Phylogeny, taxonomy, and biogeography of extant silky lacewings (Insecta: Neuroptera: Psychopsidae)

69 Pages·1993·28.7 MB·English

by  John D Oswald

MEMOIRS
OF THE
AMERICAN ENTOMOLOGICAL SOCIETY
NUMBER
40
PHYLOGENY, TAXONOMY, AND BIOGEOGRAPHY
OF EXTANT SILKY LACEWINGS
(INSECTA: NEUROPTERA: PSYCHOPSIDAE)
By
John D. Oswald
PUBLISHED BY THE AMERICAN ENTOMOLOGICAL
SOCIETY
AT THE ACADEMY OF NATURAL SCIENCES
PHILADELPHIA
1993
Paul M. Marsh
Editor
Issued 30 September 1993
Printed by BookCrafters in the United States ofAmerica
TypesettingandPageCompositionbyELYTRON
Swarthmore,Pennsylvania
TABLE OFCONTENTS
Introduction 1
Acknowledgments 2
Materials and Methods 3
Material 3
Terminalia Preparation 3
Illustrations 3
Terminology 3
CollectionAcronyms 3
Taxonomic Treatments 4
FamilyPsychopsidae Handlirsch 4
KeytoSubfamilies and Genera 6
SubfamilyZygophlebiinaeNavas 6
Genus Silveira Navas 6
Genus Cabralis Navas 8
Genus Zygophlebius Navas 8
SubfamilyPsychopsinae Handlirsch 10
Genus Baltnes Navas 10
GenusPsychopsis Newman 12
FunctionalMorphologyofMale and Female Terminalia 13
. .
Morphological Observations 14
Males 14
Females 16
Hypothesized Mechanics ofCopulationand Oviposition 19
.
Copulation 19
Oviposition 23
PhylogeneticAnalysis 26
Methods 26
Characters 27
Omitted Characters 41
Results 42
Classification 42
Biogeographic Analysis 46
Introduction 46
Biogeographic Hypotheses 48
Hypothesis Testing 49
Results 50
Discussion 50
Suggestions for Future Research 52
Literature Cited 53
Appendix 1: Synonymical CatalogofExtant Psychopsids,
and Material Examined 56
Appendix 2: Character State Data Matrix 65

Memoirs
of the
American Entomological Society
Number 40
Phylogeny, Taxonomy, and Biogeography of
Extant Silky Lacewings (Insecta: Neuroptera: Psychopsidae)
by
John D. Oswald
DepartmentofEntomology
NationalMuseumofNaturalHistory,NHB 165
SmithsonianInstitution
Washington,D.C.20560
Abstract.TheworldgeneraofthefamilyPsychopsidaearerevisedbasedonacomparativemorphologicalassessmentofadults
of21(of26)extantsilkylacewingspecies.Twosubfamiliesandfivegeneraarerecognized.ThesubfamilyZygophlebiinaecontains
threeAfricangenera,Silveira,Cabralis,andZygophlebius;thesubfamilyPsychopsinaecontainsthegeneraBalmes,fromsoutheast
Asia,andPsychopsis,fromAustralia.Agenerickeyandfulltaxonomictreatmentsofallsupraspecifictaxaarepresented,together
with a synonymical catalog of extant species. Three new species are noted. A novel male/female copulatory mechanism is
described;andseveralhypothesesareadvancedregardingthefunctionalmorphologyofthemaleandfemaleterminalia.Acladistic
analysis of the 21 examined species using 60 adult morphological characters demonstrates the holophyly of all recognized
supraspecifictaxa.Thefiverecognizedgeneraarecladisticallyrelatedasfollows:([Silveira+(Cabralis+Zygophlebius)]+[Balmes+
Psychopsis]). Alternative classifications of the family are discussed, particularly with regard to the Australian species. The
biogeographyofextantpsychopsidsisdiscussed. Thedistributionandcladogenyoflivingsilkylacewingsisconsistentwitha
Gondwananoriginoftheextantmembersofthefamily.ThepresentoccurrenceofBalmesspeciesinsoutheastAsiaisbestexplained
byeither(1)thenorthwesterlydispersalofanAustralian,probablyTertiary,ancestorthroughtheMalayArchipelagoor(2)the
northerlyraftingofaMesozoicancestoronariftfragmentofnortheasternGondwanaland.
INTRODUCTION
TheneuropterousfamilyPsychopsidae,or"silky latedate.Thefirstdescribedspecies,theattractive
lacewings," contains 26 extant species (including Psychopsis mimica from Australia, was initially
three undescribed, see Appendix 1) and a variety characterized by Edward Newman in 1842, and
of fossil forms ranging in age from Tertiary to more completely described the following year
Triassic (Table 5, page 48). This work treats the (Newman, 1843). Newman's implicit and appro-
phylogeny,taxonomy,biogeography,and several priatecomparisonofmimicatoacolorfullybanded
aspects of the functional morphology of living lepidopteranis memorializedinhis genericname
psychopsids. Psychopsis, derived from the Greek words psyche,
Becauseoftheabsenceofsilkylacewingsfrom butterfly,andopsis,appearance.In1889,Friedrich
Europeandadjacentlandareas,psychopsidscame Brauer described the firstAfrican psychopsid, ze-
to the attention of western science at a relatively bra. Two years later Robert McLachlan (1891) re-
1
SILKYLACEWINGS
corded the first silky lacewing from southeastern leled concurrent trends in other neuropterous
Asia, birmanus, and established the presence of families. Subsequently, New ([1989a]) made ex-
psychopsids inthe lastofthe three disjunctgeog- tensive use of terminalic traits to diagnose Aus-
—
raphies Australia, sub-Saharan Africa, and tralianpsychopsids.
—
southeast Asia which currently comprise the The present work builds upon this 150-year
tripartitedistributionofthelivingmembersofthis history. The major new contribution contained
family. hereisthefirstdetailedhypothesisofinterspecific
Handlirsch (1906-1908) recognized thedistinc- and intergeneric relationships within the family.
tiveness of the genus Psychopsis within the broad Thishypothesis isbased upon a cladistic analysis
concept of the family Hemerobiidae then current of 60 characters (mostly terminalic) scored for 21
and erected for it a new subfamily, the (of 26) living psychopsid species. The relative
Psychopsinae, which was subsequently adopted phylogenetic relationships inferred from this
by several contemporaries, e.g., Banks (1913) and analysisareused (1) asthebasisforanewgeneric
Nakahara (1914). Navas (1910), apparently then and suprageneric classification of the family and
unaware of all earlier work on this group, inde- (2) tointerpretthebiogeographichistoryofliving
pendently proposed two generic names, silky lacewings and to account for theirpresently
Zygophlebius and Balmes, to contain three new tripartite distribution. The descriptive aspects of
species (one each from Africa, Australia, and this work are limited to taxa of generic and
southeast Asia) that are now recognized as suprageneric rank. Consequently, full taxonomic
psychopsids. Navas placed his genera in a new treatments are provided only for the two sub-
hemerobiid tribe, the Zygophlebini. families and five genera recognized here. A com-
In the years following his 1910 paper, Navas prehensive synonymical catalog of extant
describedseveraladditionalnewpsychopsidsina psychopsid species, however, has been provided
series of general papers (e.g., Navas, 1910, 1912a, (Appendix 1), and synapomorphic characters
1912b) and one revisionary work, the "Ensayo supportingintragenericcladesamongtheexamined
monografico de la familia de los Sicopsidos." Al- species are included in the cladistic analysis.
though the latter work was not published until Readers are referred to the revisionary works of
1917 (Navas, 1917), itwas presented orally at the Tjeder (1960) and New ([1989a]) for species-level
Valladolid Congress of the Asociacion Espahola taxonomic treatments of African and Australian
para el Progreso de las Ciencias in October, 1915. psychopsids respectively. The interspecific tax-
Several aspects of this work are significant: (1) it onomy of southeast Asian psychopsids requires
contained the first elevation of psychopsids to additional study.
family rank—although, the family name
"Psychopsidae" itself was first published inde- ACKNOWLEDGMENTS
pendentlybyTillyard(1916:271),(2)itrepresented
thefirstcomprehensive,worldwide,reviewofthe I thank the following individuals for loaning
family, recognizing eight genera, 13 species, and materialforthisstudy:M.S.Moulds(AMS);Stephen
three varieties, and (3) it firmly established alar Brooks (BMNH); NormanD. Penny (CAS);James
characters, principally venation and wing pat- K. Liebherr (CUIC); M. Fletcher (DANSW); G.
J.
terning,astheprincipaldiagnosticandtaxonomic Miiller-Motzfeld(EMAU);LionelA.Stange(FSCA);
traitsusedintheearlydifferentiationofpsychopsid David G. Furth (MCZ); Les Minter (MINTER);
—
taxa an emphasis which was maintained Elaine De Coninck (MRAC); Mervyn W. Mansell
throughout the subsequent revisions of Tillyard (NCIP);UlrikeAspock(NHMW);R.Sithole(NMB);
([1919a]; Australia), Kriiger (1922; world), and M. N. Mungai (NMK);OliverS. Flint,Jr. (USNM);
Kimmins (1939;world). AndyE.Whittington(NMSA);E.C.Dahms(QMB);
More recently, Tjeder's (1960) revision of the JanvanTol (RNHL); E. G. Matthews (SAMA); M.
southern African psychopsid fauna conclusively A.Cochrane(SAMC);MartinaPusch(SMWW);M.
demonstrated the taxonomic utility of male and A.Schneider(UQIC);RoyDanielsson(UZIL);Kurt
femaleterminaliccharactersatallhierarchicallevels K.Giinther(ZMHB);andErnst-GerhardBurmeister
within the family. Tjeder's expansion of the taxo- (ZSM).
nomic character base in psychopsids to include I gratefully acknowledge Oliver Flint, Jr.
terminalic traits was a major advance and paral- (USNM), MervynMansell (NCIP),WayneMathis
JOHND.OSWALD
(USNM),TimothyNew(LaTrobeUniversity,Aus- bentback and the remaining connection severed.
tralia),andLionelStange(FSCA)forreviewingthe Careful cuts will avoid damage to the bursa, its
manuscriptandprovidingvaluablecomments.Les glands, the spermatheca, and the posterior ab-
Minter generously shared some unpublished ob- dominal chamber.
—
servations on several African species. George C. Illustrations. Terminalic drawingswerepre-
Steyskalprovidedhelpfuladviceonaproblematic pared using a drawing tube attached to a dissect-
nomenclatural question. ingmicroscopefromstructureseither(1)completely
Theresearchforthismonographwassupported embedded in glycerin jelly, or (2) partially em-
inpartby a SmithsonianPostdoctoral Fellowship beddedinpetroleumjellyandcoveredwithliquid
andwasundertakenwhiletheauthorwasinresi- glycerin.TheexcellentillustrationsfoundinTjeder
denceintheDepartmentofEntomologyattheU.S. (1960)arefrequentlycitedhereintoavoidexcessive
NationalMuseumofNaturalHistory(Smithsonian duplicationofartwork. References to figurescon-
Institution),Washington, DC. tained in Tjeder and other published works are
cited as "fig." [lower case f]; figures contained in
MATERIALSANDMETHODS thisworkarecited as "Fig." [uppercaseF].
—
Terminology. The morphological terminol-
—
Material. Thisstudywasbasedontheexami- ogy used here generally follows that of Tjeder
nationofapproximately650specimensrepresent- (1960) and New ([1989a]). A number of termino-
ing21 ofthe26knownextantpsychopsid species. logicaldifferences,however, occurin (1) themale
Theindividualsandinstitutionswho/whichloaned gonarcus/mediuncus/9th gonocoxite complex,
this material are listed below under "Collection and (2) the female ventral terminalia. A cross-
Acronyms." — referenceguidetothesedifferencesisgivenbelow.
Terminalia Preparation. The following gen- Justificationsofthetermsusedherearegiveninthe
eralsequencewasusedtopreparemaleandfemale text, particularly in the discussions of characters
terminalia: (1) remove abdomen from specimen used inthe cladistic analysis and under thehead-
[whole or only apical segments (ca. somites 6+)], ing "Functional Morphology ofMale and Female
(2) immerseovernightincold 10%KOHtosoften, Terminalia: Morphological Observations." Dis-
digest, andhydrate, (3) rinse in 70% ethyl alcohol cussionsoftheterminologyusedforthegonarcus
[EtOH],(4) [optional]staininsaturatedsolutionof aregiveninOswald(1988:396) andOswald (1993:
70%EtOHandChlorozolBlackforapproximately 155-157).
oneminute,thenrinsein70%EtOH,(5)transferto Cross-reference guide to terminalic terminol-
glycerinfor dissectionand examination. ogy (Format: Tjederian term = Current term
Rinsingisfacilitatedbyuseofasyringecontain- [(M)ale/(F)emale]): arcessus = mediuncus [M];
ingEtOHandafine,bent-tippedneedle.Removal entoprocessus = extrahemigonarcal process [M];
ofthe gonarcus/mediuncus/9thgonocoxitecom- glandulae accessoriae = bursal accessory glands
plexfromtheabdominalapexofthemalefacilitates [F]; gonapophysis lateralis = 9th gonocoxite [F];
detailed examination and illustration ofboth this paramere = 9th gonocoxite [M]; pregenitale = 8th
complexandthetergalandsternalelementswhich sternite [F]; spermatheca = bursa (membranous
enclose it. Removal can be accomplished using a portion) + spermatheca (scl—eroAtiMzeSd—portion) [F].
pairofjewelers'forcepsandafineprobetopartthe Collection Acronyms. Australian
membranesjoining these structures. Examination Museum, Sydney, NSW Australia; BMNH—The
of female terminalic structures—particularly the Natural History Museum, London, England;
—
bursa,spermatheca,posteriorabdominalchamber, CAS California AcademyofSciences, San Fran-
— —
andvarioussternalstructures isgreatlyaidedby cisco,CA,USA; CUIC CornellUniversityInsect
DANSW—
cutting the abdominal body wall in a horizontal Collection, Ithaca, NY, USA; Agricul-
planeatapproximatelythelevelofthespiracles.To tural Scientific Collections Trust (NSW Dept.
accomplishthis,firsttrimtheabdomentocontain Agric), Rydalmere, NSW, Australia; EMAU -
approximatelysomites6+.Then,withafinepairof Zoologisches Museum, Ernst-Moritz-Arndt-
—
scissors, cut horizontally through the body wall Universitat,Greifswald,Germany;FSCA Florida
fromthepleuralmembranetothedistalapexofthe StateCollectionofArthropods,Gainsville,FL,USA;
ectoproct.Withsimilarcutsmadeonbothsidesof MCZ—Museum of Comparative Zoology, Cam-
theabdomen,thedorsalhalfoftheabdomencanbe bridge, MA, USA; MINTER—Les Minter private
—
4 SILKYLACEWINGS
—
MRAC
collection, Pietersburg, South Africa; (Fig.32,vt).InthisformationthesubcostalandRl
MuseeRoyal—de1'AfriqueCentrale,Tervuren,Bel- spaces run parallel, and are subequal in width,
gium; NCIP National Collection of Insects, from near the base of the wing to a common (or
NHMW—
Pretoria,SouthAfrica; Naturhistorisches nearly common) distal point (the "anastomosis";
NMB—
Museum, Wien, Austria; NNaMtKur—al History Fig.32,ams)whereeachspaceiseitherconstricted
Museum,Bulawayo,Zimbabwe; National (by an interposed crossvein) or terminated (by
Museum ofKenya, Nairobi, Kenya; USNM—Na- brief vein fusion), and beyond (distal to) which
tional Museum of Natural History, Washington, pointtheparallel,subequal,natureofthespacesis
NMSA—
D.C., USA; NatalQMMuBs—eum, Pieter- not apparent. Other characteristic forewing traits
maritzburg, South Africa; Queensland include: (1) an unusually broad costal space, its
Museum, SouthBrisbane, Queensland, Australia; widthatleasttwicethewidthofthe"venatriplica"
—
RNHL Rijksmuseum van N—atuurlijke Historie, near the "anastomosis," i.e., in the "pterostigmal
SAMA
Leiden, Netherlands; South Australian region," (2) the presence of only one forewing
MSuAsMeCum—,SAoduetSlhaMiAdWfer,Wic—SaonutMhusAuesutmra,liaC,apAeustTroalwina,; nthyegcmoam,plloectaeteadbsbaesnaclelyofbpettewreoestnivgemiantsa,Raanndd(M4),t(h3e)
SouthAfrica; StateMuseum,Windhoek, presence of at least four (usually >10) crossveins
—
Namibia; UQIC University of Que—ensland, St. traversingthesubcostalspacebetweenitsbaseand
Lucia,Queensland,Australia; UZIL Museumof the "anastomosis."
—
ZoologyandEntomology,LundUniversity,Lund, Description. In the following treatment fa-
—
Sweden; ZMHB Museum fur Naturkunde d—er milialsynapomorphiesareindicatedbybracketed
Humboldt-Universitat, Berlin, Germany; ZSM character numbers (see Phylogenetic Analysis:
Zoologische Staats-sammlung, Miinchen, Ger- Charactersbelow). See also thediscussionsunder
many. theheading"FunctionalMorphologyofMaleand
Female Terminalia." Head: compound eyes
TAXONOMICTREATMENTS prominent; ocelli absent; 2 or 3 ocellar/cranial
pulvinae usually present (all lost in P. coelivaga);
FamilyPSYCHOPSIDAE Handlirsch antennaeshort,moniliform;toruliwithoneortwo
antennifers; maxillary palpi 5-segmented; labial
Psychopsinae Handlirsch [1906]:42 [As a subfamily of palpi3-segmented,ultimatelabialpalpomereseach
Hemerobiidae. Type genus: Psychopsis Newman,
bearingapalpimacula;mandibleswelldeveloped,
1842.].—Banks,1913:211[nomenclature];Nakahara,
slightly asymmetrical. Thorax: Pterothorax mac-
1914:491 [nomenclature].
Psychopsini.—Navas, 1912b:194 [As a tribe of ropterous; wings generally broadly triangular;
Hemerobiidae.Note:Navas'sproposalofthisname margins trichosorate; membrane fully
was apparently independent ofHandlirsch's prior microtrichose, but not macrotrichose; hind wing
useofthenamePsychopsinae. Navasproposedthe somewhat smaller than forewing. Forewing dis-
namePsychopsinitoreplacehisearliertribalname tinctlypatterned,sometimeswithbrightlycolored
Zygophlebini (based onZygophlebius Navas, 1910), patches; humeral plate prominent, sometimes
subsequent to his discovery ofthe existence of the elongate;costalspaceverybroadthroughout,width
oldergeneri—cnamePsychopsis.]. in "pterostigmal region" >2 times the combined
Psychopsidae. Tillyard, 1916:271 [nomenclature]; width of the adjacent Sc and Rl spaces [8];
Navas, 1917 [taxonomy]; Tillyard, [1919a] [tax-
pterostigma absent; costal gradate series present
onomy]; Kriiger, 1922 [taxonomy]; Kimmins, 1939
[taxonomy];Tjeder, 1960 [taxonomy];New, [1989a] [6] (occasionally secondarily reduced or lost);
proximal humeral trace recurrent andpectinately
[taxonomy].
BalmesiniNavas,1917:207[AsatribeofPsychopsidae. branched; subcostal space with more than four
Type genus: Balmes Navas, 1910.]. crossveins(butusually>10)[5];longitudinalveins
—
Psychopsididae [sic]. Martynova, 1949:161 [nomen- and costal veinlets densely macrotrichose. Legs
clature]. cursorial; paired tibial spurs present; tarsi 5-seg-
mented; pretarsus biclawed, arolium present,
—
Diagnosis. Medium-sized to large neurop- simple.
terans (forewing length 10-35 mm) readily distin- Male Terminalia (Figs. 1, 18). 8th Somite:
guished by the presence of a forewing and hind tergite narrowed, hemiannular; sternite
wingvenationalformationcalledthe"venatriplica" hemiannulartomoderatelyproducedposteriorly;
—
JOHN D.OSWALD
spiraclesopeningthroughventrallyprolongedmar- sternitealwayspresent,conformationvariable(nar-
gins of tergite [10] (secondarily free in several rowly rectangular [44], triangular, trapezoidal, or
species). 9th Somite: tergite arcuate, narrowed compactandlobed),fusedto7thsterniteinseveral
dorsally,expandedandelongatedventrally;tergal species. 9th Somite: tergite narrowed dorsally,
antecosta prominent, especially ventrolaterally, greatlyexpandedventrally,posteroventralexpan-
distal (ventral) ends articulated to anterolateral sion subtending ectoproct posteriorly; contralat-
margins of 9th sternite [14]; sternite variously eral tergal margins adpressed ventrally;
modified, usually somewhat reduced, composed posteroventral tergal surfaces inwardly revolute
principally of a pair of rigid lateral longitudinal and pilose [48]; tergal antecosta prominent for
costae [17], with various degrees ofadjacent scle- much of its length, particularly laterally and
rotization, apex frequently emarginate, lateral ventrolaterally, and giving rise to a pair of short
costae articulated proximally to antecosta of 9th broadapodemes(Fig.43,ala)abovespiraclesof8th
tergite [14],lateralcostaecontinuedinternallyasa somite [47]; a pair of slender inconspicuous
pair of short apodemes [15]. Ectoprocts: free apodemes (Fig. 43, da) issuing from near dorsal
dorsomedially, doubly (two species; e.g., Fig. 21) angles of posteroventral lobes of tergite [49];
orsinglylobeddistally, trichobothriatecalluscer- subgenitalepresent,asubrectangularplatebearing
cus always present. Gonarcus (Fig. 23): a promi- a pair of small, free, lateral lobes. Ectoprocts:
nentarcuateframeworkservingasthearticulatory proximolateral margins more or less fused to 9th
base of the mediuncus and 9th gonocoxites; tergite,developmentofsutureinthisareavariable,
extragonopons, extrahemigonarcus, and freeonsagittalmidlineofabdomen,butparasagittal
intrahemigonarcusprominent;intragonoponsab- marginsjoineddorsallybyanarrowmembranous
sentorpoorlydeveloped;extragonoponsgenerally rift which severely restricts lateral ectoproctal
protruded beyond extrahemigonarcus in lateral motion.9thGonocoxites:present,spathulate [50],
view. Mediuncus (=arcessus ofTjederand New): each with a prominent longitudinal costa [51], at
always present, proximal end articulated to pos- leastsomecochleariformsuprastylarsetaepresent
terior margin of extragonopons by means of a [54]; 9th gonocoxal styli present, each bearing a
transverse flexion line; curvature variable; apex field of short stout "digging" setae [55]. Internal
simple or bifid; apodeme of mediuncal adductor Structures:bursaalargemembranoussac,itsven-
musclealwaysdistinct[36];proximoventralsurface tral margins confluent with the adpressed dorsal
of mediuncus associated with a pair (sometimes edges of an elongate, sclerotized, slit-entry sper-
fusedsagittally)ofaccessoryscleritesdevelopedin matheca; colleterial gland present, one pair of
adjacent gonosaccal membrane [37]. 9th dorsodistal bursal accessory glands normally
Gonocoxites:alwayspresentandpaired;proximal present (see Character 56 for variants); posterior
ends articulated to posteroventral angles of abdominal chamber present [60]. Miscellaneous:
extrahemigonarcus;distalendsfusedatabdominal subanale always present.
—
midline [23] (exceptinP. coelivaga) toformamore Natural History and Immature Stages. See
or less rigid arch joining the extrahemigonarcus. individual generic treatmentsbelow.
—
Miscellaneous: hypandrium internum and Distribution. Range tripartly disjunct:
subanale alwayspresent. southern half of Africa (predominantly south of
FemaleTerminalia (Figs. 43-44). 7th Somite: the Equator), southeastern Asia (limits of distri-
tergitehemiannular,unmodified;sternitemoreor bution poorly known, reported from Burma,
lesshemiannular overall, posteriormarginwith a southernChina,Laos,andTaiwan),andAustralia
sagittal emargination and/or depression [40]; (widespread,butrec—ordspredominantlyeastern).
copulatory fovea present [42]. 8th Somite: tergite Included Taxa. Two subfamilies: Zygo-
narrowed, lateral extremities greatly prolonged phlebiinae (three genera, nine species) and
ventrally, posterior margin fused for much of its Psychopsinae (two genera, 17 species). For addi-
length (especially laterally and ventrolaterally) to tional discussion of family classification see
anteriormarginof9thtergite [46]; spiraclesopen- "PhylogeneticAnalysis:Classification"below.All
ingthroughventrallyprolongedmarginsoftergite; extant species arecataloged inAppendix 1.
SILKYLACEWINGS
KEYTOSUBFAMILIESANDGENERAOFPSYCHOPSIDAE
1 Distribution:Africa(Fig.54);Male:apexofmediuncussimple(Fig.4);Female:spermathecabearingapairof
hollowventrolaterallobes (Fig.46,vll) (Zygophlebiinae)2
Distribution:AustraliaorsoutheasternAsia(Fig.54);Male:apexofmediuncusemarginateorbifid(Fig.27);
Female: spermatheca lackinghollowventrolaterallobes,butsolid apodemalplates (Fig. 48,lap) maybe
presentventrolaterally (Psychopsinae)4
2(1) Head: vertexbearing3well-developedocellar/cranialpulvinae (Tjeder, 1960:175,fig.341);Female:hollow
ventrolaterallobesofspermathecaslender(Tjeder, 1960:179,fig. 365) SilveiraNavas
Head: vertex bearing 2 well-developed ocellar/cranial pulvinae [anteromedian pulvinus absent] (Tjeder,
1960:192,fig.405);Female:ventrolaterallobesofspermathecabroad(Fig.46) 3
3(2) Forewing:bearingdarkspotsonawhiteground(Tjeder,1960:199,fig.434A);HindWing:crossveinsclosing
"venatriplica"distallynotdarkbrown;Male:apexofmediuncusstronglydecurved(Fig.19);Female:bursa
withoutcorniformdiverticulae (Tjeder, 1960:203,fig.448) CabralisNavas
Forewing:bearingtransversebarsproximallyonayellowishorhyalineground[winglightlymottledinznsl,
barsinconspicuus] (Tjeder,1960:193,fig.412);HindWing:crossveinsclosing"venatriplica"distallydark
brown;Male:apexofmediuncusrecurved(Fig.14);Female:bursawithapairofcorniformdiverticulae(Fig.
47,cd) ZygophlebiusNavas
4(1) HindWing:bearingaprominentdarkmaculaon,beyond,orbelowanastomosisof"venatriplica"[Australia]
PsychopsisNewman
Hind Wing: dark distal macula lacking [Southeast Asia. The Australian speciesgallardi, regarded here as
Psychopsinaeincertaesedis,willalsokeyhere.] BalmesNavas
SubfamilyZYGOPHLEBIINAE Navas Genus SILVEIRANavas
Figs.1-5,39,41
Zygophlebini Navas, 1910:82 [As a tribe of
Hemerobiidae. Type genus: Zygophlebius Navas, Silveira Navas, 1912b:196 [Type species: Silveira
1910].—Navas, 1917 [taxonomy]. marmoratusNavas, 1912b:196 (=Psychopsis marshalli
— McLachlan, 1902),by monotypy. Etymology: from
Diagnosis. Cladistic analysischaracternum- the surname of Gonzalo Silveira, S. J., see Navas,
bers bracketed ([xx]), synapomorphies asterisked 1912b:196. Gender: Masculine (implied from the
(*). Head: Vertex bearing 2 (Cabralis and original combination Silveira marmoratus, Art.
Zygophlebius) or 3 (Silveira) well-developed ocel- 30d).].—Navas, 1917 [taxonomy]; Kimmins, 1939
[taxonomy];Tjeder, 1960 [taxonomy].
lar/cranialpulvinae [1];antennaltorulieachwith
Psychophasis Kriiger, 1922:44 [Type species: Psychopsis
2 antennifers, 1 medial and 1 lateral [2*, unique].
marshalliMcLachlan, 1902:234,byoriginaldesigna-
Male Terminalia: Anterolateral apodemes of 9th
tion. Etymology: unexplained, probably Psycho-
sternite continuing in line with proximal ends of (fromGr psyche,butterfly)+-phasis(fromGr.phasis
. —
lateral costae [15]; 9th Gonocoxites without true [fern.],appearance).Gender:Feminine.]. Kimmins,
superprocesses [24]; apex of mediuncus simple 1939 [synonymy].
[32]; apodeme of adductor muscle of mediuncus
—
insertingproximoventrallyonfloorofmediuncus Diagnosis. Cladisticanalysischaracternum-
[36]. Female Terminalia: Suprastylar setae of 9th bers bracketed ([xx]), synapomorphies asterisked
gonocoxite <50% (Silveira) or >50% (Cabralis and (*).Head:Vertexbearing3well-developedocellar/
Zygophlebius) cochleariform [54]; spermatheca cranialpulvinae[1].Wings:Forewingwithoutlight
bearingapair of—ventrolaterallobes [58*, unique]. browntransversebars [4];forewingcostalgradate
Distribution. —SouthernhalfofAfrica. series well developed [6]; hind wing without a
IncludedTaxa. Silveira(fourspecies),Cabralis darkdistalmacula [9]. MaleTerminalia: 8thster-
(two species, including one undescribed), and nitewithoutaposteromedianlobe [11];9thtergite
Zygophlebius (three species, including one without free posteroventral processes [12]; 9th
undescribed) [seeAppendix 1]. sternitenarrow andparallelsidedinventralview