Omaliine o e bee les in Eocene Bal ic ambe 35
Unexpec ed palaeodi e si y o omaliine
o e bee les in Eocene Bal ic ambe
(Coleop e a, S aphylinidae, Omaliinae)
Alexey V. Sha in1, Shûhei Yamamo o2
1 Ins i u e o Li e Sciences and Technologies, Dauga pils Uni e si y, Vienibas 13, Dauga pils, LV-5401, La ia
2 In eg a i e Resea ch Cen e , Field Museum o Na u al His o y, 1400 S Lake Sho e D i e, Chicago, IL
60605-2496, USA
Co esponding au ho : Alexey V. Sha in ([email p o ec ed])
Academic edi o : Adam B unke|Recei ed 19 Ma ch 2019|Accep ed 19 May 2019|Published 11 July 2019
h p://zoobank.o g/763EDE2B-5F0C-414D-8289-D37765E993E4
Ci a ion: Sha in AV, Yamamo o S (2019) Unexpec ed palaeodi e si y o omaliine o e bee les in Eocene Bal ic ambe
(Coleop e a, S aphylinidae, Omaliinae). ZooKeys 863: 35–83. h ps://doi.o g/10.3897/zookeys.863.34662
Abs ac
Fossil eco ds o he sub amily Omaliinae a e agmen a y and mos o hem a e less in o ma i e comp ession
ossils. Bal ic ambe om he mid-Eocene o no he n Eu ope is one o he mos impo an sou ces o
insec ossils, bu only wo eliably placed omaliines ha e been desc ibed. He e, we p o ide a gene al
o e iew o his sub amily in Bal ic ambe . In o al, i e new ex inc species o ou gene a in h ee ibes a e
desc ibed and illus a ed: Geod omicus bal icus sp. no . (An hophagini), Eusphale um kan isp.no . (Eus-
phale ini), Pa aphloeos iba mo osa sp. no ., Phyllod epa daedali sp. no ., and Ph. ica i sp. no . (Omaliini).
Addi ionally, we epo on ou species belonging o Eusphale um, which emain unnamed, om he same
ambe deposi . The eco ds o Eusphale um include he i s ossils o he ibe Eusphale ini, while ha
o Geod omicus may ep esen he second and he i s de ini i e ossil eco d o he genus and ibe An-
hophagini. Ou disco e ies highligh he unexpec ed palaeodi e si y o Omaliinae in Bal ic ambe , u he
ein o cing he coexis ence o he mophilous and empe a e-lo ing bee les in Bal ic ambe and po en ially
indica ing we land and ipa ian habi a s o ambe -p oducing o es s.
Keywo ds
An hophagini, Eusphale ini, Omaliini, ossil, mic o-CT
ZooKeys 863: 35–83 (2019)
doi: 10.3897/zookeys.863.34662
h p://zookeys.penso .ne
Copy igh Alexey V. Sha in, Shûhei Yamamo o. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License
(CC BY 4.0), which pe mi s un es ic ed use, dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed.
RESEARCH ARTICLE
Launched o accele a e biodi e si y esea ch
A pee - e iewed open-access jou nal
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
36
In oduc ion
Wi h 1639 species in 117 ex an and 14 ex inc gene a (Thaye 2016; A.F. New on un-
published da abase 17 Jan. 2019), he o e bee le sub amily Omaliinae (S aphylinidae)
is a ela i ely la ge g oup, cu en ly composed o se en ibes: An hophagini Thomson,
1859, Aphaenos emmini Peye imho , 1914, Co neolabiini S eel, 1950, Co yphiini Ja-
kobson, 1908, Eusphale ini Ha ch, 1957, Had ogna hini Po e in, 1929 and Omaliini
W.S. MacLeay, 1825. Howe e , McKenna e al. (2015) ecen ly demons a ed he non-
monophyly o Omaliinae wi h espec o h ee o he sub amilies (i.e., Empelinae New-
on & Thaye , 1992, Glypholoma inae Jeannel, 1962, and Mic osilphinae C owson,
1950) in he "Omaliine g oup o sub amilies based on wo molecula ma ke s. Thei
esul suppo s he ea lie esul s o Thaye (2000) based on la al mo phology (al hough
la ae o Empelinae a e s ill unknown and he e o e we e no included in he s udy).
These ou sub amilies combined o med a monophyle ic clade, sis e o P o eininae
(McKenna e al. 2015). Membe s o he Omaliinae a e dis ibu ed wo ldwide, wi h
he g ea es di e si y in he Hola c ic and O ien al egions, p edominan ly in mon ane
a eas. A e ision and clea diagnosis o Omaliinae s ill do no exis , and hus, he s a-
us o many sup aspeci ic axa is s ill unclea due o he di icul ies o o mally placing
hem wi hin ibes (New on and Thaye 1992, 1995). Omaliinae, o e en he Omali-
ine g oup, ha e o en been conside ed o be plesiomo phy- ich among S aphylinidae
(Thaye 2016). Howe e , his was no suppo ed by he comp ehensi e molecula phy-
logene ic s udy by McKenna e al. (2015). The p esence o pai ed ocelli in mos axa o
Omaliinae has o en been ega ded as one o he mos impo an cha ac e s o de ine he
sub amily. Ne e heless, i is unclea whe he ocelli should be in e p e ed as p imi i e o
e en apomo phic (New on and Thaye 1995; Leschen and Beu el 2004; Cai e al. 2013;
Thaye 2016). The ex an omaliines a e u he cha ac e ized by ha ing an ennae in-
se ed unde he la e al ma gins o he ons, a sal o mula 5-5-5 (excep Co neolabiini,
4-4-4; S eel 1950), p ocoxal ca i ies opened behind, well-de eloped p os e nal and pos -
p ocoxal p ocesses, p ocoxae conical and p ominen , abdomen wi h six isible s e ni es,
abdominal in e segmen al memb anes a ached apically and wi h b ick-wall-like pa e n
o scle i es, all spi acles well de eloped and unc ional, p esence o wing- olding pa ches
o mic o ichia on some abdominal e gi es, and an e io p ojec ion o abdominal s e -
ni e VIII wi h well-de eloped de ensi e glands (e.g. Klinge 1980; De ne and Reissen-
webe 1991), as well as some ea u es o geni alia and geni al segmen s o bo h male and
emale (Thaye 1985; New on e al. 2000; Pe is e al. 2014; Zane i e al. 2016).
A b ie his o y o ossil Omaliinae was ecen ly p o ided by Cha zimanolis (2018).
Fossil eco ds o omaliines a e ela i ely p e alen . Compa ed o he high di e si y and
abundance o ex an Omaliinae, howe e , he eco ds o ex inc omaliines a e s ill sig-
ni ican ly agmen a y. Many o hem a e desc ibed wi h sho desc ip ions, incomple e
illus a ions and p oblema ic sys ema ic placemen s (Cha zimanolis 2018). Hence, i
p e en s a compa ison o hese ossils o each o he and o ecen axa. In addi ion,
se e al ex inc gene a a e known as “ ibe ince ae sedis” and no placed in any o he
ibes mainly because o hei poo p ese a ion o di icul y in in e p e ing hei mo -
Omaliine o e bee les in Eocene Bal ic ambe 37
phology (Schau uss 1890; Tikhomi o a 1968; Ry kin 1985, 1990; He man 2001).
Cha zimanolis (2018) lis ed i e Ju assic omaliine gene a (Tikhomi o a 1968; Ry kin
1985): †A chod omus Tikhomi o a, †Globoides Tikhomi o a, and †Po hod omus Tik-
homi o a om he Uppe Ju assic o Ka a au, Kazakhs an; †Eophyllod epa Ry kin
om he Middle-Uppe Ju assic o No ospasskoe, Russia; and †Mo sum Ry kin om
he Middle Ju assic o Kubeko o, Wes e n Sibe ia. Two gene a †Daiod omus Ry kin
and †P odaia Ry kin a e known om he Uppe Ju assic o Daya, Russian T ansbai-
kalia, al hough Cha zimanolis e al. (2012) ega ded hem as Lowe C e aceous axa.
La e , Cai and Huang (2013) added he ex inc genus †Sinan hobium Cai & Huang
om he Middle Ju assic o Inne Mongolia, China. Compa ed o Ju assic comp es-
sions, only a hand ul o ossils a e known om he C e aceous, wi h only a single com-
p ession ossil, genus †Mesodeliph um Ry kin, desc ibed om he Lowe C e aceous o
Tu ga, T ansbaikalia (Ry kin 1990). In addi ion, Pe is e al. (2014) ecen ly epo ed
he i s Mesozoic ambe genus †Duocalca Pe is & Thaye (Omaliini) om opaque
Lowe C e aceous F ench (Cha en es) ambe and isualized he bee le ossil by using
phase-con as Synch o on Radia ion x- ay mic o omog aphy (PPC-SR X- ay μCT).
The Cenozoic omaliine auna is also a om well unde s ood. Scudde (1900)
desc ibed he comp ession ossil species Geod omicus abdi us Scudde (An hophagi-
ni), om he Uppe Eocene o Flo issan , USA, al hough he p ese a ion is no ad-
equa e o assess i s sys ema ic posi ion. Fo example, he whole head was los p io o
desc ip ion (Scudde 1900), which makes i s gene ic iden i ica ion highly doub ul
o impossible. Ne e heless, acco ding o he o iginal desc ip ion, he gene al body
shape o G. abdi us is simila o ha o Geod omicus o some o he ela ed gene a
( o example Mic oedus), bu he e is no in o ma ion abou s uc u e o he head and
mou hpa s. Ano he ossil species, Omalium an iquo um Wickham (Omaliini), wi h
P o einus-shaped body (Wickham 1913: pl. 5 ig. 3), is desc ibed om he same ho i-
zon (Wickham 1913), bu again, he p ese a ion o his ossil is insu icien o jus i y
i s axonomic placemen . Al hough he desc ip ion is oo sho o inal conclusions
(Wickham 1913), his axon may no belong o Omalium due o he p opo ions o
he ma kedly ans e se p ono um and wide ely a, which a e consis en wi h many
P o einus (P o eininae) species. O he p e-Qua e na y eco ds include An hophagus
giebeli Heyden & Heyden (An hophagini) om he Oligocene o Ge many (Heyden
and Heyden 1866), Omalium p o ogaeae Hee om he Miocene o C oa ia (Hee
1847) and some unnamed ossils (e.g. Hopkins e al. 1971; A chibald and Ma hewes
2000; Kisele and Naza o 2009). All hese comp ession ossils need e-examina ion as
gene ic assignmen s by he ea ly paleon ologis s a e doub ul. Fo example, A. giebeli
can be assigned nei he o An hophagus no An hophagini. Based on he small body
wi h wo da kened spo s in middle o he p ono um, i may belong o Eusphale um
o a small Omaliini (Phloeonomus-like) conside ing he shape o he e y wide abdo-
men (Heyden and Heyden 1866). Schau uss (1890) desc ibed Pseudoles eua insinuans
Schau uss om he mid-Eocene Bal ic ambe (ca 44 Ma) bu his ossil may no e en
belong o Omaliinae (Zane i e al. 2016). Un o una ely, Schau uss’s ma e ial was
likely o be los o sca e ed du ing Wo ld Wa II (Vi ali 2006); hus, i is p obably
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
38
impossible o e-examine he ype specimen. The only de ini i e omaliines in Bal ic
ambe we e ecen ly desc ibed by Zane i e al. (2016), ep esen ing wo Omaliini spe-
cies: Pa aphloeos iba elec ica Zane i e al. and Phyllod epa an iqua Zane i e al. They
we e isualized wi h he PPC-SR X- ay μCT me hod, illumina ing ine mo phologi-
cal de ails. Fu he , Hieke and Pie zeniuk (1984) no ed an “An hobium” om Bal ic
ambe iden i ied by E. Rei e ; howe e , his species has no been o mally desc ibed
(see also he his o y o con usion be ween An hobium, La h imaeum and Eusphale um
in To enham (1939) and Sha in and Sme ana (2017)).
The p esen pape p o ides an o e iew o he ema kable and unexplo ed palaeo-
auna o Omaliinae in Bal ic ambe . We epo a leas 18 ossil bee les in se en ambe
pieces, wi h he desc ip ions o i e new ex inc species in ou ex an gene a om he
ibes An hophagini, Eusphale ini and Omaliini. The new species and unnamed speci-
mens o Eusphale um K aa z ep esen he i s de ini i e ossil o Eusphale ini, while
ha o Geod omicus Red enbache may ep esen he second and he i s de ini i e
ossil eco d o he genus and An hophagini. These disco e ies a e signi ican o u u e
phylogene ic and paleon ological s udies o he sub amily Omaliinae and ela ed axa.
Ma e ials and me hods
Eigh een adul s in se en Bal ic ambe pieces we e used in ou s udy. Nea ly all s udied
ma e ial is deposi ed in he Gan z Family Collec ions Cen e , Field Museum o Na u-
al His o y (FMNH), Chicago, USA, wi h he assigned specimen numbe s om FM-
NHINS-3260628 o FMNHINS-3260632 wi h addi ion o FMNHINS-3965993,
bu a single ambe piece is de i ed om he p i a e collec ion o V.I. Aleksee (Ka-
lining ad, Russia), unde he egis a ion numbe AWI-045. Each piece o ambe is
placed in a small anspa en ec angula plas ic en elope wi h he labels wi hin. The
age o Bal ic ambe is o g ea deba e, wi h es ima es om he Lowe Eocene o Lowe
Oligocene (e.g. Pe ko sky e al. 2007; Wei scha and Wicha d 2010; Aleksee 2013;
Bog i e al. 2018), al hough i is gene ally accep ed as middle o uppe Eocene. He e
we en a i ely ollow he mid-Eocene (Lu e ian: 44.1 ± 1.1 Ma) age based on he
mos ecen es ima ions ob ained by he absolu e da ing analyses o glauconi es om
Sambia Peninsula (Wapple 2005). The s aphylinid auna in Bal ic ambe is di e se
and abundan , wi h he ollowing 12 ex an sub amilies eco ded: Aleocha inae, Eu-
aes he inae, Omaliinae, Oxypo inae, Paede inae, Pies inae, P o eininae, Pselaphinae,
Scydmaeninae, S aphylininae, S eninae, and Tachypo inae (e.g. Cha zimanolis and
Engel 2011; Aleksee 2013; Cai e al. 2017; Yamamo o and Ma uyama 2017). The
ambe pieces o igina e om he Bal ic Sea Coas : Yan a ny, Kalining ad, Russia (FM-
NHINS-3965993, FMNHINS-3260629, FMNHINS-3260632, AWI-045); Wisła
Ri e , Gdańsk, Poland (FMNHINS-3260628); and he Bal ic Sea Coas wi hou u -
he in o ma ion (FMNHINS-3260630). The second au ho (SY) u he p epa ed
wo specimens (FMNHINS-3260628 and FMNHINS-3260629) by polishing wi h
eme y pape s o di e en g ain sizes and a plas ic bu ing clo h.
Omaliine o e bee les in Eocene Bal ic ambe 39
The ollowing measu emen s a e used in his pape and abb e ia ed as ollows:
HW maximum wid h o head including eyes;
HL leng h o head ( om base o lab um o neck cons ic ion along
head midline in do sal iew o om apical ma gin o men um
o neck cons ic ion in en al iew (G. bal icus sp. no .));
OL ocula leng h (longi udinal);
PLL×PLW (II, III) leng h×wid h o segmen s II and III o labial palpi;
PML×PMW (III, IV) leng h×wid h o segmen s III and IV o maxilla y palpi;
AL leng h o an enna;
PL leng h o p ono um;
PW maximum wid h o p ono um;
ESL su u al leng h o ely a (leng h o ely a om he apex o scu el-
lum o he pos e io ma gin o su u al angle);
EW maximum wid h o ely a oge he ;
MTbL leng h o me a ibia;
MT L leng h o me a a sus;
AW maximum wid h o abdomen (a segmen IV);
TL o al leng h ( om an e io ma gin o clypeus o apex o abdomen).
All measu emen s a e gi en in millime e s and we e made wi h a s e eoscopic mi-
c oscope equipped wi h an ocula mic ome e . Some measu emen s o he body we e
di icul o do because o he specimen's pa ial isibili y and o ien a ion wi hin he
ambe pieces; he esul ing app oxima e alues a e ma ked wi h “~”, and he cases
when measu emen s we e no possible a e ma ked wi h “?”. The desc ip ion o he
p ese a ion o he ma e ial is gi en below he ype ma e ial lis ing in a sepa a e pa a-
g aph. The ype labels a e ci ed in in e ed commas and sepa a ed om each o he by
a comma, di e en lines in labels o he ypes and his o ic labels a e sepa a ed wi h ‘|’;
explana ions o he ype labels a e gi en in squa e b acke s, necessa y no es wi hin he
label a e gi en in angle b acke s.
Specimens we e examined using Nikon SMZ 745T and Nikon Eclipse E200 s e e-
omic oscopes. A digi al came a (Sony Alpha DSLR-A300) was used o pho og aphs o
habi us o Geod omicus bal icus sp. no . O he pho og aphs we e p oduced using a Can-
on 80D digi al came a wi h a Canon MP-E 65 mm mac o lens (F2.8, 1–5×), equipped
wi h a Canon MT-24EX mac o win li e lash as ligh sou ce. Then, image s acks we e
ca ied ou using CombineZM so wa e (Alan Hadley, She ield, UK). All igu es we e
modi ied using Adobe Pho oshop so wa e. Fo one pa a ype (FMNHINS-3260630) o
Eusphale um kan i sp. no ., images we e gene a ed using x- ay mic o-compu ed omog-
aphy (μ-CT), acqui ed wi h a mic o- ocus x- ay CT sys em (inspeXio SMX-100CT;
Shimadzu) h ough he cou esy o Shimadzu Co p. (Kyo o, Japan). I was scanned a
60 kV unde 60 μA, esul ing in a oxel size o 5.0 μm. Speci ic se ings o he scan a e
con iden ial and e ained by he company. Rende ing o he image olume was ca ied
ou using VGs udio max . 2.2 (Volume G aphics, Heidelbe g, Ge many).
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
40
Sys ema ic Palaeon ology
O de Coleop e a Linnaeus, 1758
Family S aphylinidae La eille, 1802
Sub amily Omaliinae MacLeay, 1825
Sys ema ic placemen o ossils. The cha ac e s o he sub amily by which he ossil
specimens desc ibe he e a e unambiguously e e ed o Omaliinae a e: shape o he body
is a iable bu in gene al mo e o less wide, wi h sho and less lexible abdomen han
mos s aphylinids; ely a a e a iable in leng h and some imes dis inc ly elonga e and
co e ing he en i e abdomen (New on and Thaye 1995); do sal su ace o he head,
mo e o less close o hind ma gin, usually wi h ocelli (e.g. Ha ch 1957; Moo e and
Legne 1979; New on e al. 2000; Leschen and Beu el 2004; Thaye 2016), educed in
some axa (see below); apical maxilla y palpome e as wide as penul ima e segmen , bu
in Co yphiini and some axa o An hophagini and Omaliini i can be dis inc ly na ow-
e ; an ennae a ached unde la e al ma gins o ons, ili o m, monili o m o cla a e;
coxal ca i ies usually open; pos coxal p ocess well de eloped (wi h some excep ions;
see New on e al. 2000); p ocoxae conical and p ominen ; epis omal su u e absen ,
pos e io ace o me acoxa e ical (New on and Thaye 1995; Thaye 2016); a sal
o mula 5-5-5 (4-4-4 in Co neolabiini); abdomen wi h six isible s e ni es; abdominal
e gi es h ee o se en, usually wi h one pai o pa a e gi es; in e segmen al memb anes
a ached apically and wi h b ick-wall s uc u es (e.g. Hammond 1971).
T ibe An hophagini Thomson, 1859
Genus Geod omicus Red enbache , 1857
Type species. S aphylinus plagia us Fab icius, 1798
†Geod omicus bal icus Sha in & Yamamo o, sp. no .
h p://zoobank.o g/16755333-4CBD-4485-B51A-6B64459253B3
Figu es 1, 2, 17–20
Type ma e ials examined. Holo ype: emale, FMNHINS-3965993, comple e speci-
men as inclusion in a piece o ligh yellow Bal ic ambe , 3.4 cm × 2.4 cm × 0.5 cm in
size (Figs 1, 2), wi h glued small pape on plas ic en elope labeled “6083”, wi h h ee
colou pho og aphs o habi us o he bee le ( wo o do sal and one o en al iew)
wi h ec angula s amp on he back o each labeled “Ce i ica e 6083 [handw i en
in blue] | Na u al Bal ic Ambe wi h Inclusions | expe Jonas Damzen | In e na-
ional Ambe Associa ion | Names o Inclusions: | S aphylinidae [handw i en in
blue] | Ro e bee le [handw i en in blue]” <wi h addi ional ound s amp on he le
Omaliine o e bee les in Eocene Bal ic ambe 41
Figu es 1–16. Ambe specimens wi h inclusions o Omaliinae: 1, 2 Geod omicus bal icus sp. no . 3, 4 Eus-
phale um kan i sp. no . 5, 6 Pa aphloeos iba mo osa sp. no . 7–9 Phyllod epa daedali sp.no . 10, 11 Ph.ica i
sp. no . 12, 13 Eusphale um sp. 2 (sp2), Eu. sp. 3 (sp3) and Eu. sp.4 (sp4) 14–16Eu.sp. 1 (specimens 1 o 9
(in he ex : no. 1 o no. 9). Abb e ia ions: hl = holo ype, p = pa a ype, sn = syninclusion. Scale ba s: 1.0cm
(1–6, 10–16), 0.5 cm (7–9).
side: “+SOCIETAS SVCCINORVM+INTERNATIONALIS”]>, wi h he ollowing
labels: “Bal ic ambe | Yan a ny, Kalining ad | Russia | (S. Yamamo o Coll.) | ?Geo-
d omicus | Omaliinae, An hophagini | P o obiae wi h minu e | hai s | a si wi h long
hai s” < ec angula label; handw i en on bo h sides o he label>, “HOLOTYPE |
Geod omicus | bal icus sp. no . | Sha in A. & Yamamo o S. des. 2019” < ed ec an-
gula label, p in ed> (FMNH).
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
42
P ese a ion. The specimen is poo ly isible because i is pa ially co e ed wi h
whi e cloud o mic obubbles c ea ed by decay p oduc s in e ac ing wi h esin, a cha -
ac e is ic o au hen ic Bal ic ambe (Cai and Huang 2013). This is especially no iceable
on he an e io hal o he body, unde he apical and basal po ions o he head, in-
cluding he usual loca ion o ocelli, and mos o he p ono um. The abdominal e gi es
a e no isible do sally, as hey a e co e ed by he hind wings. The en al side o he
specimen is isible in de ail excep o he basal po ion o he ho acic scle i es.
Locali y and ho izon. Bal ic ambe om Yan a ny, Kalining ad, wes e nmos
Russia; mid-Eocene (ca 44 Ma; Wapple 2005).
Desc ip ion. Measu emen s: HW ( en al): 0.76; HL ( en al): ~0.40; OL ( en-
al): 0.25; PLL×PLW (II, III): II: 0.05 × 0.03, III: 0.08 × 0.02; PML × PMW (III, IV):
III: 0.10 × 0.06, IV: 0.16 × 0.05; PL ( en al): ~0.47; PW ( en al): ~0.87; ESL: 1.40;
Figu es 17, 18. Habi us pho og aphs o Geod omicus bal icus sp. no . 17 Do sal iew 18 Ven al iew.
Scale ba s: 1.0 mm.
Omaliine o e bee les in Eocene Bal ic ambe 43
EW: 1.51; MTbL: 1.00; MT L: 0.36 (I–IV: 0.20; V: 0.16); AW (IV): 1.41; TL: 3.80
(head o specimen sligh ly ou o p ono um, hus he o al leng h likely o be sligh ly
sho e ). An ennome es wi h leng hs × wid hs: 1: ? × 0.07; 2: 0.16 × 0.06; 3: 0.11 × 0.06;
4–5: 0.15 × 0.05; 6–7: 0.15 × 0.07; 8: 0.14 × 0.07; 9–10: 0.12 × 0.07; 11: 0.25 × 0.07.
Body elonga e; o ebody con ex. Specimen da k-b own and glossy, wi h an enno-
me es b own, mou hpa s eddish-b own, legs yellow-b own wi h a somewha da k-
ened ibia. Habi us as in Figu es 17–20.
Head ans e se, sligh ly ele a ed in middle, abou wice as wide as long, wi h
sho emples, mode a ely s ongly na owing owa d neck, wi h diagonal mode a ely
deep g oo es ( isible only apical pa o le g oo e), eaching le el o apical hi d o
eye; gula su u es sligh ly sepa a ed a na owes poin on le el o basal hi d o leng h
o eyes (Fig. 20). Eyes la ge and widely con ex, wi h medium-sized ace s. Medio-
Figu es 19, 20. Habi us d awings o Geod omicus bal icus sp. no . 19 Do sal iew 20 en al iew. Ab-
b e ia ions: a1–a11 = an ennome es 1–11; alp = apical labial palpome e; gc = gonocoxi e; g = g oo e; gs =
gula su u e; hw = hind wing; ip = in e coxal p ocess; lb = lab um; lp = labial palpi; md = mandibles; mp =
maxilla palpus; m 1, m 5 = me a a some es; nk = neck; s3–s8 = s e ni es 3–8; s = s ylus. Scale ba : 1.0 mm.
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
50
Figu es 24–29. Eusphale um kan i sp. no . (holo ype: 24–26, 29 pa a ype: 27, 28) 24 habi us, do sal
iew 25 habi us, en al iew 26 ho acic scle i es and legs, en al iew 27 apical pa o ely a, abdo-
men and hind legs, pos e odo sal iew 28 head and an ennae, do sola e al iew 29 head, an ennae and
o elegs, do sal iew. Abb e ia ions: a1–a11 = an ennome es; amp = apical maxilla y palpome e; gc =
gonocoxi e; gs = gula su u e; ip = in e coxal p ocess; nk = neck; m = me a emu ; m = me a ibia; m =
me a en i e; oc = ocellus; p = p o ibia; p 1, p 5 = p o a some es 1 and 5; s3–s8 = s e ni es III–VIII; sc
= scu ellum; s = s ylus; 4– 8 = e gi es IV–VIII. Scale ba s: 1.0 mm (24, 25), 0.3 mm (26–29).
Abdomen dis inc ly na owe han ely a (Figs 27, 40); apical ma gin o e gi e
VII wi h indis inc b ick-wall sculp u e; abdominal e gi es wi h spa se small punc-
u es and no isible mic osculp u e (Fig. 27); s e ni es VII and VIII o bo h males and
emales wi hou modi ica ions (Fig. 44).
Male. Ely a as in Figu e 24; apical ma gin o ely a widely ounded (Figs 30, 31).
Apical ma gin o abdominal e gi e VIII somewha s aigh . Apical ma gin o abdomi-
nal s e ni e VIII widely ounded (Fig. 27).
Omaliine o e bee les in Eocene Bal ic ambe 51
Figu es 30–33. Eusphale um kan i sp. no . (holo ype, male: 30, 31 pa a ype, emale: 32, 33): 30,
32 ely a (schema ic d awings) 31, 33 apical pa o ely a. Scale ba s: 0.3 mm.
Female. Ely a as in Figu e 21; apical ma gin o ely a dis inc ly p olonged a su-
u al apex (Figs 32, 33). Apical ma gin o abdominal e gi e VIII and s e ni e VIII
(Figs 27, 40) s aigh . Geni al segmen wi h ma kedly elonga e gonocoxi es and e y
small s yli (Figs 27, 40).
E ymology. Pa onymic, he species is named in honou o he g ea Ge man philoso-
phe Immanuel Kan (1724–1804), he au ho o he doc ine o anscenden al idealism.
Rema ks. The pa a ype o Eu. kan i sp. no . was isualised h ee-dimensionally
using a mic o-CT scan. Al hough he esul was no e y sa is ac o y, we could obse e
he ossil om mul iple addi ional angles (Figs 34, 44). Based on his scan, we could
desc ibe mo e cha ac e s ha we e no isible wi h ligh mic oscopy. The ossil was as-
signed o he ibe Eusphale ini and genus Eusphale um based on he gene al shape o
he body, shapes and leng h o sho and sligh ly widened a some es 1–4, wi h dense
and elonga e en al se ae, oge he abou as long as apical a some e, and shape o he
ely a o emale sligh ly longe han ha o male, wi h p olonged po ion a su u al
apex (Figs 32, 33). This lo icolous genus con ains 260 alid species (Zane i 2014)
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
52
Figu es 34–44. Eusphale um kan i sp. no ., pa a ype, econs uc ions om x- ay mic o-compu ed omog-
aphy (μ-CT) 34 habi us, do sal iew 35 habi us, en al iew 36 habi us, la e al iew 37 o ebody, do sal
iew 38 head and p o ho ax, en al iew 39 head and p ono um, an e odo sal iew 40 ely a and abdo-
men, pos e odo sal iew 41 p e o ho acic scle i es, en al iew 42 head, la e al iew 43 neck and an e io
po ion o p ono um, do sal iew 44 abdomen, la e o en al iew. Copy igh 2015 Shimadzu Co po a ion.
Omaliine o e bee les in Eocene Bal ic ambe 53
dis ibu ed in he Hola c ic Region. Ea lie , he genus was subdi ided in o wo sub-
gene a: Eusphale um and Pa eusphale um Coi ai , 1959 (Zane i 1987), bu because
se e al species o sensu s ic o and Pa eusphale um we e o e lapping in some mo pho-
logical cha ac e s, he la e was synonymized wi h he nomino ypical axon (T onque
and Zane i 2008). Based on gene al mo phological ea u es o he aedeagus, emale
accesso y scle i e and, in some cases, shapes o he modi ied apical abdominal s e ni es,
se e al species g oups ha e been e ec ed o many species o he genus (e.g. Zane i
1987, 1993, 2014). Howe e , o da e, his di e se genus emains insu icien ly s udied
globally and is in need o u he phylogene ic e ision because o unclea ela ions
be ween bo h species g oups and he ibe Eusphale ini wi h ela ed Omaliini.
The new species is di icul o compa e wi h ex an species as hey ypically di e
om each o he by he mo phology o he aedeagus and emale geni al s uc u es.
Howe e , based on he shape o he s ongly elonga e and dimo phic ely a, Eu. kan i
sp. no . is like membe s o he ollowing species g oups: No h Ame ican con exum
(Zane i 2014; ou species dis ibu ed in Canada and USA) and wes e n Palaea c ic
amplipenne (Zane i 1993; one species known om Tu key), longipenne (Zane i 1987;
six species dis ibu ed in Middle and Sou h Eu ope), mon i agum (Zane i 1987,
1992, 1993, 2004, 2012a; 10 species dis ibu ed in Cen al and Sou he n Eu ope and
Tu key) and anale (T onque and Zane i 2001; h ee species om he cen al-wes e n
pa o Eu ope). The new species di e s om he con exum g oup by he p esence o
he pos ocula ca ina, by he do sal po ion o head wi hou isible imp essions, by he
shape o he apical a some e sligh ly longe han ha in species o con exum g oup
and by he abdominal s e ni e VII o male wi hou modi ica ions. I di e s om he
amplipenne g oup by i s somewha smalle and da ke body, spa se punc a ion o he
o ebody and shape o me a a sus o male, sligh ly cu ed in Eu. amplipenne (see Zan-
e i 1993: ig. 13). The new species sha es simila leng h o he body and pos ocula
ca ina wi h some species o he longipenne g oup, bu di e s by he da ke body and
longe apical a some es. Based on he da k body, gene al cha ac e s o punc a ion and
mic osculp u e o head and p ono um, Eu. kan i sp. no . is somewha like some spe-
cies o he mon i agum and anale g oups, o example Sou he n Eu opean Eu. scha z-
may i (Koch, 1938), Eu. anale (E ichson, 1840), Eu. b andmay i (Zane i, 1981), and
Eu. coi ai i Nicolas, 1974, bu i di e s by he la ge body (body leng h o membe s o
he mon i agum and anale g oups a ies om 1.50 o 2.50 mm) and mo e ans e se
p ono um. F om all hese g oups, Eu. kan i sp. no . di e s by he absence o dis inc
g oo es in on o he ocelli and elonga e an ennome es 2–4 (Fig. 28).
T ibe Omaliini MacLeay, 1825
Genus Pa aphloeos iba S eel, 1960
Type species. Pa aphloeos iba ma ianicola S eel, 1960.
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
54
†Pa aphloeos iba mo osa Sha in & Yamamo o, sp. no .
h p://zoobank.o g/722E3364-B487-4F94-8BA6-5A38BD4A1C00
Figu es 5, 6, 45–53
Type ma e ials examined. Holo ype ( emale), FMNHINS-3260632, comple e speci-
men as inclusion in a piece o small yellow Bal ic ambe , 15.6 mm × 13.1 mm × 4.0
mm in size (Figs 5, 6), wi h he ollowing labels: “14[p in ed] 11[handw i en]- SYAC
00 [p in ed]06 [handw i en] | Bal ic / Bu mi e | O he : | La a / Adul | Omaliinae
[handw i en] | Kalini[n]g ad [handw i en] | Shûhei Yamamo o’s | Ambe Collec ion”
<la ge ec angula label, p in ed>, “Kalining ad, RUSSIA | Shûhei Yamamo o’s | Am-
be Collec ion | (SYAC0006)” <small ec angula label, p in ed>, “[FMNH ba code a
le side o label] FMNHINS | 3260632 | AMBER [handw i en] | FIELD MUSEUM
| We ” <small ec angula label, p in ed>, “HOLOTYPE | Pa aphloeos iba | mo osa sp.
no . | Sha in A. & Yamamo o S. des. 2019” < ed ec angula label, p in ed> (FMNH).
P ese a ion. The specimen is loca ed a an angle wi h he head somewha deepe
in he ambe piece (Figs 5, 6); he specimen is clea ly isible om bo h do sal and en-
al sides. Syninclusions: ound and elonga e ai bubbles nea he specimen on om
he en al side o i s body, and elonga e piece o plan ma e ial loca ed close o he
do sal su ace o ambe piece nea he bee le.
Locali y and ho izon. Bal ic ambe om Yan a ny, Kalining ad, wes e nmos
Russia; mid-Eocene (ca 44 Ma; Wapple 2005).
Desc ip ion. Measu emen s: HW: 0.36; HL: 0.29; OL: 0.17; AL: 0.51; PML ×
PMW (III, IV): III: 0.03 × 0.03, IV: 0.06 × 0.02; PL: 0.31; PW: 0.74; ESL: 0.52; EW:
0.77; MTbL: 0.38; MT L: 0.15 (I–IV: 0.07; V: 0.08); AW: 0.75; TL: ~1.80. An en-
nome es wi h leng hs × wid hs: 1: 0.07 × 0.03; 2: 0.05 × 0.02; 3: 0.05 × 0.01; 4: 0.03 ×
0.02; 5: 0.04 × 0.02; 6–7: 0.03 × 0.03; 8: 0.03 × 0.04; 9–10: 0.04 × 0.05; 11: 0.10 × 0.05.
Body mode a ely wide, glossy (Fig. 45), black, wi h mou hpa s, emo a, and api-
cal pa s o abdominal e gi es eddish-b own, and a si yellow-b own. Body la e ally
as in Figu es 47 and 48. Body wi hou isible mic osculp u e and se a ion excep o
pa a e gi es and abdominal e gi e VIII wi h long e ec se ae (Fig. 48).
Head 1.2 imes as wide as long, wi h sligh ly con ex pos e io po ion, dense and
small punc a ion and pos ocula ca ina (Fig. 49). Head la e ally as in Figu e 49, an e -
o en ally as in Figu e 50, and en ally as in Figu e 51. Eyes la ge, wi h medium-sized
ace s, b oadly con ex, wi h dis inc in ao bi al ca ina (Figs 49–51). Ocelli mode a ely
la ge, si ua ed a le el o pos e io hi d o eyes, dis ance be ween ocelli abou wice as
long as dis ance be ween ocellus and medial ma gin o eye (Fig. 45). Lab um wi h wide-
ly ounded apical ma gin (Figs 46, 51). Apical maxilla y palpome e dis inc ly longe
and sligh ly na owe han penul ima e, swollen in middle and elonga e, om middle
g adually na owing owa d ounded apex (Figs 49–51). Submen um la ge, apezoidal;
apical labial palpome e elonga e, om middle na owing apicad (Fig. 49). Gula su u es
wi h ma kedly ounded pos e io pa s widely sepa a ed om each o he below le el o
pos e io ma gins o eyes (Figs 50, 51). Gena wi h ugose isodiame ic mic osculp u e
(Figs 50, 51). An enna mode a ely sho , exceeding basal po ion o p ono um, wi h
Omaliine o e bee les in Eocene Bal ic ambe 55
Figu es 45, 46. Habi us o Pa aphloeos iba mo osa sp. no . 45 do sal iew 46 en al iew. Scale ba s: 1.0 mm.
spa se elonga e se a ion; basal an ennome e swollen, mo e han wice as long as wide,
an ennome e 2 elonga e, sligh ly widened apicad, 3 hin, as long as 2, 4, and 5 ma k-
edly widened apicad, 6 and 7 as long as wide, 8–10 sligh ly ans e se, apical an en-
nome e wide, om apical hi d sligh ly na owing owa d ounded apex (Figs 45–49).
P ono um wi h sligh ly con ex su ace, ma kedly ans e se, mo e han wice longe
han b oad, wice wide han head, om middle sligh ly mo e na owed an e ad han
pos e ad, wi h widely ounded an e io and sca cely ounded pos e io angles; apical ma -
gin widely ounded, dis inc ly sho e han somewha conca e pos e io ma gin; pa amed-
ian longi udinal imp essions indis inc , wide and long, occupying mos o middle po -
ion; la e al ma gins na owly ema gina e, wi h indis inc ly conca e la e obasal ma gins;
pos e io angles wi hou dep essions (Fig. 45). Do sal su ace o p ono um wi h mo e o
less egula small and dense punc a ion, dis inc ly dense han in pos e io po ion o head
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
56
Figu es 47, 48. Habi us o Pa aphloeos iba mo osa sp. no ., la e al iew. Abb e ia ion: oc = ocellus. Scale
ba s: 1.0 mm.
(Fig. 45). P os e num wi h widely open p ocoxal issu es, exposing ochan ins, and e y
long in e coxal p ocess, wi h acu e apex eaching apical pa o p ocoxae (Figs 46, 51). Me-
dian pa o meso en i e somewha con ex, wi h e y long acu e in e coxal p ocess, each-
ing mo e han hal way along he leng h o he mesocoxae and mode a ely wide apex o
me a en al p ocess (Fig. 46). Scu ellum la ge and iangula , wi h ounded apex and dense
punc a ion in apical pa (Fig. 45). Me a en i e wi h mode a ely wide me acoxal ca i ies
(Fig. 46). Median pa o p os e num and me a en i e wi h indis inc and spa se small
punc u es; mesanepis e num wi h diagonal mic osculp u e; median po ions o p os e -
num and me a en i e, including in e coxal p ocesses, wi h ans e se meshes (Fig. 46).
Ely a e iden ly la ened, 1.4 imes as wide as long, 1.6 imes as long as p ono-
um, wi h mode a ely pa allel la e al sides (Fig. 45), wi h widely ounded apical angles
(Fig. 52), eaching apical ma gin o abdominal e gi e III, wi h apical ma gins sligh ly
oblique owa d su u e (Fig. 45). Punc a ion as ha in p ono um, sligh ly spa se in
basal po ion and nea scu ellum.
Omaliine o e bee les in Eocene Bal ic ambe 57
Figu es 49–53. Pa aphloeos iba mo osa sp. no . 49 head and an enna, la e al iew 50 head, an e o en al
iew 51 head and p o ho ax, en al iew 52 apical pa o ely on and abdominal e gi es IV–V, do sal
iew 53 apex o abdomen and hind legs, en al iew. Abb e ia ions: a1–a11 = an ennome es 1–11; ai =
an ennal inse ion; alp = apical labial palpome e; bws = b ick-wall sculp u e on in e segmen al memb ane;
gs = gula su u e; ip = in e coxal p ocess; m m = men um; m = me a ibia; s7–s8 = s e ni es VII–VIII;
4– 6 = e gi es IV–VI. Scale ba s: 0.2 mm.
Legs mode a ely long and slende , wi h wide emo a and slende ibiae, g adually
widened apicad, co e ed by elonga e se ae on bo h inne and ou e ma gins and wi h
a ew s ong se ae on ou e ma gins (Figs 46–48); a si sho , wi h small se ae on a -
some es 1–4, apical me a a some e sligh ly longe han p eceding a some es oge he ;
a sal claws simple, widely cu ed and elonga e (Figs 46, 53).
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
58
Abdomen con ex, sligh ly na owe han ely a, wi h wide b ick-wall sculp u e on
in e segmen al memb anes be ween e gi es III–VI (Fig. 52) and s e ni es III–VI (Fig.
45). Abdominal e gi es wi h mode a ely dense and deep small punc a ion and dis inc
ne -like mic osculp u e (Fig. 52); abdominal s e ni es wi h indis inc spa se punc a-
ion, wi h shallow isodiame ic mic osculp u e (Figs 45, 52).
Male unknown.
Female. Apical ma gin o abdominal e gi e VIII ounded. Apical ma gin o ab-
dominal s e ni e VIII b oadly conca e (Fig. 53). Geni al segmen wi h mode a ely
wide apical po ions o gonocoxi es; shape and leng h o s yli in isible, each wi h e y
long se a (Figs 47, 48).
E ymology. The speci ic epi he is he La in adjec i e mo osus, -a, -um (s ange). I
e e s o somewha b oad body wi h ma kedly ans e se p ono um o he new species.
Rema ks. Based on he shape o body and maxilla y palpome es (see also Zan-
e i 2012: ig. 55l), sligh ly con ex p ono um, punc a ion and mic osculp u e o he
su ace o body, he ossil p esumably belongs o he genus Pa aphloeos iba. The genus
was e ec ed by S eel (1960a) and was compa ed wi h Phloeos iba Thomson, 1858 and
Phloeonomus Hee , 1839. I di e s om Phloeos iba by he shape o sho maxilla y
palpome e 3 and ela i ely elonga e apical palpome e, and om Phloeonomus by a di -
e en shape o ligula and maxilla y palp ( o de ails see S eel 1960a). Pa aphloeos iba
includes mo e han 30 species dis ibu ed in he Palaea c ic, Madagascan, Nea c ic,
and p edominan ly in O ien al, Aus alian, and Oceanic egions (S eel 1960a; He -
man 2001; Sha in and Sme ana 2016; Sha in 2017b); one species, P. gayandahense
(W.J. MacLeay, 1873) is widely ad en i e a ound he wo ld o New Zealand, se e al
coun ies o Eu ope, and he USA (He man 2001). The new species is di icul o
compa e eliably wi h known species as hese mos ly di e by he s uc u e o he ae-
deagus, and shapes o accesso y scle i e and spe ma heca. The apical an ennome es o
P. mo osa sp. no . a e sligh ly ans e se, beginning wi h an ennome e 8 (Figs 48, 49)
while o he known species ha e ans e se an ennome es beginning wi h 6 o 7. Based
on he punc a ion and mic osculp u e o he p ono um and shape o an ennome e
10 (Fig. 49), he new species is simila o P. specula is (Be nhaue , 1915), known om
New B i ain (Bisma ck A chipelago o Papua New Guinea) bu di e s by he some-
wha la ge and wide body, he absence o la e obasal p ono al dep essions, and he
mo e ans e se p ono um (Fig. 45). Based on he shape and colo a ion o he body,
simila punc a ion, p ono um wi hou dep essions on basal po ions, and somewha
con ex meso en i e, P. mo osa sp. no . is also simila o P. elec ica Zane i e al., 2016,
ecen ly desc ibed om Bal ic ambe , om which i di e s by he wide body wi h
mo e ans e se p ono um, as well as elonga e an ennome es 4–7, and wide apical and
penul ima e palpome es.
Pa aphloeos iba equi es e ision due o unclea mo phological bounda ies be ween
desc ibed species and ela ed gene a, as well as many undesc ibed species om he O i-
en al and Aus alian egions deposi ed in ins i u ional and p i a e collec ions. The new
species is en a i ely a ibu ed o his genus, making i he second ex inc ep esen a-
i e o he genus a e P. elec ica.
Omaliine o e bee les in Eocene Bal ic ambe 59
Genus Phyllod epa Thomson, 1859
Type species. S aphylinus lo alis Paykull, 1789
†Phyllod epa daedali Sha in & Yamamo o, sp. no .
h p://zoobank.o g/BDE372B6-F773-433D-B98E-C0E272FB8FA7
Figu es 7–9, 54–64
Type ma e ials examined. Holo ype (male), FMNHINS-3260629, comple e specimen
as inclusion in e y small piece o ligh yellow Bal ic ambe , 9.3 mm × 5.9 mm × 2.9
mm in size (Figs 7–9), wi h he ollowing labels: “16 [p in ed] 02 [handw i en] SYAC
0 [p in ed] 294 [handw i en] | Bal ic / Bu mi e | O he : | La a / Adul | Omalium sp.
[handw i en] | Omaliinae [handw i en] | Yan a ny, Kalining ad [handw i en] | Shûhei
Yamamo o’s | Ambe Collec ion” <la ge ec angula label, p in ed>, “[FMNH ba code
a le side o label] FMNHINS | 3260629 | AMBER [handw i en] | FIELD MUSE-
UM | We ” <small ec angula label, p in ed>, “HOLOTYPE | Phyllod epa | daedali sp.
no . | Sha in A. & Yamamo o S. des. 2019” < ed ec angula label, p in ed> (FMNH).
P ese a ion. The specimen is ela i ely well p ese ed and many de ails a e is-
ible, om he do sal, en al and la e al sides (Figs 7–9). Howe e , mos body pa s,
excep he do sal su ace o he head, a e co e ed wi h cloud o milky subs ance, espe-
cially mos o he en al side.
Locali y and ho izon. Bal ic ambe om Yan a ny, Kalining ad, wes e nmos
Russia; mid-Eocene (ca 44 Ma; Wapple 2005).
Desc ip ion. Measu emen s: HW: 0.32; HL: 0.22; OL: 0.11; AL: 0.52; PML ×
PMW (III, IV): III: 0.02 × 0.02, IV: 0.07 × 0.02; PL: 0.35; PW: 0.48; ESL: 0.56; EW:
0.51; MTbL: 0.31; MT L: 0.20 (I–IV: 0.08; V: 0.12); AW: 0.50; TL: ~1.80. An enno-
me es wi h leng hs × wid hs: 1: 0.08 × 0.03; 2: 0.06 × 0.02; 3–4: 0.05 × 0.02; 5: 0.04
× 0.02; 6: 0.04 × 0.03; 7: 0.03 × 0.03; 8–10: 0.03 × 0.04; 11: 0.08 × 0.04.
Body elonga e and sligh ly con ex, glossy (Fig. 54), eddish-b own, wi h da ke
head and abdomen; mou hpa s, an ennae, legs and apical ma gins o abdominal scle-
i es yellow-b own. Body la e o en ally as in Figu e 55 and la e ally as in Figu e 56.
La e al ma gins o p ono um (Figs 54, 57), pa a e gi es and abdominal e gi e VIII
(Fig. 64) wi h se e al long e ec se ae.
Head 1.4 imes as wide as long, wi h sligh ly con ex median po ion and sligh o al
la e oapical imp essions (Fig. 57), wi h spa se, small and mode a ely deep punc a ion,
wi h shallow pos ocula ca ina. Eyes la ge and b oadly con ex (Figs 55–57, 60). Ocelli
la ge and con ex, si ua ed a le el o pos e io ma gins o eyes, dis ance be ween ocelli
much mo e han wice as long as dis ance be ween ocellus and pos e io ma gin o eye;
g oo es in on o ocelli p esen , mode a ely deep and sho (Fig. 57). Apical segmen
o maxilla y palp signi ican ly longe han small penul ima e segmen , om swollen
middle g adually na owing apicad (Figs 56, 57, 63). An enna mode a ely sho , jus
su passing basal ma gin o p ono um, wi h spa se e y long se ae on an ennome es
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
66
Figu es 67–72. Phyllod epa ica i sp. no . 67 head and p ono um, an e odo sal iew 68 le an enna,
do sal iew 69 hind legs and abdomen, en al iew 70 le ely on, do sal iew 71 apex o abdomen,
do sal iew 72 apex o abdomen, en al iew. Abb e ia ions: a1–a11 = an ennome es 1–11; bws = b ick-
wall sculp u e on in e segmen al memb anes; gc = gonocoxi e; lb = lab um; lp = labial palpi; mp2–mp4 =
maxilla y palpome es 2–4; m 5 = me a a some e 5; m = me a emu ; m = me a ibia; m c = me acoxa;
m = me a en i e; nk = neck; oc = ocellus; s3–s6 = s e ni es III–VI; sc = scu ellum; s = s ylus; 6– 8 =
e gi es VI–VIII. Scale ba s: 0.2 mm.
ow impunc a e longi udinal a ea (Figs 65, 67). Scu ellum la ge, wi h iangula apex,
wi hou punc u es o mic osculp u e (Fig. 65).
Ely a 1.2 imes as long as wide, eaching apical ma gin o abdominal e gi e III,
sligh ly widened apicad, wi h widely ounded apicola e al angles and apical ma gins
unca e a su u e (Figs 65, 70). Punc a ion dense , ma kedly la ge and deepe han
ha on p ono um, smalle in basal and apical, and spa se in la e al po ions. Su ace
be ween punc u es wi h dense isodiame ic mic osculp u e.
Omaliine o e bee les in Eocene Bal ic ambe 67
Ta si long, wi h apical a some e ma kedly longe han p e ious a some es o-
ge he (Figs 66, 69).
Abdomen sligh ly con ex, as wide as ely a o sligh ly wide , in e segmen al mem-
b anes be ween e gi es IV–VII wi h b ick-wall sculp u e (Fig. 65). Abdominal e gi es
wi h indis inc small and e y spa se punc a ion and mic osculp u e, and wi h spa se
and sho pubescence, wing- olding pa ches no isible.
Male unknown.
Female. Fi s ou meso a some es 1–4 wi hou modi ied se ae (Figs 66, 69). Apical
ma gin o abdominal e gi e VIII sligh ly ounded (Fig. 71). Apical ma gin o abdomi-
nal s e ni e VIII widely conca e (Fig. 72). Geni al segmen wi h ma kedly elonga e
and wide gonocoxi es, wi h e y small na ow s yli, each wi h e y long se a (Fig. 72).
E ymology. The speci ic epi he is he La inized name o Ica us (Ika os), son o
Deaedalus in G eek my hology.
Rema ks. Despi e he shape o an ennome e 3 and he pos e io angles o he p o-
no um, which a e usual in membe s o he genus Ac olocha Thomson, in o he ex e nal
cha ac e s (see de ails abo e), he new species belongs o he genus Phyllod epa. The
ossil sha es wi h ha genus sligh ly p o uded an e io angles o he p ono um wi h
imp essed la e obasal po ions (Fig. 67), simila o hose o ex an No h Eu opean Ph.
sahlbe gi Luze, in addi ion o simila colo a ion o he body and p opo ions o an en-
nome es 4 and 5 (Fig. 68). Howe e , he ossil di e s om ha species by he signi i-
can ly smalle and sligh ly mo e con ex body (Fig. 65), wi h mo e p ominen eyes (Fig.
67), coa se punc a ion o he ely a and p ono um, p ono um wi h less sinua e la e al
ma gins (Figs 65, 67), less ans e se an ennome es 6–10 (Fig. 68), and longe ely a
(Fig. 70). Based on he shape o i s small and pale body, and la ge and deep punc a-
ion o he ely a (Figs 65, 70), Ph. ica i sp. no . is simila o Ph. daedali sp. no ., om
which i di e s by he pale abdomen, na ow apical maxilla y palpome e (Fig. 67), he
shape o an e io angles o he p ono um p o uding apicad, spa se punc a ion o he
p ono um, dis inc ly ans e se head and p ono um (Fig. 67), sho e an ennome es 4,
5, and 11 (Fig. 58), and sho e ely a.
Unnamed species
Eusphale um sp. 1
Figu es 14–16, 73–80
Ma e ials examined. 2 males (no. 6, no. 9), 2 emales (no. 3, no. 7), 5 unsexed specimens
(no. 1, no. 2, no. 4, no. 5, no. 8), FMNHINS-3260631, comple e specimens as inclu-
sions in yellow Bal ic ambe 31.3 mm × 20.6 mm × 12.6 mm in size (Figs 14–16), wi h
he ollowing labels: “15[p in ed]03[handw i en]-SYAC 00[p in ed]8[handw i en]
| Bal ic / Bu mi e | O he : | La a / Adul | Omaliinae 11 exs, [handw i en] | Bal ic
Sea Coas [handw i en] | Axel (Ge many) [handw i en] | Shûhei Yamamo o’s | Am-
be Collec ion” <la ge ec angula label, p in ed>, “Axel Niggeloh” < ec angula label,
p in ed>, “[FMNH ba code a le side o label] FMNHINS | 3260631 | AMBER
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
68
Figu es 73–75. Habi us o Eusphale um sp.1: 73 o ebody, do sal (specimen no. 5) and la e al (speci-
men no. 6) iew 74 p ono um and ely a o specimens no. 1 and no. 2, do sal iew 75 body, la e al iew
(specimen no. 6). Scale ba s: 1.0 mm.
[handw i en] | FIELD MUSEUM | We ” <small ec angula label, p in ed>, “Euspha-
le um | sp. 1 | Sha in A.V. de . 2018” < ec angula label, p in ed> (FMNH).
P ese a ion. The specimens a e isible om one su ace o he piece o ambe
(specimens we e numbe ed as in Figs 14–16). One o he bes p ese ed specimens
(male, no. 6) is loca ed do sola e ally: he su ace o he body, le an enna, and pa
o he abdomen in la e al iew a e clea ly isible. Eigh o he specimens a e p esen
wi h di e ing deg ees o isible de ails. A male (no. 9) is loca ed deep in he piece
o ambe a he le el o i s median con exi y; i s p ono um, ely a, and abdomen
a e clea ly isible do sola e ally, and he an ennae, mou hpa s, legs and pa s o he
ho acic segmen s and abdomen a e isible en ally. A emale (no. 3) is la e ally
o ien ed close o he ou e su ace o he ambe piece; i s basal an ennome es o he
Omaliine o e bee les in Eocene Bal ic ambe 69
igh an enna a e pa ly isible, and he mou hpa s, including maxilla y and labial
palpi, la e al side o he ely a, and pa ly ho acic scle i es, and legs ela i ely a e
isible. Ano he emale (no. 7) is loca ed la e ally deep in he piece o ambe and,
he e o e, he do sal side o i s body is isible bu s ongly cloudy and dis o ed; de-
ails o he s uc u e o maxilla y palpus, igh an enna, i e legs pa ly, some de ails
o he ho ax and abdomen, including apical po ion wi h he geni al segmen a e
mo e o less isible om la e al iew. An unsexed specimen (no. 1) is loca ed do -
sally nea he ou e su ace o he piece o ambe ; only he p ono um and ely a a e
pa ly isible. Ano he unsexed specimen (no. 2) is loca ed do sally nea he ou e
su ace o he piece o ambe nea specimen no. 1; he apical segmen s o i s igh
an enna, p ono um, and ely a a e clea ly isible. An unsexed specimen (no. 4) is
loca ed a li le deepe in he piece o ambe , close o specimen no. 3; i s hind wings
a e ex ended and co e he en i e abdomen. The body is no clea ly isible excep
o he p ono um and ely a. An unsexed specimen (no. 5) is loca ed do sally nea
specimen no. 6, wi h he le an enna, pos e io po ion o head, p ono um, and
ely a pa ially isible. Ano he unsexed specimen (no. 8) is loca ed deep in he piece
o ambe , nea specimen no. 9, and has i s hind wings ex ended so as o co e he
do sal side o he abdomen; he e is a milky co e ing on he la e al side o he body,
and, he e o e, he middle and hind legs a e only pa ly isible in la e al iew and he
p ono um and ely a a e pa ly isible in do sal iew. Syninclusion nea ou e su ace
o he piece o ambe close o i s ma gin: imago o Dip e a abou 2.80 mm leng h,
including wings (Fig. 14).
Desc ip ion. Measu emen s (n = 9): HW: 0.67 (no. 5); HL: ?; OL: 0.18 (no. 6);
AL (no. 6): 0.74; PML × PMW: ?; PL: 0.41–0.46; PW: 0.87 (no. 5); ESL: 0.83–0.96;
EW: 0.71–0.77; MTbL (no. 8): 0.40; MT L (no. 8): 0.28; AW: 0.68–0.74; TL: ~2.50–
3.20. An ennome es wi h leng hs × wid hs (no. 6): 1: 0.15 × 0.05; 2: 0.08 × 0.04; 3:
0.06 × 0.03; 4–6: 0.05 × 0.03; 7: 0.05 × 0.04; 8: 0.06 × 0.04; 9 0.05 × 0.05; 10: 0.06
× 0.05; 11: 0.08 × 0.05.
Body mode a ely wide, con ex (Figs 73–76). Body la e ally as in Figu es 77 and
79, do sally as in Figs 73, 74, and 76 and do sola e ally as in Figu e 78. The specimens
appea b own o black. Body glab ous, wi hou isible se a ion.
Head ans e se, wi h sligh ly con ex middle po ion, wi hou g oo es in on o
ocelli (Fig. 75); punc a ion o pos e io po ion o head i egula , small, and spa se;
su ace be ween punc u es wi h ela i ely la ge ans e se ine mic osculp u e, dis inc -
ly la ge and coa se on neck. Eyes la ge, widely con ex (Figs 75, 77). Ocelli ela i ely
small, sligh ly con ex (Figs 73, 75). Apical segmen o maxilla y palp elonga e, sligh ly
na owe and dis inc ly longe han penul ima e segmen , om middle g adually na -
owed apicad, wi h mode a ely acu e apex (Fig. 77). An enna eaching basal ma gin
o ely a; basal an ennome e ma kedly wide, an ennome e 2 swollen and sligh ly elon-
ga e, 3 wi h hin basal po ion, sligh ly widened apicad, an ennome es 4–6 sligh ly
sho e han 3, 7 sligh ly wide han 6 and 8, 10 sligh ly ans e se in apical po ion,
apical an ennome e wide han penul ima e segmen , om apical hi d g adually na -
owed apicad (Figs 73, 75, 78, 79).
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
70
Figu es 76–80. Eusphale um sp. 1 76 habi us, do sal iew (specimen no. 4) 77 habi us, la e al iew
(specimen no. 3) 78 habi us, do sola e al iew (specimen no. 8) 79 habi us, la e al iew (specimen no. 9)
80 p ono um and scu ellum, do sal iew (specimen no. 1). Scale ba s: 1.0 mm (76–79), 0.2 mm (80).
P ono um sligh ly con ex and dis inc ly ans e se, abou wice as wide as long,
dis inc ly b oade han head, wides in middle, mo e na owed pos e ad han an e ad;
apical ma gin sligh ly ounded, abou as b oad as pos e io ma gin, an e io (Fig. 75)
and pos e io angles (Fig. 80) widely ounded; la e obasal ma gins sligh ly conca e;
la e al ma gins in middle na owly ma gina e; median disc o p ono um wi h e y
indis inc ans e se and la e obasal po ions wi h indis inc wide imp essions (Figs
73–76, 80). P ono um wi h mo e o less egula small and spa se punc a ion, some-
imes wi h wide impunc a e longi udinal a ea on disc, wi h dis inc and mode a ely
la ge ans e se and diagonal mic osculp u e (Figs 73–76, 80). Scu ellum wi hou is-
ible punc u es, wi h dis inc isodiame ic mic osculp u e (Figs 73, 74, 76, 80).
Omaliine o e bee les in Eocene Bal ic ambe 71
Ely a sligh ly con ex, dis inc ly longe han b oad, wice as long as p ono um,
om middle sligh ly widened apicad, eaching apical ma gin o abdominal e gi e IV,
wi h widely ounded apical angles and s aigh apical ma gin unca ed a su u e (Figs
73–76, 78). Punc a ion ma kedly dense and deepe han ha on p ono um, smalle
on basal and apical ma gins and nea scu ellum; mic osculp u e as ha on p ono um
(Figs 73, 74, 76, 78).
Abdomen sligh ly na owe han ely a, wi h small, mode a ely spa se punc a ion
and ine indis inc mic osculp u e.
Male. Apical ma gin o abdominal e gi e VIII ounded. Apical ma gin o abdomi-
nal s e ni e VIII sligh ly sinua e.
Female. De ails o shapes o apical abdominal segmen no isible.
Rema ks. The p esen unique piece o ambe con ains an in e es ing and a e ag-
g ega ion o omaliine specimens which appa en ly belong o one species. Based on
he shape o he body and o he s uc u es (an ennae, maxilla y palpus), ea u es o
punc a ion and mic osculp u e, e c., he species belongs o Eusphale ini o Omaliini.
Ta si o o e- and middle legs a e pa ly isible in one specimen (Fig. 77); a si o his
specimen has long and indis inc ly dense se ae on la e al po ions o a some es 1–4
ha a e common in species o he genus Eusphale um. Based on he shape o he body
and o he mo phological de ails, and lack o addi ional mo phological da a, we ha e
no ound simila species among ex an ep esen a i es o he genus, so we he e ea
his axon as Eusphale um sp. 1. We did no obse e sexual dimo phism in he shape
o apical po ions o he ely a, which o en occu s in Eusphale um, as was obse ed
o Eu. kan i sp. no . abo e. Fu he mo e, he mo phology o he aedeagus should be
s udied, as species o he genus a e eliably dis inguished by he shapes o he median
lobe o he aedeagus and he pa ame es.
Eusphale um sp. 2.
Figu es 12, 13, 81–85
Ma e ials examined. One male, comple e specimen as an inclusion in a piece o yel-
low Bal ic ambe 35.4 mm × 21.5 mm × 7.5 mm in size (Figs 12, 13), wi h glued e y
small quad a e pape on plas ic bag labeled “AWI | 045”, wi h he ollowing labels:
“AWI-045 | Phyllod epa (?) | 3 spec.” < ec angula handw i en label>, “D . Vi alii
Aleksee 's | Collec ion” < ec angula handw i en label>, “Eusphale um sp. 2 | Sha in
A.V. de . 2018” < ec angula label, p in ed>. The specimen is deposi ed in he p i a e
collec ion o Vi alii I. Aleksee (Kalining ad, Russia), egis e ed as AWI-045.
P ese a ion. The single specimen is a male loca ed close o he ou e su ace o
he piece o ambe , wi h many de ails isible in bo h do sal and en al su aces. The
ely a a e somewha de o med and seem la ened, and he igh ely on is dep essed
in o he ho ax. Addi ionally, he piece o he ambe con ains wo males o Eu. sp. 3
and Eu. sp. 4 (see below), and a syninclusion loca ed nea he na owes side o he
ambe : nymph o small mi e abou 0.50 mm in leng h (Figs 12, 13).
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
72
Figu es 81, 82. Habi us o Eusphale um sp. 2 81 oblique do sal iew 82 do sola e al iew. Scale ba s: 1.0 mm.
Desc ip ion. Measu emen s: HW: 0.53; HL: 0.20; OL: 0.11; AL: 0.69; PML ×
PMW (III, IV): III: 0.03 × 0.01, IV: 0.07 × 0.02; PL: 0.43; PW: 0.67; ESL: 0.85; EW:
0.81; MTbL: 0.36; MT L: 0.27 (I–IV: 0.14; V: 0.13); AW: 0.79; TL: 2.06. An ennome -
es wi h leng hs × wid hs: 1: 0.12 × 0.02; 2: 0.07 × 0.02; 3: 0.06 × 0.02; 4: 0.05 × 0.02;
5–6: 0.05 × 0.03; 7: 0.04 × 0.04; 8: 0.05 × 0.04; 9–10: 0.05 × 0.05; 11: 0.10 × 0.04.
Body elonga e, somewha la ened (Fig. 81), glossy and glab ous, wi hou isible
se a ion. Body appea s da k-b own, wi h basal po ions o p ono um and legs eddish-
b own. Body do sola e ally as in Figu e 82 and en ally as in Figu e 83.
Head s ongly ans e se, dis inc ly mo e han wice as wide as long, wi h sligh ly
con ex middle po ion and pos e io pa s o in ao bi al idges, wi hou isible g oo es
in on o ocelli and pos ocula ca ina (Fig. 84). Head wi h indis inc , small and spa se
punc a ion, wi h mode a ely coa se and la ge isodiame ic mic osculp u e be ween
punc u es becoming mo e ans e se owa d middle pa o neck. Eyes la ge, s ongly
p o uding la e ad. Ocelli small, con ex, si ua ed e y close o in ao bi al idges a
le el o abou middle leng h o eyes, dis ance be ween ocelli abou wice as long as
dis ance be ween ocellus and ma gin o eye (Figs 82, 84). Lab um ans e se. Apical
segmen o maxilla y palp elonga e, sligh ly wide in middle han penul ima e seg-
men , om apical hi d g adually na owed owa d mode a ely acu e apex (Figs 82,
Omaliine o e bee les in Eocene Bal ic ambe 73
Figu es 83–85. Eusphale um sp. 2 83 habi us, en al iew 84 le an enna, head and p ono um, do sal
iew 85 abdomen, do sal iew. Scale ba s: 1.0 mm (83), 0.4 mm (84, 85).
84). Gula su u es no ully isible, widely sepa a ed om each o he (Fig. 83). An enna
mode a ely long, dis inc ly exceeding shoulde s o ely a, wi h elonga e se ae, dis inc ly
longe on an ennome es 6–11; basal an ennome e wide, g adually widened apicad, an-
ennome e 2 simila in wid h, swollen and elonga e, 3 wi h hin basal po ion, sligh ly
widened apicad, 4 sligh ly sho e han 3, 5, and 6 sligh ly wide han 4, 7 sho and
mode a ely ounded, 8–10 sligh ly ans e se, apical an ennome e mo e han wice as
long as b oad, om abou middle s ongly na owed owa d acu e apex (Figs 81–84).
P ono um 1.5 imes as wide as long, sligh ly b oade han head, wides in middle,
ma kedly mo e na owed pos e ad han an e ad; apical ma gin sligh ly and widely ound-
ed, abou as b oad as pos e io ma gin, an e io and pos e io angles widely ounded; la -
e obasal ma gins sligh ly conca ed, wi h e y indis inc small c enula ion; la e al ma gins
na owly explana e; la e al po ions wi h indis inc semio al imp ession abou middle (Figs
81, 84). P ono um wi h somewha egula small and spa se punc a ion and wi h isodiame -
ic g ound sculp u e sligh ly coa se han ha on head (Figs 81, 84). P os e num wi h mod-
e a ely wide and p o uded p os e nal p ocess (Fig. 83). Scu ellum la ge and wide (Fig. 81).
Alexey V. Sha in & Shûhei Yamamo o / ZooKeys 863: 35–83 (2019)
74
Ely a li le longe han wide, abou wice as long as p ono um, g adually widened
apicad, eaching basal o apical ma gins o abdominal e gi e IV, wi h widely ounded
apicola e al angles; shoulde s mode a ely widely ounded; la e al edges na owly ex-
plana e (Fig. 81). Punc a ion o ely a in isible in de ails bu appea s sligh ly dense
and deepe han ha on p ono um.
Legs mode a ely long and slende , emo a ma kedly widened in middle, ibiae mod-
e a ely sho and hin, g adually widened apicad, sligh ly sho e han emo a, co e ed
by elonga e se ae, wi h a ew s ong se ae on apical ma gins nea apex; a some es 1–4
dis inc ly wide, wi h dense and long se ae; apical me a a some e long, ye sligh ly sho e
han leng h o p eceding a some es oge he ; a sal claws simple, elonga e (Figs 81–83).
Abdomen (Fig. 85) sligh ly na owe han ely a; abdominal e gi es wi h spa se
and mode a ely small punc u es, no wing- olding pa ches a e isible.
Male. Apical ma gin o abdominal e gi e VIII sligh ly ounded. Apical ma gin o
abdominal s e ni e VIII sligh ly sinua e.
Female unknown.
Rema ks. As in he p e ious species, his specimen has e y long and mode a ely
dense se ae on la e al po ions o a some es 1–4, dis inc ly de o med body (especially
ely a) and unusually s ongly p o uded eyes.
Eusphale um sp. 3
Figu es 12, 13, 86, 89
Ma e ials examined. One male, an inclusion in he same piece o he Bal ic ambe
ha con ains a specimen o Eu. sp. 2 and Eu. sp. 4, wi h an addi ional label: “Euspha-
le um sp. 3 | Sha in A.V. de . 2018” (p i a e collec ion o Vi aly Aleksee (Kalinin-
g ad, Russia), egis e ed as AWI-045).
P ese a ion. The specimen is loca ed do sola e ally close o he ma gin o he
piece o ambe (Figs 12–13). An ennae, la e al po ion o he p ono um and ely a wi h
de ails o he s uc u e o punc a ion and mic osculp u e, abdomen and legs (pa ly)
a e isible in a do sal iew o he body (Fig. 86); eyes, an ennae, some de ails o ho ax,
legs and abdomen a e ela i ely isible in a la e al iew o he body (Fig. 89).
Rema ks. This specimen is abou 2.30 mm long (Figs 86, 89). I is simila o Eu.
kan i sp. no . and Eu. sp. 4 in he shape o he body, eyes and an ennome es. Because
some mo phological de ails o head, p ono um, ho acic scle i es, and legs, as well as
punc a ion and mic osculp u e, a e no isible, we lea e his specimen unnamed.
Eusphale um sp. 4
Figu es 12, 13, 87–88, 90
Ma e ial examined. One male, as an inclusion in he same piece o he Bal ic ambe
ha con ains Eu. sp. 2 and Eu. sp. 3, wi h an addi ional label: “Eusphale um sp. 4 |
Omaliine o e bee les in Eocene Bal ic ambe 75
Figu es 86–90. Eusphale um spp. (86, 89 Eu. sp. 3; 87, 88, 90 Eu. sp. 4) 86 habi us, do sola e al iew
87 habi us, oblique do sal iew 88 habi us, en al iew 89 habi us, la e al iew 90 abdominal apex,
en al iew. Scale ba s: 1.0 mm (86–89), 0.2 mm (90).
Sha in A.V. de . 2018” (p i a e collec ion o Vi aly Aleksee (Kalining ad, Russia),
egis e ed as AWI-045).
P ese a ion. The specimen is loca ed wi h i s do sal side nea he wides ou e
ma gin o he piece o ambe (Figs 12, 13). I is ela i ely clouded wi h many de ails
no isible bo h do sally (Fig. 87) and en ally (Fig. 88).
Rema ks. This specimen is abou 2.30 mm long (Figs 87, 88). Based on he ela i ely
na ow body and shapes o an ennome es, as well as he punc a ion and mic osculp u e
o he o ebody, i is simila o Eu. sp. 2. Howe e , we conside his specimen belongs o
a di e en species, because he eyes o his specimen a e widely ounded as in Eu. kan i
sp. no . and Eu. sp. 3, and because some de ails o he body such as do sal po ion o he
head and shapes o on and middle a si a e poo ly isible. Apical pa o he abdomen
( en al iew) as in Figu e 90, wi h s e ni e VII dis inc ly ema gina ed medioapically.
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