogeny is p elimina y and based on a single gene only, bu shows
He e olepisma inae as a monophyle ic g oup and sis e o C enolepisma inae.
Lepisma inae appea as sis e o He e olepisma inae + C enolepisma inae and
his does no suppo he hypo hesis o Mendes (1991) who sugges ed He e o-
lepisma inae as he mos plesiomo phic g oup and sis e o he o he sub am-
ilies o Lepisma idae. Ou mo phological and molecula analyses also suppo
he assignmen o he wo newly desc ibed Chilean gene a o He e olepisma i-
nae. Al hough he e a e some mo phological simila i ies be ween Aus alian
Vis olepisma and Chilean Lapidisma gen. no ., molecula da a do no suppo
a pa icula close ela ionship be ween Aus alian and Chilean He e olepisma i-
nae. The Aus alian axa appea as sis e o all analysed Sou h Ame ican axa,
including H. andinum.
He e olepisma inae now o mally consis s o six gene a: He e olepisma, Vis-
olepisma, Ma i isma, Sil es isma gen. no ., Lapidisma gen. no ., and Cac is-
ma gen. no ., which we e all conside ed in ou e ision, including a i s molec-
ula analysis using COI gene sequences (excep o Ma i isma). Al hough his
ep esen s he mos comp ehensi e e ision o his sub amily o da e, he ax-
onomy o He e olepisma inae is a om being cla i ied. Some open axonomic
ques ions a e discussed below.
Ou mo phological and molecula esul s show ha Sil es isma gen. no .
dese es o be sepa a ed om he opo ypic Sou h Ame ican He e olepisma.
The cha ac e s p esen ed he e o H. andinum a e likely sha ed wi h H. pam-
peanum, excep o he u os e nal chae o axy, al hough he co ec s a e o his
la e ai in H. pampeanum is doub ul. Wygodzinsky (1948) and S ach (1933)
men ioned 1+1 mac ochae ae on u os e ni es o H. pampeanum, bu he o igi-
nal desc ip ion by Sil es i (1902) indica es 1+1 small combs o se e al se ae
on he same posi ion, which is also he hypo hesis p e e ed by Smi h (2013),
bu his needs o be con i med. Ne e heless, o he momen we a ibu e o
He e olepisma s. s . he cha ac e s p esen ed in his wo k o H. andinum. The
a ailable specimen o his species is no in a pe ec condi ion, so a comple e
edesc ip ion o he species H. andinum canno be gi en ( o example, we can-
no con i m he s a e o cha ac e s such as he a angemen o papillae on he
las segmen o labial palps, shape and a io leng h/wid h o ho acic s e ni-
es, chae o axy o u o e gi es I, V, VI, VII, IX, u os e ni es I, VII, shape o do sal
scales, leng h o an ennae and ce ci), bu an impo an numbe o o he cha -
ac e is ics o his insec om A gen ina has been p o ided, in o de o compa e
he species wi h mo e ecen ly desc ibed axa o He e olepisma inae, including
he new Chilean gene a desc ibed in his wo k.
The di e ences o Sil es isma gen. no . o Ma i isma and Vis olepisma
we e de ailed in Smi h and Mi chell (2019) and in Smi h e al. (2021), espec-
i ely and summa ised in Tables 2, 3. Tables 4, 5 also lis he di e ences o
Lapidisma gen. no . and Cac isma gen. no .; he absence o chae o axy in he
pos e io ma gins o ho acic no a o he Chilean gene a is one o he mos
ema kable di e ences, as well as he absence o lanceola e apically acu e
scales in he Chilean gene a (excep o he e minal ilamen s o Lapidisma
gen. no .), ha a e eplaced by ounded o sub iangula , apically unca e
scales. The p esence o chae o axy in pos e io ma gins o ho acic no a in
Sil es isma gen. no . is also a di e ence o he A gen ine He e olepisma (con-
i med in H. andinum), he la e co esponding o he genus He e olepisma s.
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s . Fu he di e ences o He e olepisma s. s . will be con i med when he de-
sc ip ion o he ype species o he genus (H. pampeanum) will be upda ed, bu
he cha ac e s obse ed in H. andinum and hose a ailable o H. pampeanum
sugges ha he di e en ype o s yloconic sensilla on he maxilla y palps and
he di e en shape o u o e gi e X, which is sho e in He e olepisma s. s ., as
well as he p obably la ge pa ame es (cha ac e s sha ed wi h he new Chilean
gene a) can be p esen ed as key cha ac e s o dis inguish bo h gene a.
Ne e heless, he cha ac e s desc ibed as a iable in he diagnosis o Sil-
es isma gen. no . indica e ha he genus could spli in o wo o mo e gene a
o subgene a. Ou molecula analyses o COI gene sequences o wo species
o Sil es isma gen. no . e ealed a dis inc gene ic dis ance (>17%) (see Sup-
pl. ma e ial 2) ha could be co ela ed wi h he a iable mo phological ai s
(e.g., p esence o absence o lanceola e scales, o median gap wi hou se ae in
ons, o median comb on u os e ni e I).
He e olepisma zelandicum (Tillya d, 1924), desc ibed om New Zealand as
No olepisma zelandica, could be congene ic wi h Sil es isma gen. no ., and in
his case he name No olepisma would ha e p io i y o designa ing he Aus-
alian genus, bu his species equi es e ision since he o iginal desc ip ion
and he commen s la e gi en by Wygodzinsky (1961) do no p o ide enough
in o ma ion abou he mo phological di e ences o simila i ies wi h species
om Aus alia.
The name Isolepisma Esche ich, 1905 is also disca ded o he new Aus-
alian genus o He e olepisma. This name was used as a synonym o He e o-
lepisma and F. Sil es i used i o desc ibing some species om Ame ica and
o he a eas. The genus Isolepisma was in oduced o he species I. ise osum
Esche ich, 1905, a species ha can be conside ed species inqui enda since he
cha ac e s de ining he species a e e y con using and i has been ci ed om
e y sepa a ed geog aphic egions (B azil, Angola, Indonesia) (Mendes 2011).
Fu he species desc ibed om o he geog aphic egions (Ca ibbean a ea,
A ica, e c.) as belonging o He e olepisma should be analysed in u u e e-
sea ch o check he possibili y o being congene ic wi h He e olepisma s. s . o
wi h o he gene a o He e olepisma inae, i.e., Ma i isma in Smi h and Mi chell
(2019) and Visma (now Vis olepisma) in Smi h e al. (2021), o new gene a
should be e ec ed o hem. Conside ing wha is al eady known abou he p e-
iously desc ibed species ound ou side Aus alia, we can al eady s a e ha
H. annec ens om Juan Fe nández Islands ag ees e y well wi h Lapidisma
gen. no . due o he absence o pos e ola e al mac ochae ae on ho acic no a,
he size o pa ame es and he numbe o pai s o s yli, and i is p oposed he e
o assign i o he genus Lapidisma.
Mo eo e , we can hypo hesise ha :
– The undesc ibed He e olepisma inae om Talcahuano, Chile (36°43'S,
73°07'W), included in ou phylogene ic ee (Fig. 21), is a leas gene ically,
a he closely ela ed o Lapidisma gen. no ., bu his axon s ill needs o
be desc ibed mo phologically o decide whe he i is congene ic wi h Lapi-
disma paposanum sp. no . o dese es i s own genus, gi en he la ge ge-
ne ic dis ance o mo e han 20% o L. paposanum (see Suppl. ma e ial 2).
–Species o He e olepisma om he Asia-Paci ic egion (H. japonicum
(Uchida, 1968), H. mum o di (Sil es i, 1935), H. ouxi (Sil es i, 1915),
