The blueprint of the Amphiesmenoptera – Tarachoptera, a new order of insects from Burmese amber

17 Pages·2017·25.26 MB·English

Foss.Rec.,20,129–145,2017
www.foss-rec.net/20/129/2017/
doi:10.5194/fr-20-129-2017
©Author(s)2017.CCAttribution3.0License.
The blueprint of the Amphiesmenoptera – Tarachoptera, a new
order of insects from Burmese amber (Insecta, Amphiesmenoptera)
WolframMey1,WilfriedWichard2,PatrickMüller3,andBoWang4,5
1MuseumfürNaturkunde,LeibnizInstituteofEvolutionandBiodiversityResearchattheHumboldtUniversity,
Invalidenstr.43,10115Berlin,Germany
2UniversityofCologne,InstituteofBiology,Gronewaldstr.2,50931Cologne,Germany
3independentresearcher:Friedhofstraße966894Käshofen,Germany
4StateKeyLaboratoryofPalaeobiologyandStratigraphy,NanjingInstituteofGeologyandPalaeontology,ChineseAcademy
ofSciences,Nanjing210008,China
5KeyLaboratoryofZoologicalSystematicsandEvolution,InstituteofZoology,ChineseAcademyofSciences,Beijing
100101,China
Correspondenceto:WolframMey([email protected]),WilfriedWichard([email protected]),PatrickMüller
([email protected]),andBoWang([email protected])
Received:7October2016–Revised:16February2017–Accepted:24February2017–Published:24March2017
Abstract. Based on a total of 14 inclusions from Burmese 1 Introduction
amberthenewinsectorderTarachopteraisestablished.The
family Tarachocelidae previously described from Burmese
amber and then placed in Amphiesmenoptera incertae sedis Since 2009, we have had an enigmatic fossil insect from
isassignedtothisneworder.ThegenusKinitocelisgen.nov. Burmeseamberonourdeskforstudy.Thespecieswasiden-
is established to accommodate three new fossil species: K. tified as being a representative of the Amphiesmenoptera, a
hennigi spec. nov., K. divisinotata spec. nov. and K. brevi- superorderthatcontainsthetwoordersTrichopteraandLep-
costata spec. nov. The new genus differs from Tarachocelis idoptera. The term for the superorder was coined by Hen-
gen.nov.bytheabsenceofandroconialscalesonthewings nig (1969), who simultaneously provided a number of au-
and the loss of Cu2 in the forewings. The species are de- tapomorphiccharactersforthistaxon.Healsopresentedthe
scribedindetailandthecriticalcharactersareillustratedby firstcladogramonthephylogenyoftheAmphiesmenoptera
linedrawingsandphotos.Bothmalesandfemaleswerede- in which he explained the differences between genuine
scribed.Thespeciescanbedistinguishedbytraitsinthewing and false stem-group members (Fig. 7). Kristensen (1984)
venation.TheneworderTarachopteraisplacedinthesuper- added further autapomorphies and constructed a hypothe-
orderAmphiesmenopterabasedonthepresenceofsevenam- sizedgroundplanfortheLepidoptera,whichincluded21am-
phiesmenopteran apomorphies and nine tarachopteran apo- phiesmenopteranautapomorphies.Althoughthefossilspeci-
morphies. Apomorphic characters of Trichoptera and Lep- menwasrelativelywellpreservedwithmanycharactersvis-
idoptera could not be disclosed, which suggests an inde- ible,itcouldbeassignedtoneithertheTrichopteranortothe
pendent origin and evolution from an amphiesmenopteran Lepidoptera. Only one autapomorphy is shared with Lepi-
ancestor which was not the ancestor of the Trichoptera- doptera: the presence of scales on the wings. It is the most
Lepidoptera clade. The species of Tarachoptera are tiny in- conspicuous and remarkable feature of the Lepidoptera and
sectswithawingspanof2.3–4.5mmbuthighlyspecialized wewereinclinedtogroupthefossilspeciesclosertotheLep-
according to their aberrant morphology. Aspects of the pre- idopterathantotheTrichoptera.However,theexternalmor-
sumedlifehistoryoftheadultswerededucedfromsomeof phologyoftheinsectdiffersinnearlyallaspectsfromLepi-
thederivedmorphologicaltraitsthatcouldbeinterpretedas dopteraandweeventuallycametotheconclusionthatitwas
adaptationstoahighlystructuredmicro-environment. besttotreatthisspeciesasamemberofahithertounknown
lineage of the stem group of Trichoptera/Lepidoptera. The
PublishedbyCopernicusPublicationsonbehalfoftheMuseumfürNaturkundeBerlin.
130 W.Meyetal.:TheblueprintoftheAmphiesmenoptera–Tarachoptera
species was recently described as Tarachocelis microlepi- Thefossilswereexaminedunderincidentandtransmitted
dopterellaMeyetal.(2017)andplacedinthesimultaneously light using a stereomicroscope (Leica MZ125). Line draw-
establishedfamilyTarachocelidaeofAmphiesmenopterain- ingswereproducedwithaLeicacameralucidaanddigitally
certaesedis. processedusingAdobePhotoshopCS4.Measurementswere
Shortly after completing the manuscript we received, to made with the ocular micrometer of the stereomicroscope.
oursurprise,additionalinclusionsofthisBurmeseamberin- The fossils will be embedded in rectangular pieces of
sect. This new material was very welcome and enabled us artificial resin to make the inclusions more clearly visible
to continue our study on this remarkable insect group. Dur- andtoensureitsdurability.
ingtheinitialinspectionsitbecameobviousthatthematerial
containedmalesandfemalesofatleastthreespecieswhich Acronymsofdepositories:
belong to Tarachocelidae but were not conspecific with the
recently described Tarachocelis microlepidopterella in Mey PM–collectionofPatrickMüller;
etal.(2017).Thefossilsexhibitedsomenew,previouslyun- WW–collectionofWilfriedWichard;
observable characters and provided the first true indications MfN–MuseumofNaturalHistory,Berlin;
for an estimation of the variability in morphological traits NIGP – Nanjing Institute of Geology and Palaeontology,
in this group. In a reassessment of the systematic position Nanjing.
ofTarachocelidaewerecognizedtheunusualmorphologyas
significantanddistinctivetoconsiderthegroupasanorderof
itsown.Herewedescribethisnewfossilorderofinsectsand 3 Descriptionofnewtaxa
give a discussion of the evidence which led us to establish
thishigher-ranktaxon. Tarachoptera,ordonov.
Wehaveincludedinthepresentarticleareducedversion Typefamily:Tarachocelidaefam.nov.(Burmeseamber)
of the descriptions of Tarachocelidae and Tarachocelis mi-
crolepidopterella gen. nov. spec. nov. in order to avoid the Description (Figs. 1–14): Head flat, dorso-ventrally de-
establishmentsofnominanuda.Thedetaileddescriptionsof pressed, with coronal, frontal and frontogenal sulci present,
thetaxaarepublishedinMeyetal.(2017). one pair of elongate setose areas on vertex and frons, ocelli
absent; antenna filiform, with barrel-like flagellomeres;
anteclypeolabrum separated by horizontal furrow from
2 Materialandmethods
frontoclypeus and kneed in frontal direction; mandibles
vestigial; maxillary palpi very small, three-segmented,
BurmeseamberhasbeenrecordedfromtheShwebo,Thayet-
galea triangular; labial palpi long, porrect or ascending,
myo, Pakokku, Magway, and Pegu districts in Myanmar
three-segmented; cervix enlarged with laterocervicalia and
(Rossetal.,2010).However,theonlycommercialsourceis
additional pair of small cervical sclerites on dorsal side.
the Hukawng Valley in Tanaing Township, Myitkyina Dis-
Thoracal segments inclined frontad, pronotum flat, divided
trictofKachinState.Theamberstudiedhereisfromanam-
bymembranous,transversefurrowintoananteriorandpos-
berminelocatednearNoijeBumVillage,TanaingTownship
terior plate, both connected with lateropleuron small setose
(Kaniaetal.,2015).Thesedepositshavebeendatedindetail
areas on lateral sides of pronotum. Wings homoneurous,
byShietal.(2012).Wetentativelyfollowtheagedetermina-
with three radial veins forming a long radial cell closed
tionasearlyCenomanian(98.8±0.6millionyears)givenby
apicallybycrossveinr1–r2;mediaunbranched,wingsurface
U-Pbdatingofzirconsfromthevolcanoclasticmatrixofthe
covered by small scales, which are spindle-shaped with
amber (Shi et al., 2012), but the geological age of Burmese
acute or slightly rounded tips, hindwing and wing margin
amber should be slightly older than the zircon date (Ross,
scales lanceolate and 2 times longer, but with only three to
2015).
four primary ridges; veins devoid of hairs or scales; large
Thefossilspecimensareembeddedinsmallamberblocks
and erect androconial scales on radial and medial veins
cut out from larger Burmese amber pieces. Some adult in-
of male fore- and hindwing present or absent. Hindwings
sects are nearly completely preserved and often visible in
incompletely covered by scales, smallest scales with a size
ventralanddorsalview.Themalegenitaliaareflattened.The
of28–35µmandmorepigmented(darker)thanotherscales;
hindwingsareoftenpartlycoveredbythesomewhatdistorted
allscalesonthewingsbentandslightlyarched,whichisin
forewings.Thehead,thoraxandabdomenshowoftensigns
contrasttotheflatscalesofLepidoptera;wingmembraneon
ofdecompositionandmaceration.Antennaeandlegsareof-
both wings densely covered by tubercles, giving the wings
tenincomplete.
a granulate appearance; scales present on antennae, head,
PhotosweretakenusingaLeicastereomicroscopeM420
thoraxandlegs;legsslender,tibiaofalllegswithoutspurs,
ApozoomincombinationwithaCanonEOS600D,EOSutil-
pretarsus with simple claws and pulvilli. Abdomen of male
ity software and the Zerene Stacker software or were taken
with sternal processes on segment IV–VI. Male genitalia
withaKeyenceVHX-900Fdigitalmicroscope.
small, retracted into segment IX, and covered dorsally
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W.Meyetal.:TheblueprintoftheAmphiesmenoptera–Tarachoptera 131
by single, long plate; segment IX with comb of spines the upper side of the hindwings. Membrane of forewing
on ventro-caudal margin. Female genitalia with paired or sparsely covered by spindle-shaped and piliform scales
unpaireddorsalplate. without perforations. Humeral vein in the forewing present
andapicalsectionofCu2inforewingabsent.
Diagnosis: The characters 21–29 in Figs. 4–9 discussed
below are interpreted as apomorphies of the new order. Tarachocelismicrolepidopterellaspec.nov.
Seven characters (1, 2, 4, 8, 9, 12, 13) were identified as (urn:lsid:zoobank.org:act:2CADD125-72AA-4F9B-81B7-
amphiesmenopteran apomorphies (see discussion below). 106B980B4C12)
ThreeapomorphiesoftheTrichoptera/Lepidopteraclade(3, (Figs.13–14)
6, 14) are in a plesiomorphic state, with the architecture of
theprothoraxasthemostremarkablecharacter(seeFigs.2, Material: Holotype male, Burmese amber, S. Anderson
9). Coll. (No. 47), Reg. No. G2010.20.36, National Museums
Scotland,Edinburgh.
Tarachocelidaefam.nov.
(urn:lsid:zoobank.org:act:8C5E04FF-2B78-4BD4-A55F- Etymology: The name refers to the general similarity
2480D4BC90C9) oftheinsecttovariousmicrolepidopteranspecies.
Typegenus:Tarachocelisgen.nov., Description: Length of forewing 3mm; length of hind-
wing 2.9mm; head elongate and somewhat flattened
Description: Head elongate, mouthparts slightly hypog- dorso-ventrally, scape and pedicel together as long as eye
nathous, vertex without setal warts, eyes oval, antenna diameter, scaled dorsally, 23 barrel-shaped flagellomeres
filiform, maxillary palpi very short, three-segmented, of present, basal segments (1–8) with scales, subsequent seg-
equal length, ciliated, last segment pointed; labial palpi ments unscaled, each flagellomere with short cilia, evenly
long, three-segmented, all segments of about equal length, distributed over the entire length; spurs and smaller spines
withlonghairs,terminalsegmentwithpointedapex;labrum absent on all tibiae, tarsal segments with terminal pair of
large, produced medially as a quadroangular and volumi- ventral bristles; crossveins cu–r1 and r1–r2+3 present, the
nous organ, not set as a prolongation of the clypeus but latter closing an elongate, triangular, radial cell; crossvein
protruding by nearly 30◦ from the clypeus; pilifers absent, m–cu1apresent;basalpartofCu2fusedwithCu1a+b;anal
pretarsus with slender ungues and stalked arolium, pulvilli veins (A1 and A2) apparently with basal loop. Hindwing
absent; fore- and hindwings of nearly the same length with venationsimilartoforewingvenation.
rounded apices, pterostigma absent, venation homoneurous
concerning branching of R and M veins, three radial and Kinitocelisgen.nov.
a single median vein present; small jugum present, folded (urn:lsid:zoobank.org:act:22FA942B-37FA-4C47-932B-
under base of wing, forewing with Sc a thin vein branched 26519E0393A3)
apically and with humeral (h) and subapical crossvein sc–r;
radiusdividedintotwothickveins,bearingbroad,triangular, Typespecies:K.hennigispec.nov.
putative androconial scales on the underside of the wing, Gender:Female.
upper R vein unbranched (R1), lower R vein with two Etymology: The name is derived from the Greek kinitos,
terminalbranchesR1 andR2(orR2+3andR3+4),ending agile,andcelis,asasuffixofgenericnames.
onwingmarginbeforeandatapexrespectively.
Description: Small adult insects with 2.3–4.5mm wing
Tarachocelisgen.nov. length, and 2.5–4.3mm body length. Wings held above
(urn:lsid:zoobank.org.act:BDE44937-0710-426A-B2A5- abdomeninaflatandoverlappingmode.
C4850CAFFF6A) Head: Elongate, moderately dorso-ventrally depressed,
hypognathous,fronsandvertexwithsemi-erectscales,eyes
Typespecies:T.microlepidopterellaspec.nov. hemispherical,ocelliabsent,antennafiliform,20–26barrel-
Gender:Female. shaped, scaled flagellomeres; head capsule with coronal
Etymology: The name is derived from the Greek ταραχη and frontal sutures, fronto-genal sutures also present, ante-
(tarachi), frightened, staggering, which concerns the pre- rior tentorial pits large, deep furrow between frontoclypeus
sumedflightoftheadults. and clypeolabrum, proximal part of clypeolabrum sclero-
tized, distal part membranous, mandibles vestigial, reduced
Description:Asdescribedforthefamily. to small, conical processes, lying laterally of clypeolabrum
and obviously nonfunctional, galea of maxilla an enlarged,
Diagnosis: Males with presumed androconial scales on nearlytriangular,flatlobewithablade-likeanteriormargin,
R and M veins on the underside of the forewings and on maxillaryandlabialpalpithree-segmentedandscaled,labial
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132 W.Meyetal.:TheblueprintoftheAmphiesmenoptera–Tarachoptera
Figure 1. (1–5) Wing venation. (1) Kinitocelis hennigi spec. nov. Figure 2. (6–8) Kinitocelis divisinotata spec. nov.: (6) head and
(B113); RC – radial cell. (2) Kinitocelis divisinotata spec. nov. pronotum,dorsal;(7)head,latero-frontal;(8)head,ventro-frontal.
(B44).(3)Kinitocelisdivisinotataspec.nov.(B67).(4)Kinitocelis
brevicostataspec.nov.(BUB319).(5)Kinitocelisbrevicostataspec.
nov.(BUB399).
Genitalia: Male – dorsal part (uncus?) an elongate, roof-
like plate, ventral part (vinculum?) with a bulbous, distal
palpi much longer than the former, haustellum or proboscis marginequippedwithatransversecombofabout15–20stiff
absent. spines, valvae short, in vertical position, hardly protruding
Thorax:Thoracalsegmentsinclinedtowardshead,cervix beyondcombofspines.
extensible,laterocervicalscleritefrombroadbaseonepister- Females – dorsal part of final, abdominal segment pro-
num spanning to hind margin of head, pronotum seemingly duceddistadwithapairofovalplatesattachedtothelateral
divided transversally into an anterior and posterior notum, sides, or with a single, apically rounded plate; ventral part
corner of anterior notum connected ventrally with epister- broadly rounded bearing a large, triangular process in the
num; fore- and hindwings translucent, of nearly the same middle,orwithoutprocess,butwithnotchedventralmargin.
length with rounded apices, humeral vein and pterostigma
absent,jugallobetransformedintorod-likeappendageorab- Diagnosis. Kinitocelis gen. nov. can be separated from
sent,venationhomoneurousconcerningbranchingofRand Tarachocelis gen. nov. by characters of wing venation: the
Mveins,threeradialandasingle,unbranchedmediapresent, humeral vein is absent, the radial and median veins are
forewingScwithorwithoutapicalfork(Sc1andSc2),bases without enlarged, androconial scales and the postcubital
of R1+2 and R3 and crossvein r2–r3 forming radial cell vein (Cu2) has lost its free, apical section. An advanced
(RZ);spindle-shapedscalesandpiliformorlanceolatescales characteristhedensescalingontheforewingsincontrastto
onthewingmembrane,thescaleswithoutperforations,some thesparselyscaledwingsofTarachocelis.
scatteredscalesalsoontheventralside,hindwingscalesand
fringes mostly of the piliform type, hindwing without jugal Kinitocelishennigispec.nov.
lobe,Cu1bformingafalsecrossveinbetweenCu1aandA1, (urn:lsid:zoobank.org:act:71B3ADEE-519A-4682-A105-
Cu2 absent; legs slender, epiphysis on foreleg absent, spurs F2180C539195)
andadditionalspinesabsentontibiae,basaltarsomerenearly (Figs.1,4(16–17),8)
aslongasremainingtarsalsegmentstogether,alltarsalseg-
mentswithapicalpairofshortspines. Material:Holotype♀,B113,depositedNIGP164790
Abdomen: Female abdomen without sternal processes,
male sternum of abdominal segment V with pair of short, Etymology: Named in honor of Willi Hennig (1913–
lateralprocesses,segmentsVI–VIIwithsmallprotuberances 1976), in recognition of his ideas on the phylogeny of
inshallowdepressionsonlateralsides. AmpiesmenopterapublishedinHennig(1969).
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W.Meyetal.:TheblueprintoftheAmphiesmenoptera–Tarachoptera 133
Figure 4. (16–22) Female genitalia. (16–17) Kinitocelis hennigi
spec.nov.(B113),lateral(16)andventral(17).(18–19)Kinitocelis
divisinotataspec.nov.,lateral(18)andventral(19).(20)Kinitocelis
brevicostataspec.nov.(BUB399),ventral.(21)Maleabdomenof
K. brevicostata spec. nov., lateral view (B384). (22) Wing scales,
dorsal side of K. brevicostata spec. nov. (BUB 843). Hindwing –
A–B:dorsalside;C:lateralside.Forewing–D–E:dorsalside;F:
lateralside.
Figure3.(9–14)Kinitocelisbrevicostataspec.nov.(BUB319):(9)
Kinitocelisdivisonotataspec.nov.
headandthorax,dorsal;(10)head,fronto-lateral;(11)pterothorax,
(urn:lsid:zoobank.org:act:2DE92E61-C740-414E-8658-
lateral;(12)maleabdomenwithgenitalia,lateral.Kinitocelisbrevi-
costataspec.nov.(BUB399):(13)head,dorsal;(14)head,ventral. 9DB441D79C55)
Kinitocelisdivisinotataspec.nov.(BUB730):(15)head,dorsal. (Figs.1(2–3),2(6–8),3(15),4(18–19),9)
Material:Holotype♀,B44x,depositedinNIGP164789

Paratypes: 1 ♀, BUB 773, deposited in PM; 1♂, BUB
Description (holotype): Length of forewing 2.9mm;
730,depositedinMfN;1♀,B6 7and1♀,B390deposited
length of body 3.0mm. Antenna with 21 flagellomeres; in
inNIGP164785,NIGP165287
forewing (Fig. 1(1)) subcosta apically furcate, right wing
withasinglecostalcrossvein,whichisabsentonleftwing,
Etymology: The specific name refers to the seemingly
distance between sc–r and r1–r2 is 2 times the length of
splitprototumindorsalview.
the latter crossvein, A1 and A2 looping, jugum in form
of a rod-like appendage; hindwings slightly broader than
Description (holotype): Length of forewing 3.5mm;
forewings,crossveinr1–r2veryshort.
length of body 3.5mm. Antenna with 22 flagellomeres; ter-
Female genitalia (Fig. 4(16–17)): Dorsal part of final,
minal segment of labial palpi thickened; pronotum divided
abdominal segment produced distad with a pair of oval
intoanteriorandposteriorplatesbytransversefold,anterior
plates attached to the lateral sides, ventral part broadly
plate connected ventrally with episternum; in forewing
rounded bearing a large, triangular process in the middle,
(Fig. 1(2–3)) subcosta simple, not forked, ending on wing
directeddistad.
margin before or after crossvein r1–r2; distance between
sc–r and r1–r2 variable, 1 to 4 times the length of the latter
Diagnosis: The species differs from all other species of
crossvein,A1andA2looping,scalessparselydistributedon
thegenusbythepresenceofanapicallyforkedsubcostaand
wing surface; hindwings as broad as forewings, crossvein
therod-likejugumoftheforewing.Theabsenceofthecostal
r1–r2aslongassc–r1.
crossvein in one wing qualifies this character as regressive
andoflimitedtaxonomicsignificance.
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134 W.Meyetal.:TheblueprintoftheAmphiesmenoptera–Tarachoptera
Figure5.(23–25)Headinlateralview(withoutflagellum).(23)RhyacophilasibiricaMcLachlan,1879;(24)Micropterixcalthella(Lin-
naeus,1758);(25)Kinitocelisbrevicostataspec.nov.(BUB319).
(Figs.1(4–5),3(9–14),4(20–21),10–12)
Material:Holotype♀,BUB399,depositedinMfN;

Paratypes: 1♂BUB 319 PM, 1♂B 307 WW, 1♂352
NIGP164786,1♀,B371NIGP164787,1♂,B384NIGP
164788,1♂,BUB843PM,deposite dNIGP,WWandPM.
Etymology: The specific name is composed of the Latin
Figure 6. Cladogram of the phylogenetic relationship of Tara-
brevis, short, and costa, vein referring to the atrophied or
choptera, Lepidoptera and Trichoptera (numbers correspond with
thoseforsynapomorphiesandautapomorphiescitedinthetext). weaklysclerotizedapexofsubcostalvein.
Description (holotype): Length of forewing 4.5mm;
length of body 4.3mm. Antenna with 26 flagellomeres, the
Female genitalia (Fig. 4(18–19)): Dorsal part of final paratypeswith22–23flagellomeres;erectpiliformscaleson
abdominal segment a single, elongate, oval plate, ventral fronsarrangedintwolines,rightmandibleasmall,rounded
partbroadlyrounded,upturnedandnotchedinthemiddle. cone sitting in a deep cavity close to the lateral side of
clypeolabrum, segments of labial palpi sparsely scaled; in
Diagnosis: The club-shaped labial palpi, the female forewing(Fig.1(4–5))subcostaforked,bothbranchesclose
genitaliaandthelongapicalsectionofthesubcostabeyond together and weakly sclerotized, distance between sc–r and
crossveinsc–rarediagnosticforthespecies. r1–r2isthelengthofthelattercrossvein,A2andA3looping,
asmallrod-likejugumpresent,scalesdenselydistributedon
Remarks: The wing venation appears to be variable in wing surface; hindwings broad as forewings, only on costal
the length of the radial cell (RZ). The positions of the area with dense scaling, crossvein r1–r2 shorter than sc–r1.
crossveins are altered adequately. In Fig. 1(3) the aberrant Male abdomen with sternites V-VII enlarged dorsally and
wingvenationofB67isdepicted.Itisleftforfuturestudies with a bulbous protuberance displaying a round orifice in a
and more material to decide whether the forewing venation shallowdepression.
isindeedavariablecharacterofK.divisinotataspec.nov.or Female genitalia (Fig. 4(20)): Dorsal part of abdomi-
asecondspeciesisinvolved. nal segment with pair of oval plates, ventral part broadly
rounded,upturnedandnotchedinthemiddle.
Kinitocelisbrevicostataspec.nov.
(urn:lsid:zoobank.org:act:B7D7B85C-630F-456D-B010-
74B5E41DD76D)
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W.Meyetal.:TheblueprintoftheAmphiesmenoptera–Tarachoptera 135
Figure7.CladogramofHennig(1969)onthepresumedphylogeny
ofAmphiesmenoptera,illustratingthetopographicdifferencesbe-
tweengenuineandfalse(=unechte)stemgroups.Thecladogram
stillappliesandthepositionofTarachopteraisinserted,demonstrat-
ingitsearlybranchingfromanamphiesmenopteranancestor.
Additional characters provided by paratype specimens Figure 8. Kinitocelis hennigi spec. nov., holotype♀(B113): (a)
BUB319: dorso-lateralviewand(b)ventralsideoftipoffemaleabdomen.
Male genitalia (Fig. 3(12)): Dorsal part of abdominal seg-
mentIXaflat,elongateplatewithtruncateapex,overarching
the genital cavity; ventral part with a short vinculum, and identified at that time (1984–2002), 19 are adult and 3 are
short vertical processes (? valvae); ventral hind margin of larval characters. Most of them have no relevance in the
segment VIII with a comb of about 20 stiff spines, directed studyofamberfossils,becausetheyconcernembryological,
distad. anatomicalandcytologicalstructuresunobservableininclu-
sions. Characters of the exoskeleton can only be used for
Diagnosis: The reduction of the apical section of the comparisons if well preserved and not covered by wings or
subcosta in both veins is a distinguishing character of the other structures. None of the examined specimens displays
new species. The individuals of this species are slightly the full set of the critical traits. Characters are well visible
largerthantheprecedingspecies.Thereissomevariationin in one specimen but not in the other. The descriptions and
the wing venation among the specimens, which is probably discussions are based always on the individual that shows
due to different embedding positions rendering direct the character in the clearest way. The following characters
comparisons and productions of corresponding illustrations in the sequence and numbering of Kristensen (1984) are
difficulttomake. applicabletoTarachoptera:
1. Prelabium fused with hypopharynx. This composite
4 Discussion structure leads to the formation of the haustellum. The
haustellum of primitive Trichoptera and of the aglossatan
Amphiesmenopterancharacters Lepidoptera families (Micropterigidae, Agathiphagidae,
Heterobathmiidae) has a similar structure (see Kristensen
Kristensen (1984) and Kristensen and Skalski (1998) and Nielsen, 1979). The fossil specimens do not show a
summarizedtheautapomorphiccharactersofthesuperorder clear haustellar lobe which allows a direct recognition of
Amphiesmenoptera. In addition, Ivanov (2002) mentioned this autapomorphy, but in the holotypes of K. divisinotata
one trait (most of the forewing Sc branches lost) from his spec. nov. and K. brevicostata spec. nov. the opening of the
paleontologicalperspective.Thepresenceofwell-developed preoralcavityisvisible,situatedventrallyofthegalealobes
silkglandswasfoundtobeanembryologicalautapomorphy (Figs. 2(7–8), 3(10, 14)). In extant Trichoptera the preoral
(Kobayashi and Ando, 1988). Of the 23 autapomorphies cavity is filled with the eversible haustellum, in which no
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136 W.Meyetal.:TheblueprintoftheAmphiesmenoptera–Tarachoptera
space is left for a large opening or a large preoral cavity The morphology of the pronotum in Tarachoptera differs
that can be seen from frontal view. As a consequence, an considerably from Trichoptera/Lepidoptera and represents
assumed haustellum of Tarachoptera must be a small organ veryprobablytheplesiomorphicstateinAmphiesmenoptera,
too, similar to that of Micropterigidae. The preoral cavity preservedinTarachoptera.
appears to be large enough to provide space for something
like an infrabuccal pouch in the hypopharynx, which plays 4. Pterothoracic episterna with characteristic suture
a role in crushing food particles and which was considered pattern. The paracoxal sulcus and a short, ventrad running
by Kristensen (1984) as plesiomorphic for the Amphies- sulcus depicted by Kristensen (1984) are also present in
menoptera or Panorpida. However, the vestigial mandibles Tarachoptera.
ofTarachopteraareobviouslynotmoveableandthusunable
to interact with an infrabuccal pouch. This function of 5. Secondary furcal arms in pterothorax fused with
the mandibles could have probably been adopted by the posterior margin of corresponding epimera. This autapo-
triangular galeas, which show blade-like ventro-median morphybecomesvisibleonlybylookingatthebacksidesof
margins. If this is correct, the food particles were probably thesesegments.Noneoftheavailablespecimensallowsthis
not pollen grains but something of a softer consistency like observation.
sporangiaoflichensorfungi.
6. Metathorax with a setose sclerite in the wing base mem-
2. Lower posterior corner of laterocervicale produced branebelow/behindthesubalare.Asetosescleriteisabsent
towards the prosternum. The morphology of this cervical in this position, an observation possible only in specimen
sclerite corresponds largely with the sclerites in extant BUB 319 of K. brevicostata. However, the subalare in both
Lepidoptera and Trichoptera. The cervix was observed thoracal segments is not discernable, but the dorsal side of
in two fossil specimens (BUB 730, B 352) as a long, theepimeronisunusuallylargeandslightlyconcave(B113).
eversibleorganwithapairofadditionalcervicalscleritesor
protuberancesonthedorsalside(Fig.3(15)).Thesesclerites 7. Pretarsus above the claws with a pseudempodium, a
are not known to occur in extant species of Trichoptera strong seta on a socket. The pseudempodium is not clearly
and Lepidoptera, but are present in distantly related orders visible. Observing this character is further complicated by
like Homoptera, Zoraptera and Mantodea (Matsuda, 1970). the presence of several setae arising from the dorsal side of
The sclerites as supporting structures probably developed thepretarsus.
togetherwiththeelongatedcervicalmembrane,whichledto
a greater movability of the head. The cervix and its dorsal, 8. Wings with extensive covering of setae. Setae are
cervical sclerites seems to be an adaption to an unknown present on both the fore- and hindwings. They are dis-
foraging behavior and might have evolved independently persed randomly among the more numerous scales. Like
fromotherlineages. in Lepidoptera and some Trichoptera the scales on the
wings of Tarachoptera are modified setae. Their dorsal
3. Pronotum with paired setose warts. In contrast to surfacehas4–8primaryridgeswithoutprovidinganyvisible
Trichoptera and Lepidoptera, which have vertically upright perforations between ridges. They are spindle-shaped (like
standingpronotawiththewartsinthemiddle,thepronotum inNeopseustidae)andoccurindifferentlengths(Fig.4(22)).
ofTarachopteraisaflatplate,withsetosepartsonthelateral The forewing scales are smaller and more dense than their
sidesonly,butwithoutwartsordistinctprotuberances(Figs. hindwing counterparts. The latter are somewhat elongate,
2(6), 3(9, 15)). In a number of Trichoptera families (e.g., withthelongestcomprisingthefringesonthewingtermen.
Philopotamidae, Calocidae) the pronotum carries two pairs The fringes of the anal hindwing margin are comprised of
of setose warts, a condition which is considered a ground- longhairs.
plancharacterofTrichoptera(Ross,1956).Thepronotumin
Tarachopteraisdividedintoanteriorandposteriorplatesby 9. Anal veins of the forewings apparently looping up
amembranousfossaorfold(Fig.2(6)).Theplatescanliein into a double-Y configuration. The anal veins of the
the same plain or are angled on the transverse membrane. forewing are amalgamated, but they differ in the studied
On the lateral sides, the pronotal plates seem to come close specimens (Fig. 1). Apart from a double-Y form, a simple
totheanepisternumandepimeronofthepropleuron.Trans- Y configuration is the dominant pattern, with or without a
versesuturesofthepronotumaredescribedfromMecoptera short,freeapicalpieceofA3,whichrunstothewingmargin.
(Matsuda, 1970). In the family Hepialidae (Lepidoptera)
the pronotum includes an anterior dorsal plate (Nielsen and 10. One ventral neck muscle originating on the fore
Kristensen, 1989) which could be interpreted as a remnant coxa; 11. conical furcopleural muscle in the mesothorax
of a larger and transversely divided pronotom. However, with broad end on the pleural ridge. These anatomical
the median sulcus of this anterior plate is not present in characters are not observable in fossils, but could possibly
Tarachoptera,makinghomologyofbothstructuresdoubtful. observedbyusingmicro-CTscans.
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W.Meyetal.:TheblueprintoftheAmphiesmenoptera–Tarachoptera 137
Table 1. Seven characters are identified as being present, which
12. Presence of paired glands opening on sternum V. clearlysupporttheplacementofTarachopterawithintheAmphies-
Dorsalprocessesonthesternitesoftheabdominalsegments menoptera.Threecharacterswerefoundtorepresenttheplesiomor-
are interpreted as being the location of gland openings. phicstate,andtwofurtheronesremainunclear.Thesymplesiomor-
phiesdeservefurtherstudy,becausetheyimplyanadvancedstatein
In Trichoptera similar sternal processes occur in several
Trichoptera and Lepidoptera, and may provide additional synapo-
primitive families (e.g., Hydrobiosidae, Polycentropodidae)
morphies for supporting the Trichoptera–Lepidoptera sister-group
(IvanovandMelnitsky,2002;Djernaes,2011).However,we
relationship.
cannotdecidefromtheTarachopterafossilswhichsegments
areinvolvedbecauseintersegmentallinesordepressionsare
AutapomorphiesofAmphiesmenoptera
obscuredandadistinctionbetweenmembranesandsclerites
is not possible. Nevertheless, at least two segments have present absent undecided
sternalprocesses,whichpointstothepresenceofmorethan
Characters 1,2,4,8,9,12,13 3,6,14 5,7
one gland pair (Figs. 3(12), 4(21)). Only the males exhibit
thesesternalprocesses,asituationalsofoundinLepidoptera,
Agathiphagidae(DjernaesandKristensen,2011).
13. Male abdominal segment IX with tergum and sternum
fused,formingaclosedring.Thischaracterispresentinthe
examinedmalespecimensofTarachoptera(Fig.3(12)).
14. Anterior margin of female segments VIII and IX
with long, rod-like apodemes, accommodating the insertion
of protractor/retractor muscles of the extensible oviscapt
(= ovipositor). The studied females of Tarachoptera do not
possess papillae anales nor any elongated structures, which
wouldimplytheexistenceofanovipositor(Fig.4(16,18)).
If apodemes are present, they must be small and not of
the long, rod-like shape as in extant Trichoptera and Lep-
idoptera. In Trichoptera vestiges of apodemes are retained
in several families (e.g., Philopotamidae, Phryganeidae,
Apataniidae), which are reason enough to consider the lack
of these apodemes as secondary losses. This explanation
is probably not applicable to the lacking apodemes in Figure9.Kinitocelisdivisinotataspec.nov.,holotype♀(B44):(a)
aglossatan Lepidoptera families (Micropterigidae, Het- dorsalsideofadult,(b)detailofthoraxandhead,and(c)detailof
erobathmiidae) (Kristensen 1998) and in the primitive headandprothorax,ventrolateral.
Hydrobiosidae of the Trichoptera. However, the ovipositor
withlongapodemesonsegmentVIIIandIXisanancestral
characterfoundinfossilspecimensfromtheUpperJurassic ofLepidopterathroughasuiteof26synapomorphies.Most
to Lower Cretaceous and was used to establish the extinct charactersarefoundinadults,andatotalof17concernthe
LepidopterafamilyEolepidopterigidae(Rasnitsyn,1983). exoskeleton.Threeofthesecharacterscanalsobeobserved
inTarachoptera:
Kristensen (1984) discussed a further five characters
from visceral anatomy and cytology, including the female 15. Wings with covering of broad scales. Fore- and
heterogamety as the classical synapomorphy, which are, hindwingsinTarachopterabearscaleswhicharesparselyor
however,ofnorelevancehere. denselydispersedinasinglelayerwithonlyalittleoverlap.
The results of the above discussion on the pres- Scalesarealsopresentontheundersideofthewings,onthe
ence/absence of Amphiesmenopteran autapomorphies in head,antenna,thorax,andlegs.Thewingvestiture,however,
Tarachoptera are summarized in Table 1. The numerals re- resembles the Lepidoptera vestiture only superficially. On
fertocharacterenumerationinthetext. closerinspection,thearrangementofscalesandscaleshape
in Tarachoptera do not correspond with conditions in basal
AutapomorphiesofLepidoptera Lepidoptera(seeFig.11e),butasdemonstratedbySimonsen
(2001) the evolution of wing vestiture is rather complex in
Kobayashi and Ando (1988) found three embryological basalLepidoptera.
traits which corroborate the monophyly of Lepidoptera.
KristensenandSkalski(1998)demonstratedthemonophyly
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138 W.Meyetal.:TheblueprintoftheAmphiesmenoptera–Tarachoptera
16. Male gonopod (= valva) primarily undivided. Only
thetipsofthemalevalvaearevisible,barelyprotrudingbe-
yondthehindmarginsofsegmentIX(Fig.3(12)).Thevalves
arelikelyverysmallandwithoutanyhintsofsegmentation.
ThischaracterwaslistedbyKristensen(1984)asapossible
autapomorphy in contrast to the two-segmented gonopods
of ancestral Trichoptera. In both orders, the gonopods in
the basal lineages are usually large and protruding and
serve as grasping organs. The gonopods in Tarachoptera
differclearlyfromsuchastructure.Theassumedundivided
gonopods is either a convergence or a symplesiomorphy at
theAmphiesmenopteralevel.Inanycase,thecharacterdoes
notsupportarelationshiptoLepidopteraandTrichoptera.
17. Cerci lacking in both sexes. This regressive charac-
ter is probably of little significance. In Trichoptera it is
present in female Annulipalpia and Spicipalpia but missing
in almost all families of Integripalpia. The cerci are usually
flanking the anal opening or are placed behind it, lying on
a membranous short or telescoping segment X (ovipositor).
Amembranousterminalsegmentorovipositorisnotvisible
in Tarachoptera, and probably not present (Fig. 4(16, 17)).
Even in a retracted position deep inside the genital–anal
cavityoftheabdomen,thetipsofthepapillaeanalesshould
bediscernableatleast.
In summary, we did not find any characters that sup- Figure 10. Kinitocelis brevicostata spec. nov., holotype ♀
port a sister-group relationship between Tarachoptera and (BUB399): (a) ventral view of adult, (b) ventral side of tip of fe-
maleabdomen,and(c)head,ventralside.
Lepidoptera. The presence of wing scales is a shared
character,buttheirdifferentmorphologysuggestsadifferent
origin and makes it unlikely to assume common ancestry.
A predisposition for the development of scales could be As already mentioned in the discussion on character 1, a
thoughtorheldforthepresenceofanintrinsictraitinances- clearly developed haustellum is not discernible in Tara-
tral Amphiesmenoptera. The occurrence of wing scales in choptera. It is probably in a primitive state, small and not
derivedtaxaofTrichoptera,unrelatedtothatofLepidoptera, visibleexternally,asituationalsofoundinthenon-glossatan
canberegardedasevidenceofthispredisposition. Lepidoptera.
AutapomorphiesofTrichoptera 19. Absence of adult mandibular articulations. Sclerotized
and functional mandibles are plesiomorphic groundplan
Arguments for the monophyly of Trichoptera were first charactersofAmpiesmenoptera.Theyareretainedinprimi-
summarized by Ross (1956: 9), who provided nine adult tiveLepidopteraandhavewell-developedarticulationswith
characters.Weaver(1984)andMorse(1997:434)addedfive the head capsule. The mandibles of Trichoptera are non-
further homologues supporting monophyly for Trichoptera. functional (Klemm, 1966) and exhibit an array of different
Kobayashi and Ando (1988) provided four embryological stages of reduction spanning from fully sclerotized ones
characters.Threemorphologicalcharactersareapplicableto lying beneath the anteclypeolabrum to small membranous
Tarachopteraordeservesomenotes: lobes. The mandibles of Tarachoptera are also reduced;
however,thisreductionismoreadvancedandtheremainsof
18. Modified haustellum. The haustellum of Trichoptera is the mandibles appear as conical stumps in grooves beside
one of the main apomorphies of the order. It is a membra- thelabrum(Fig.3(14)).
nous, flexible organ, with a specialized fine structure on its
surfacethatfacilitatestheuptakeoffluidsubstancesorsmall 20. Nygma. The presence of a single nygma (= corneus
particles. In basal families the haustellum is small, with spot) in the base of apical cell 3 or fork 2 in the forewings
somewhat enlarged lateral sides (Klemm, 1966; Kubiak et is regarded a trichopteran synapomorphy. According to
al., 2015), whereas in families of Integripalpia the haustel- Kristensen (1984) nygmata occur in other holometabolan
lum is a large, bulbous organ resembling a short proboscis. orderstoo,whichmakesthisstructureaplesiomorphyatthe
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