phylogeny and higher classiﬁcation of suborder...

Zoological Journal of the Linnean Society

2002

136

371ndash400 With 75 figures

copy 2002 The Linnean Society of London


2002

136

371ndash400

371

Blackwell Science Ltd

Oxford UK

ZOJZoological Journal of the Linnean Society

0024-4082The Linnean Society of London 2002

136

Original Article

K YOSHIZAWAPHYLOGENY OF PSOCOMORPHA (lsquoPSOCOPTERArsquo)

Correspondence E-mail psocidresagrhokudaiacjp

Phylogeny and higher classification of suborder Psocomorpha (Insecta Psocodea lsquoPsocopterarsquo)

KAZUNORI YOSHIZAWA

Systematic Entomology Graduate School of Agriculture Hokkaido University Sapporo 060-8589 Japan

Received January 2002 accepted for publication May 2002

Phylogenetic relationships among all 24 families of suborder Psocomorpha (Insecta Psocodea lsquoPsocopterarsquo) areinferred based on adult morphology Monophyly of Psocomorpha is strongly supported by six autapomorphies Thepresently accepted four infraorders

minus

Psocetae Homilopsocidea Epipsocetae and Caeciliusetae

minus

are regarded asmonophyletic but Archipsocidae and Hemipsocidae previously assigned to Homilopsocidea and Psocetae respec-tively are regarded as the basalmost clades of the suborder Based on the results of the cladistic analysis a higherclassification for Psocomorpha is proposed Six infraorders (two new

minus

Archipsocetae Hemipsocetae

minus

and thefour aforementioned) are recognized Four new superfamilies are recognized within Homilopsocidea

ElipsocoideaLachesilloidea Pseudocaecilioidea

and

Peripsocoidea

Two superfamilies are recognized within CaeciliusetaeAsiopsocoidea and Caeciliusoidea Descriptions of taxa above family level are provided copy 2002 The LinneanSociety of London


2002

136

371

minus

400

ADDITIONAL KEYWORDS ndash new infraorder ndash superfamily ndash phylogeny ndash Psocomorpha ndash Psocoptera ndash

systematics

INTRODUCTION

The order Psocoptera (psocids booklice or barklice) isa small order of insects comprising

c

4110 describedspecies (Garciacutea Aldrete 1996) They range from 1to 10 mm in length and are characterized by a well-developed postclypeus long antennae pick-likelacinia reduced prothorax and well-developedpterothorax

Psocomorpha is the largest suborder in Psocopteracontaining 24 of the 37 psocopteran families Psoco-morphan families are classified into four groups Epip-socetae Caeciliusetae Homilopsocidea and PsocetaeThis taxonomic system was first proposed by Pearman(1936) and remains widely adopted with some minoralterations

Phylogenetic relationships within Psocoptera werefirst extensively studied by Smithers (1972) Howeverin his proposed dendrogram some lineages werebased on plesiomorphic or homoplastic characterstates nearly 20 years later he admitted (Smithers

1991) that his phylogenetic system needed revisionPsocomorphans exhibit a range of fascinatingbehaviours including stridulation aggregation sub-sociality and nesting Investigation into the evolution-ary aspects of these behaviours requires a reliablephylogenetic system

In the present paper I infer the phylogenetic rela-tionships among families of Psocomorpha based onadult morphology Six infraorders (two of them new)are recognized In addition six superfamilies arerecognized two within Caeciliusetae and four (new)within infraorder Homilopsocidea

HISTORY OF THE HIGHER CLASSIFICATION OF PSOCOMORPHA

The presently accepted taxonomic categories abovefamily level within Psocoptera were first proposed byPearman (1936) Unlike earlier efforts based on a fewprominent characters such as wing venation andnumber of tarsomeres his classification was based onan analysis of a wide range of external morpholo-gical characters He proposed eight family groups(infraorders of the present paper) in Psocoptera ofwhich four

minus

Epipsocetae Caecilietae (

=

Caeciliuse-

372

K YOSHIZAWA



2002

136

371ndash400

tae) Homilopsocidea and Psocetae

minus

are presentlyassigned within Psocomorpha (Table 1)

Roesler (1944) proposed three suborders in Psocop-tera corresponding to the presently acceptedTrogiomorpha Troctomorpha and Psocomorpha(

=

Eupsocida

sensu

Roesler) In Psocomorpha he rec-ognized three family groups Epipsocetae Psocetaeand Caecilietae (

=

Caeciliusetae) Psocetae andHomilopsocidea of Pearmanrsquos system were included inPsocetae (

sensu

Roesler) He also grouped or splitsome of Pearmanrsquos families as shown in Table 1

The taxonomic system proposed by Badonnel (1951)is a combination of those of Pearman and Roesler Heretained all Pearmanrsquos families and family groupsand arranged them into Roeslerrsquos subordersBadonnelrsquos system has been widely accepted with onlytwo modifications Smithers (1967) transferred Calop-socidae from Caeciliusetae to Homilopsocidea andMockford (1976) transferred Hemipsocidae fromHomilopsocidea to Psocetae

The monograph by Smithers (1972) is the mostextensive study of the higher classification of Psocop-tera He also investigated the phylogenetic relation-ships among all families and genera of PsocopteraHowever as mentioned above some lineages in his

dendrogram were defined by symplesiomorphic orhomoplastic characters and his phylogenetic classifi-cation has not been accepted

The classifications adopted by Smithers (1996) andLienhard (1998) are based on Badonnel (1951) andinclude the previous updates A few years prior to thisMockford (1993) raised the status of Dasydemellidaeformerly a subfamily of Amphipsocidae but this wasnot followed by Smithers and Lienhard

Within the family groups of Psocomorpha the fol-lowing phylogenetic hypotheses have been proposed

(1) Pseudocaeciliidae and Calopsocidae comprise amonophyletic group (Smithers 1967 Thornton ampSmithers 1984)(2) Elipsocidae and Mesopsocidae comprise a mono-phyletic group with Philotarsidae the sister group(Badonnel amp Lienhard 1988)(3) Caeciliusidae Stenopsocidae and Amphipsocidaecomprise a monophyletic superfamily Caeciliusoideawith Asiopsocidae the sister group (Mockford amp GarciacuteaAldrete 1976)(4) Bryopsocidae is the sister group of the clade com-prising Pseudocaeciliidae and Calopsocidae (Mockford1984)

Table 1

History of the higher classification of Psocomorpha

PHYLOGENY OF PSOCOMORPHA (lsquoPSOCOPTERArsquo)

373



2002

136

371ndash400

(5) Closer relationships between Philotarsidae and(a) Mesopsocidae or (b) Elipsocidae are also possible(Mockford 1984)(6) Lachesillidae and Elipsocidae may be phylogeneticsister groups (Mockford amp Sullivan 1986)(7)

Ectopsocus

and its close allies appear to be closerto the lachesillids than to

Peripsocus

(Mockford 1972)

Eertmoed (1973) proposed phenetic relationshipsfor the Epipsocetae group

minus

a classification which hasbecome widely accepted

MATERIAL AND METHODS

Lists of the taxa examined in this study or selectedfrom the literature are available at httpinsect3agrhokudaiacjppsoco-webdataindexhtmlAll internet resources in this paper are also availablefrom the author on request Terminology mainly fol-lows Matsuda (1965 1970 1976) and Smithers (1991)

Dried and wet specimens were used for the studyFor the observation of external structures the mate-rial was placed in a 5 solution of KOH at 45

deg

C for1ndash3 h depending on size It was then washed with dis-tilled water and stored in 80 ethanol Dissectedstructures were stained with Delafieldrsquos Hematoxylinthen observed and illustrated For the study of inter-nal structures the wet material was dissected in 80ethanol and stained with methylene blue A LeicaMZ12 stereoscopic microscope was mainly used forobservation and illustration For extremely smallstructures the material was slide-mounted in euparaland an Olympus BX50 compound light microscopeused for observation and illustration

As a rule exemplars (Yeates 1995) for the analysiswere selected from all psocomorphan families basedon the following criteria

(1) Selection to follow the taxonomic system adoptedby Mockford (1993)(2) At least two exemplars from each family one fromthe nominotypical genus and the other from a genusassumed to be distantly related to it(3) Only one exemplar when the family is representedby only one genus or comprises uniform taxa(4) More than two exemplars when the family ishighly diverse (strategy 3 of Hills 1998)

In some cases exemplars and character informationwere selected from published descriptions and illus-trations They are listed in the next section

The cladistic analysis was based on the externalmorphology and musculature or other internal struc-tures of adults Table 2 features the data matrix itis also available at httpinsect3agrhokudaiacjppsoco-webdataindexhtml Based on this matrixmost parsimonious trees were found using

PAUP

40b8 (Swofford 2001) A heuristic search was per-formed with TBR and the addition sequences lsquorandomrsquooptions (1000 replications) chosen Character stateswere optimized using MacClade 40 (Maddison ampMaddison 2000) The cladogram was translated toa phylogenetic system following the annotatedLinnaean system method (Wiley 1981)

CLADISTIC ANALYSIS

E

XEMPLARS

In this section I list the criteria for selecting exem-plars from each family with minimum reduction ofmorphological diversity

Outgroup exemplars were selected from the remain-ing two psocopteran suborders Trogiomorpha andTroctomorpha As the former is regarded as monophyl-etic and apparently distantly related to Psocomorphaonly one primitive exemplar

Echmepteryx lunulata

was selected from it Mockford (1967) regardedAmphientomidae of Troctomorpha as the sister groupof Psocomorpha and two exemplars

Paramphiento-mum yumyum

and

Tineomorpha

sp (from Malaysia)were selected from this family because the sistergroup has the strongest effect upon estimating thecharacter states of the ingroup node (Maddison

et al

1984) Two other exemplars

Tapinella

sp (fromTaiwan) and Troctopsocidae Gen sp (from Malaysia)were selected from other troctomorphan families eachof which is regarded as distantly related to Amphien-tomidae The list selected on the basis of the four rulesin the preceding section is as follows

(1) Archipsocidae

Archipsocus

sp (from Mexico) and

Pararchipsocus pacificus

(2) Hemipsocidae comprises two genera

Hemipsocus

and

Anopistoscena

the latter characterized only byautapomorphic forewing venation In contrast all spe-cies of

Hemipsocus

are very similar to each otherexcept for autapomorphic modifications of the malegenitalia Thus only one exemplar

Hemipsocus chlo-roticus

was selected(3) Myopsocidae

Myopsocus

sp (from Japan) and

Lichenomima muscosa

(4) Psilopsocidae comprises only one genus

Psilopso-cus

one exemplar

Psilopsocus

sp (from Indonesia)was selected(5) Psocidae one of the most diverse families in Pso-comorpha with many potential exemplars Howevermost modifications are autapomorphic for each taxonand provide little phylogenetic information for highercategories Thus two exemplars

Psocus

sp (nr

bipunc-tatus

from Japan) and

Psocidus

sp (from Japan)were selected(6) Elipsocidae

Elipsocus abdominalis

and

Hem-ineura dispar

374

K YOSHIZAWA



2002

136

371ndash400

Table 2

Data matrix for phylogenetic analysis of Psocomorpha (

=

missing data)

10

20

30

40

50

60

Echmepteryx

0000000000 0000000000 020000000 0000000000 000000010 0000000110 00000000

Tapinella

0000000000 000000000 000000000 0000200000 0000000100 0000000000 00000000

Troctopsocidae

0000000000 0000000000 100000000 0000200000 0000000100 0000000000 00000000

Tineomorpha

0000000000 0000000000 020000000 0000000000 000000010 0000000000 00000000

Paramphientomum

0000000000 0000000000 020000000 1000000000 000000010 0000000000 00000000

Archipsocus

1000000010 0002100100 021110000 1111000001 1100000102 000000001 20000000

Pararchipsocus

1000000010 0002100100 021110000 1111000001 1100000102 000000001 00000000

Hemipsocus

1011000010 0010000200 0000100001 1011000001 100100010 0010000000 00000000

Psilopsocus

1010000010 0010010111 1000100000 1011200010 1001000102 001001100 00100000

Psocus

1010100010 0010010111 1000100002 1011200011 0001100002 0010010100 00100001

Sigmatoneura

1010100010 0010010111 1000100002 1011200011 0001100002 0010010100 00100001

Myopsocus

1010000010 0010010111 1000100002 1011200010 100100010 0011010000 00000000

Lichenomima

1010000010 0010010111 1000100002 1011201010 1001000000 0011010000 00000000

Elipsocus

1000000010 0011110111 0000100000 1011100000 0000000000 0000100000 01000000

Hemineura

1000000010 0011110111 0000100000 1011100000 0000000000 0000100000 01000000

Ectopsocus

1010000010 0111110111 0000100010 1011201001 1100000000 1000200000 01000100

Eolachesilla

1010000010 0111110111 0000100000 1011200000 1000000000 1000000000 01000100

Nanolachesilla

1011100010 0111110111 0000100000 1011200001 1000000000 100030001 100100

Lachesilla

(Vietnam)

1010100010 0111110111 0000100000 1011200001 000000012 000000001 100100

Lachesilla

1010100010 0111110111 0000100000 1011200001 100000012 000000001 200100

Trichopsocus

1010100010 0011110211 0001100000 1011000101 1110000000 1000000100 01011000

Pseudocaecilius

1010100010 0011110211 0011100000 1011010001 1110111000 0000200101 01011000

Ophiodopelma

1010100010 0011110211 0011100000 1011010001 1100110000 1000200101 01011000

Heterocaecilius

1010100010 0011110211 0011100000 1011010001 1110111000 1000200101 01011000

Calopsocus

0010101110 0011110211 0111100000 1011010001 1010111000 1000200101 01011000

Bryopsocus

1 0100100000 1100 00000000 100030100 0102000

Kaestneriella

1011000010 0011110111 0000100010 1011200001 0000000001 1100300100 01012000

Peripsocus

1010000010 0011111111 1000100010 1011000001 0000000001 1100300100 01012000

Philotarsus

1010100010 0011110111 0011100000 1111010100 0000100100 0100300100 01012000

Aaroniella

1010100010 0011110111 0011100000 1111010100 0000000100 1100300100 01012000

Idatenopsocus

1000000010 0011110111 1000100000 1011100100 0000000000 0100300102 01012000Mesopsocus 1000000010 0011110111 1000100000 1011100000 0000000100 0100300102 01012000Epipsocus 1011111110 1011010111 0001100100 1111020001 0000000000 000000001 00000120Epipsocopsis 1011111110 1011010111 0001100100 1111020001 0000000000 000000001 00000120Dolabellopsocus 1010111110 1011010111 0001100100 0011020001 1000000000 000000001 00000120Isthmopsocus 1010111110 1011010111 0001100100 0011020001 1000000000 000000001 00000120Cladiopsocus 1010111110 1011010111 0001100100 0011020001 00000000 0000000000 00000120Spurostigma 1010111110 1011010111 0001100000 0011020001 0000000000 0000000000 00000110Triplocania 1011111110 1011010111 0001100100 0111020000 000000110 0000000000 00000110Ptiloneura 1011111110 1011010111 0001100100 0111020000 000 000000000 00000110Notiopsocus 1010001111 0111111211 0000100010 1011000001 2100000000 0000000000 10000200Asiopsocus 1101111 11 1000100000 10120001 0100000 000000000 1000020Stenopsoucs 1000001110 1111111111 0000101001 1011100001 1110000000 0000001000 0000020Graphocaecilius 1000001110 1111111111 0000101001 1011200001 1110000000 0000001000 00000200Amphipsocus 1100001110 1111111111 0001100000 1111020001 1110000000 0000000000 0000020Kolbia 1100001110 1111111111 0001100000 1111020001 1110000000 0000000000 0000020Matsumuraiella 1100001110 1111111111 0101100000 1111100001 1110000000 0000000000 00000200Caecilius 1000001110 1111111111 0000100000 1111000001 1110000000 0000000000 0000020Kodamaius 1100001110 1111111111 0001100001 1111000001 1110000000 0000000000 0000020Dasypsocus 0000001110 1111111111 0000100000 1111000001 1110000000 0000000000 0000020

PHYLOGENY OF PSOCOMORPHA (lsquoPSOCOPTERArsquo) 375

copy 2002 The Linnean Society of London Zoological Journal of the Linnean Society 2002 136 371ndash400

(7) Lachesillidae exemplars selected from all threetribes of this highly diverse family Lachesilla pedicu-laria and L sp (pedicularia group from Vietnam)were chosen from the highly specialized and divergentgenus Lachesilla The others were Nanolachesillananciae of Graphocaeciliini and Eolachesilla chilensisof Eolachesillini(8) Ectopsocidae only one exemplar Ectopsocus sp(from Japan) as the family consists of uniform taxa(9) Trichopsocidae comprises only one genus Tri-chopsocus one exemplar T dalii was selected(10) Pseudocaeciliidae one of the most diverse fami-lies in the Homilopsocidea both morphologically andbiologically Potential exemplars from both dead-foliage (Ophiodopelma glyptocephalus) and living-foliage dwellers As most taxa fall into the latter cat-egory two exemplars Pseudocaecilius kagoshimensisand Heterocaecilius anomalis were selected(11) Calopsocidae although the forewing charactersare highly specialized and variable among generaother morphological characters are rather similarthroughout the family Thus only one exemplar Calop-socus infelix was selected(12) Bryopsocidae comprises only one species Bry-opsocus townseni Specimens were unavailable for thisstudy and character information was extracted fromthe literature(13) Peripsocidae Peripsocus sp (from Japan) andKaestneriella guatemalensis(14) Philotarsidae Philotarsus flaviceps andAaroniella sp (from Japan)(15) Mesopsocidae Mesopsocus hongkongensis andIdatenopsocus orientalis(16) Cladiopsocidae Cladiopsocus garciai and Spuro-stigma epirotica(17) Dolabellapsocidae Dolabellapsocus roseus andIsthmopsocus sp (from Mexico)(18) Ptiloneuridae Triplocania spinosa and Ptilo-neura splendida The male of the latter species isunknown(19) Epipsocidae Epipsocus sp (from Malaysia) andEpipsocopsis sp (from Taiwan)(20) Asiopsocidae Asiopsocus sonorensis and Notiop-socus aldretei Character information for Asiopsocuswas selected from the literature as specimens wereunavailable(21) Stenopsocidae Stenopsocus sp (nr aphidiformisfrom Japan) and Graphopsocus cruciatus(22) Amphipsocidae ss (= sensu Mockford 1993)Amphipsocus rubrostigma and Kolbia fusconervosa(23) Dasydemellidae specimens of the nominotypicalgenus were unavailable One exemplar Matsumu-raiella radiopicta was therefore selected as judgingfrom descriptions and illustrations Matsumuraiella isvery similar to Dasydemella The distinction is con-sidered to be of little phylogenetic consequence

(24) Caeciliusidae one of the largest families ofPsocomorpha containing widely morphologicallydivergent species Additionally no decisive autapomor-phy supporting monophyly of this family has been pro-posed Thus three exemplars Caecilius fuscopterusDypsocus coleopteratus and Kodamaius directus all ofwhich are considered distantly related were selected

CHARACTERS

Of the 68 characters used in the cladistic analysis 11were multistate and the remainder binary Characterswere equally weighted a priori but a posteriori weight-ing was applied as discussed below All multistatecharacters were initially treated as unordered for theanalysis However if analysis and morphologicalobservation suggested that a character was possiblyordered it was optimized as ordered on the resultantcladogram Descriptive or analytical informationconcerning characters their states or polarities isincluded in Remarks Uninformative characters wereexcluded from the matrix but will be discussed below

Character length (L) consistency index (CI) reten-tion index (RI) and polarity of character states areincluded and were calculated using MacClade 40(Maddison amp Maddison 2000)

1 Vertex (0) sharply angled (Fig 1) (1) rounded(Fig 2) L = 3 CI = 033 RI = 067 (1) is an autapo-morphy of Psocomorpha but reversals have occurredin Calopsocidae and in a few genera of Caeciliusidae2 Vertex (0) without concavity (1) with pair of con-cavities L = 1 CI = 1 RI = 1 (1) is an autapomorphyof Amphipsocidae sl3 Internal ridge of epistomal suture (0) broad (Fig 3)(1) narrow (Fig 4) L = 4 CI = 025 RI = 083 (1) is anautapomorphy of the clade which comprises all psoco-morphan families excluding Archipsocidae However(0) is considered secondarily derived independently inElipsocidae Mesopsocidae and Caeciliusoidea4 Epistomal suture (0) complete (Figs 3 4) (1)absent dorsally (Fig 5) L = 4 CI = 020 RI = 033 (1)is an autapomorphy supporting the monophyly ofHemipsocidae and has evolved independently inNanolachesilla of Lachesillidae and two families inEpipsocetae5 Position of anterior tentorial pit (0) on ventral mar-gin of cranium (Fig 1) (1) separated from ventralmargin of cranium (Fig 2) L = 5 CI = 020 RI = 079(1) is regarded as the apomorphic condition in Psoco-morpha but highly homoplastic6 Labrum (0) without paired longitudinal sclerites(Fig 4) (1) with paired longitudinal sclerites (Fig 5)L = 2 CI = 050 RI = 088 (1) is an autapomorphy ofEpipsocetae and was derived independently also inAsiopsocus Remarks within the species examined in

376 K YOSHIZAWA


Figures 1ndash14 (1ndash5) Head of Echmepteryx lunulata (1) and Psococerastis nubila (2) lateral view E lunulata (3) P nubila(4) and Epipsocopsis sp (5) anterior view (6ndash7) mandible of P nubila (6) and Matsumuraiella radiopicta (7) anterior view(8ndash12) maxilla of P nubila (8) and M radiopicta (9) lateral view Paramphientomum sp (10) and P nubila (11) showingmuscles attached to galea anterior view Notiopsocus aldretei (12) showing apex of lacinia (13ndash14) labium of P nubila (13)and Stenopsocus sp (14) posterior view Abbreviations ata = anterior tentorial arm cly = clypeus fr = frons g = galea l =lacinia lr = labrum

this study (1) is observed only in Epipsocetae andAsiopsocus but Mockford (1977) and Mockford ampGarciacutea Aldrete (1976) noted that it is also observed insome Caeciliusoidea7 Mandible (0) outer margin rounded and posteriormargin not hollowed (Fig 6) (1) outer margin angled

and posterior margin hollowed (Fig 7) L = 2CI = 050 RI = 094 (1) is an autapomorphy of theclade Epipsocetae + Caeciliusetae and independentlyderived in Calopsocidae8 Galea (0) flat (Fig 8) (1) ball-shaped (Fig 9)L = 2 CI = 050 RI = 094 (1) is an autapomorphy



of the clade Epipsocetae + Caeciliusetae and wasderived independently also in Calopsocidae Remarkscondition of this character appears strongly correlatedwith character 79 Stipito-galeal muscle (s-g7) (0) present (Figs 10)(1) absent (Fig 11) L = 1 CI = 1 RI = 1 (1) is an auta-pomorphy of Psocomorpha10 Lacinia (0) without broadened region (Fig 9) (1)with externally broadened subapical region (Fig 12)L = 1 CI = 1 RI = 1 (1) is an autapomorphy ofAsiopsocidae11 Labial palpus (0) rounded (Fig 13) (1) triangular(Fig 14) L = 2 CI = 050 RI = 093 (1) is a possibleautapomorphy of the clade Epipsocetae + Caeciliuse-tae Remarks Epipsocetae and Caeciliusidae have asomewhat triangular externally expanded labialpalpus In contrast Notiopsocus has a rounded labialpalpus The state of this character for Asiopsocus ispresently unknown Consequently the state of thischaracter for the common ancestor of Epipsocetae +Caeciliusetae cannot yet be determined12 Preepisternum of prothorax (0) short (Fig 15) (1)elongate (Fig 16) L = 2 CI = 050 RI = 091 (1) isderived independently at least twice in Caeciliusetaeand Lachesillidae + Ectopsocidae and regarded as anautapomorphy of each clade13 Mesothorax (0) not strongly bulged (1) greatlybulged dorsally (Fig 17) L = 1 CI = 1 RI = 1 (1) is anautapomorphy of the clade which contains all pso-comorphan families except Archipsocidae RemarksMockford (1967) regarded (1) as an autapomorphy ofPsocomorpha However the pterothorax of Archipso-cidae is less developed and its dorsal margin is almostlevel with the vertex14 Mesothoracic dorso-ventral flight muscle (0) com-prises one or two muscles inserted into the base oftrochantin (Fig 19) (1) divided into three musclesexternal two inserted into the precoxal bridge andinternal one into the trochantin (Fig 20) (2) dividedinto three muscles internal two inserted into theprecoxal bridge and external one into the trochantin(Fig 21) L = 3 CI = 1 RI = 1 (1) is derived for theclade comprising Homilopsocidea + Epipsocetae +Caeciliusetae and regarded as autapomorphic (2) isunique to Archipsocidae and supports its monophylyRemarks in Trogiomorpha Troctomorpha Hemipso-cidae and Psocetae a less developed dorso-ventralflight muscle (probably corresponding to t-p 5 6 andt-ti 2) is inserted into the base of the trochantin(Fig 19) In contrast the dorso-ventral flight musclesof other psocomorphan families are split into two oneattached to the trochantin and the other to the pre-coxal bridge (Fig 20) In Archipsocidae the largerinner muscle is inserted into the precoxal bridge andthe smaller outer muscle into the trochantin (Fig 21)whereas in the other families the smaller inner mus-

cle is inserted into the trochantin and the larger outermuscle into the precoxal bridge (Fig 20) This sug-gests that splitting of the dorso-ventral flight muscleshave evolved independently at least twice15 Precoxal bridge (0) narrow (Fig 22) (1) broad(Fig 23) L = 3 CI = 033 RI = 089 (1) is apomorphicand observed independently in Archipsocidae Hom-ilopsocidea and Caeciliusetae Remarks see previouscharacter and lsquoMonophyly of Psocomorpha and rela-tionships of infraordersrsquo below16 Membranous region of metapleuron (0) narrow(Fig 18) (1) broad (Fig 17) L = 1 CI = 1 RI = 1 (1)is an autapomorphy supporting the clade com-prising Psocetae Homilopsocidea Epipsocetae andCaeciliusetae17 Campaniform sensilla on forewing radius (0)evenly distributed (1) divided into two groups L = 2CI = 050 RI = 089 (1) is an autapomorphy ofCaeciliusetae and was also derived independently inPeripsocus18 Apex of first axillary sclerite of forewing (0)without minute process proximally (Fig 28) (1) withminute process proximally (Fig 29) (2) broadened(Fig 30) L = 4 CI = 050 RI = 080 (1) is an auta-pomorphy of Psocomorpha and (2) evolved indepen-dently from (1) at least three times in HemipsocidaePseudocaecilioidea and Notiopsocus Remarks inmost insects the apex of first axillary sclerite (1Ax)has a long neck region that curves externally Thiscondition is also observed in Trogiomorpha and Troc-tomorpha (Fig 28) In contrast 1Ax of Psocomorphais subtriangular and often has a minute processproximally In some taxa such as Hemipsocidae andPseudocaeciliidae the apex of 1Ax is broadened andthe proximal minute process is inconspicuous (state2) This character state is regarded as being derivedfrom state 119 Second axillary sclerite (2Ax) and proximalmedian plate (PMP) of forewing (0) articulated witheach other (Fig 31) (1) fused with each other (Fig 32)L = 1 CI = 1 RI = 1 (1) is an autapomorphy of theclade comprising all psocomorphan families exceptArchipsocidae and Hemipsocidae Remarks Brodsky(1994) mentioned that the roof-like folding of thewings had presumably appeared in the common ances-tors of the Paraneoptera and fusion of 2Ax and PMPwas regarded as one of the most important changes forproviding this folding However fusion of 2Ax andPMP is never observed in outgroup suborders Appar-ently Brodsky (1994) misidentified the homology ofwing base structures of Hemiptera and 2Ax and PMPare articulated with each other in Thysanoptera andHemiptera (Yoshizawa amp Saigusa 2001) Thus fusionof 2Ax and PMP did not appear in the common ances-tor of Paraneoptera but is regarded as an autapomor-phy of the clade comprising all psocomorphan families

378 K YOSHIZAWA


except Archipsocidae and Hemipsocidae (0) and (1)can be easily distinguished externally without dissec-tion In psocids possessing (0) the posteroproximalmargin of 2Ax and the proximal margin of DMP (distalmedian plate) are closely approximated with eachother when the wings are closed (Fig 31) In psocids

possessing (1) 2Ax and DMP are separated by PMPwhen the wings are closed (Fig 32)20 Posterior margin of folded forewings proximal tonodulus (0) closely approximated with each other(Fig 31) (1) separated from each other (Fig 32) L = 1CI = 1 RI = 1 (1) is an autapomorphy of the clade

Figures 15ndash27 (15ndash16) prothorax of Psococerastis nubila (15) and Ectopsocus sp (16) lateral view (17) thorax of Pnubila lateral view (18) metapleuron of Hemipsocus chloroticus lateral view (19ndash21) mesothoracic dorso-ventral flightmuscle of Paramphientomum sp (19) Stenopsocus sp (20) and Parachipsocus pacificus (21) (22ndash23) mesothoracic precoxalbridge and trochantin of Psococerastis nubila (22) and S sp (23) (24ndash25) apex of hind tibia of S sp (24) and Psocus sp (25)distal tarsomere and claws of Psocus sp (26) and S sp (27) Abbreviations aes = anepisternum cx = coxa em = epimerones = episternum kes = katepisternum pb = precoxal bridge pes = preepisternum tr = trochantin



comprising all psocomorphan families except Archip-socidae and Hemipsocidae21 Forewing margin and veins (0) setose (Fig 37)(1) bare (Fig 36) L = 5 CI = 020 RI = 056 (1) isregarded as a derived condition but highly homoplastic22 Membranous region of forewing (0) bare (Figs 36minus40) (1) sparsely setose (2) densely setose (Fig 35)L = 5 CI = 040 RI = 050 (1) is derived indepen-dently at least three times within Psocomorpha (2) isconsidered as the most primitive condition of Psoco-morpha Remarks exemplars with scaly forewingwere coded as (2) because scales can be considered asmodified setae Because a possible sister group of Pso-comorpha Amphientomidae has scaly wings and thebasalmost psocomorphan clade Archipsocidae has adensely setose forewing (2) is considered to be thebasic condition of Psocomorpha but further study isrequired23 Marginal setae of forewing (0) not crossing(Fig 39) (1) crossing on apical margin (Fig 37) L = 3CI = 033 RI = 071 (1) is apomorphic and is derived

independently at least three times in ArchipsocidaePseudocaecilius + Calopsocidae and Philotarsidae andregarded as an autapomorphy of each clade Remarksexemplars with bare forewings were scored lsquo0rsquo24 Setae on veins of forewing (0) 0ndash1 row (1) 2 ormore rows (Fig 39) L = 4 CI = 025 RI = 083 (1) isregarded as the derived condition and derived inde-pendently at least four times Remarks although thischaracter is rather homoplastic it is consistent withina family superfamily or infraorder (1) is regarded asautapomorphic for Pseudocaecilioidea PhilotarsidaeEpipsocetae and Amphipsocidae sl and no reversalhas been deduced The character state of exemplarswith scaly forewings (eg Amphientomidae) or setae(Archipsocidae) could not be determined and wasscored lsquorsquo25 Pterostigma (0) not thickened (1) thickenedL = 1 CI = 1 RI = 1 (1) is an autapomorphy of Psoco-morpha Remarks although a thickened pterostigmais observed in Archipsocidae it appears to be muchthinner than in other families of Psocomorpha In this

Figures 28ndash34 (28ndash30) right first axillary sclerite of Paramphientomus sp (28) Peripsocus quercicola (29) and Pseudocae-cilius kagoshimensis (30) (31ndash32) mesothorax and forewing of Hemipsocus chloroticus (31) and Psococerastis nubila (32)dorsal view (33ndash34) nodulus of Paramphientomum sp (33) and Psococerastis nubila (34) Abbreviations 2ax = second axil-lary sclerite dmp = distal median plate fw = forewing pmp = proximal median plate

380 K YOSHIZAWA


instance I did not code them separately but differentdegrees of thickness of the pterostigma may providefurther evidence for the phylogenetic placement ofArchipsocidae as the basalmost clade of Psocomorpha

26 Forewing veins (0) normal (Figs 36minus40) (1)reduced (Fig 35) L = 2 CI = 1 RI = 1 (1) is an auta-pomorphy of Archipsocidae27 Forewing R1ndashR2+3 cross vein (0) absent (Fig 39)

Figures 35ndash40 Forendash (left) and hindwing (right) of Pararchipsocus pacificus (35) Lichenomima muscosa (36) Philotarsusquercicola (37) Ectopsocus sp (38) Triplocania spinosa (39) and Stenopsocus sp (40) Abbreviations ap = areola postica ps= pterostigma



(1) present (Fig 40) L = 1 CI = 1 RI = 1 (1) is anautapomorphy of Stenopsocideae28 Forewing Rs and M (0) fused (Fig 40) (1) con-nected by crossvein (Fig 39) L = 1 CI = 1 RI = 1 (1)is regarded as the derived condition and observed inEpipsocetae only29 Areola postica (0) present (Fig 37) (1) absent(Fig 38) L = 3 CI = 033 RI = 033 (1) was derivedindependently at least three times in EctopsocidaePeripsocidae and Notiopsocus Remarks Ectopso-cidae Peripsocidae and Notiopsocus (Asiopsocidae)had once been classified in a single family Peripso-cidae based only on the absence of the areola postica(Smithers 1972) The present analysis clearly showsthat absence of the areola postica does not indicatephylogenetic affinities between families30 Forewing veins CuA1 and M (0) free from eachother (Fig 37) (1) connected by crossvein (Fig 40)(2) fused with each other (Fig 36) L = 5 CI = 040RI = 050 (1) was derived independently at least threetimes in Hemipsocidae Stenopsocidae and Kod-amaius (Amphipsocidae sl) (2) is observed in Psoce-tae only but the character state of their commonancestor is unresolved Remarks Mockford (1996)reported a fossil Myopsocidae having the areola pos-tica separated from M If the venation of the fossilspecimen represents the ancestral condition of theMyopsocidae (0) should be assigned for the commonancestor of Psocetae31 Forewing vein A2 (0) present (Fig 39) (1) absent(Fig 40) L = 3 CI = 33 RI = 078 (1) is regarded asthe plesiomorphic condition of Psocomorpha and (0)as an autapomorphy of Epipsocetae but reversed to (1)in Epipsocidae Remarks Epipsocetae has beenregarded as one of the most primitive groups of Pso-comorpha mainly because of the presence of A2 inmany families of this group (Smithers 1972) Thepresent analysis suggests that its presence is the apo-morphic condition within Psocomorpha32 Ventral setae of forewing (0) absent (1) presentL = 5 CI = 020 RI = 069 (1) is regarded as an apo-morphic condition but highly homoplastic33 Nodus (0) absent (1) present (Fig 38) L = 1CI = 1 RI = 1 (1) is an autapomorphy of Psocomorpha34 Nodulus (0) a set of pointed separated spines ortruncated spines set closely together (Fig 33) (1) ahook formed by of truncated spines fused at their base(Fig 34) L = 1 CI = 1 RI = 1 (1) is an autapomorphyof Psocomorpha as indicated by Mockford (1967)35 Marginal setae of hindwing (0) setose all around(Fig 39) (1) setose between R2+3 and R4+5 only(Fig 40) (2) bare (Fig 36) L = 10 CI = 020 RI = 058(1) and (2) are regarded as derived conditions buthighly homoplastic Remarks Badonnel amp Lienhard(1988) regarded (1) as a synapomorphy of Elipsocidaeand Mesopsocidae However the present analysis

shows that (1) is highly homoplastic and provides littlephylogenetic information36 Setae on hindwing veins (0) none (Fig 36) (1) onerow (Fig 37) (2) two rows (Fig 39) L = 4 CI = 050RI = 086 (1) is regarded as an autapomorphy ofPhilotarsidae and Pseudocaeciliidae + Calopsocidaerespectively (2) is regarded as an autapomorphy ofEpipsocetae and Amphipsocinae respectively37 Hindwing Rs and M (0) fused (Fig 37) (1) con-nected by crossvein (Fig 36) L = 2 CI = 050 RI = 0(1) is an autapomorphy of the family Ectopsocidae andwas also derived independently in some genera ofMyopsocidae38 Hindwing Rs and M + Cu (0) fused with eachother basally (Fig 38) (1) separated from each otherbasally (Fig 37) L = 3 CI = 033 RI = 033 (1) isregarded as the apomorphic condition and observed inTrichopsocidae Philotarsidae and Idatenopsocus(Mesopsocidae)39 Ctenidia of hind tibia (0) fine (Fig 24) (1) broad(Fig 25) L = 1 CI = 1 RI = 1 (1) is regarded as anautapomorphy of Psocetae40 Tarsus (0) three-segmented (1) two-segmentedL = 7 CI = 013 RI = 059 (1) is regarded as an auta-pomorphy of Psocomorpha but this character is highlyhomoplastic within the suborder Remarks an earlyhigher classification proposed by Enderlein (1903)attached great importance to the number of tarsom-eres and psocids were first divided into two groupsbased on this feature The phylogenetic relationshipproposed by Smithers (1972) also attached importanceto this character and Psocomorpha was divided into alsquo2-segment tarsi linersquo and lsquo3-segment tarsi linersquo nearthe base of his cladogram The present analysis clearlyshows that this character is highly homoplastic andnot worthy for the estimation of phylogenetic relation-ships in Psocomorpha41 Pulvillus (0) fine (Fig 26) (1) broad (Fig 27) (2)absent L = 9 CI = 022 RI = 068 (1) is regarded asthe most plesiomorphic condition of Psocomorpha butthis character is highly homoplastic42 Preapical tooth on pretarsal claw (0) present(Fig 26) (1) absent (Fig 27) L = 5 CI = 020RI = 075 (1) is regarded as apomorphic but highlyhomoplastic43 Abdominal eversible vesicles (0) absent (1)present L = 3 CI = 033 RI = 082 (1) is an autapo-morphy of Pseudocaecilioidea (= Trichopsocidae +Calopsocidae + Pseudocaeciliidae) and Caeciliusoidearespectively and a reversal has occurred in a genus ofPseudocaeciliidae Remarks the presence of abdomi-nal eversible vesicles is often associated with living-foliage dwellers (New 1987) and a change in habitatpreference likely occurred at the basal node ofPseudocaecilioidea and Caeciliuseoidea respectivelyThe result of the present analysis also supports this

382 K YOSHIZAWA


idea concerning the evolution of this habitat-relatedcharacter44 Male paraproct (0) rounded (Fig 41) (1) with dis-tal process (Fig 42) L = 2 CI = 050 RI = 080 (1) isapomorphic and observed in Hemipsocidae Psilipso-cidae Myopsocidae and Psocidae Remarks (1) wasnot considered synapomorphic for Hemipsocidae andPsilipsocidae + Myopsocidae + Psocidae contrary toMockford (1993) As Hemipsocidae and Psocetae (inthe present sense) branched successively most parsi-monious optimization cannot establish whether (1) ishomologous or homoplastic I think the latter is morelikely because similar paraproctal processes are alsoobserved in some species of Lachesillidae which areapparently distantly related to Hemipsocidae andPsocetae45 Hypandrium (0) fused with clunium laterally(Fig 41) (1) articulated with clunium laterally(Fig 42) L = 3 CI = 033 RI = 067 (1) is regardedas an autapomorphy of each Psocidae andPseudocaeciliidae + Calopsocidae and independentlyderived in Philotarsus Remarks when the hypan-drium is articulated with the clunium its posteriormargin always possesses processes or lobes whereaswhen it is fused with the clunium the distal margin issmoothly rounded Therefore presence of an articu-lation appears to be strongly correlated with the pres-ence of such processes or lobes Betz (1983) studied thegenital coupling mechanism of Trichadenotecnumalexanderae and reported that the hypandrial distalprocesses and the female gonopore plates are inter-locked with each other during coupling The articula-tion between hypandrium and clunium probablyfacilitates hypandrial movement and this morpholo-gical change may have enabled the evolution of thecomplicated hypandrial distal processes that lead toan interlocking mating system46 Hypandrial lateral bristles (0) absent (Fig 41)(1) present (Fig 43) L = 1 CI = 1 RI = 1 (1) isregarded as an autapomorphy of cladePseudocaeciliidae + Calopsocidae47 Pair of lateral hypandrial processes (0) absent(Fig 41) (1) present (Fig 43) L = 2 CI = 050RI = 067 (1) is regarded as an autapomorphy of cladePseudocaeciliidae + Calopsocidae and independentlyderived in Ptiloneuridae48 Apex of aedeagus (0) pointed (Fig 46) (1) rounded(Fig 47) L = 4 CI = 025 RI = 057 (1) is regarded asthe apomorphic condition and observed in Archipso-cidae Psilopsocidae Philotarsidae and Mesopsocus(Mesopsocidae) Remarks this character is scored asunknown for exemplars which lack the aedeagus49 Aedeagus (0) present (Fig 46) (1) absent(Fig 44) L = 6 CI = 017 RI = 029 (1) is regarded asa derived condition but highly homoplastic50 Paramere (0) rod-like (Fig 47) (1) strap-like

(Fig 48) (2) reduced (Fig 45) L = 4 CI = 050RI = 071 (1) is regarded as an autapomorphy ofPeripsocidae and (2) as an autapomorphy of Archip-socidae and Psocidae + Psilopsocidae51 Endophallus (0) without rod-like sclerites(Fig 47) (1) with rod-like sclerites (Fig 46) L = 5CI = 020 RI = 060 (1) is apomorphic supporting theclade containing homilopsocid families except Elipso-cidae These sclerites have been secondarily reducedin Lachesilla Pseudocaeciliius Mesopsocidae andsome genera of Philotarsidae52 Female epiproct (0) rounded (Fig 49) (1) square-shaped (Fig 50) L = 1 CI = 1 RI = 1 (1) is an auta-pomorphy of the clade containg PeripsocidaePhilotarsidae and Mesopsocidae53 Female paraproct (0) rounded (Fig 51) (1) withconical distal projection (Fig 52) L = 2 CI = 050RI = 080 (1) is apomorphic and observed in Hemip-socidae and Psocetae Remarks Hemipsocidae hasbeen assigned to Psocetae on the basis of (1) by Mock-ford (1976) In the present analysis Hemipsocidae andother families of Psocetae do not comprise a monophyl-etic group and Hemipsocidae and Psocetae are posi-tioned on successive branches most parsimoniousoptimization cannot therefore nestablish whether (1)is homologous or homoplastic54 Apex of dorsal valve of gonapophyses (0) notstrongly extended posteriorly (Fig 55) (1) stronglyextended posteriorly to reach posterior margin ofparaproct (Fig 54) L = 1 CI = 1 RI = 1 (1) is anautapomorphy of Myopsocidae55 Dorsal extension of subgenital plate (0) absent(Fig 53) (1) weakly projected pair of tubercles bearinga few apical setae (Fig 56) (2) well developed bi-lobed(Figs 57 58) (3) well developed single-lobed (Fig 59)L = 10 CI = 030 RI = 079 (1) is an autapomorphy ofHomilopsocidea but reversed in Trichopsocidae andsome lachesillid taxa Remarks (1ndash3) are only presentin Homilopsocidea The egg guide of Elipsocidae(Fig 56) comprises a pair of small tubercles bearingapical setae (state 1) whereas those of EctopsocidaePseudocaeciliidae and Calopsocidae are well projectedposteriorly bilobed apically with each lobe bearing api-cal setae (state 2) (Figs 57 58) The structure of the eggguidersquos apex in (2) is basically identical to that in (1)and is therefore regarded as derived from it morpho-logically The egg guide of Peripsocidae Philotarsidaeand Mesopsocidae is well projected posteriorly single-lobed and usually lacks apical setae (state 3) (Fig 59)The egg guide of some peripsocids is single-lobed butslightly hollowed apically and somewhat intermediatebetween states 2 and 3 Therefore a single-lobed eggguide is considered as being derived from state 2 Tosummarize a transformation series of the egg guidefrom states 0 to 3 can be hypothesized By optimizingthis character as ordered on the obtained cladogram



(1) was considered to be an additional autapo-morphy of Homilopsocidea and (2) an autapomorphyof Peripsocoidea (= Bryopsocidae + Peripsocidae +Philotarsidae + Mesopsocidae)56 Ventral extension of subgenital plate (0) absent(Fig 53) (1) present (Figs 52 55) L = 1 CI = 1 RI = 1(1) is an autapomorphy of Psocetae Remarks exter-nally (1) is similar to (3) of character 55 Howeverthey are recognized as nonhomologous In Psocetaethe egg guide is regarded as an extension of the ven-trodistal margin of the subgenital plate and the ven-tral margin of the subgenital plate (including the eggguide) is straight or smoothly arched in lateral aspect(Fig 52) In contrast the egg guide of Homilopsocideais regarded as an extension of the dorsodistal marginof the subgenital plate the ventral margin of which isstrongly dorsally bent at the base of the egg guide inlateral aspect (Fig 51)57 Spermathecal sac (0) without lateral pouch (1)with lateral pouch at junction of sac and duct (Fig 60)L = 1 CI = 1 RI = 1 (1) is an autapomorphy ofStenopsocidae

58 Gonapophyses (0) dorsal and ventral valves do notform the ovipositor (Fig 54) (1) dorsal and ventralvalves form the ovipositor together with the egg guideof the subgenital plate (Fig 55) L = 3 CI = 033RI = 087 (1) is regarded as an apomorphic conditionand derived independently twice within PsocomorphaThe ovipositor has also evolved at least once in theoutgroup but lacks the egg guide of the subgenitalplate59 Ventral valve of gonapophyses (0) present(Fig 67) (1) absent (Fig 68) L = 4 CI = 025RI = 067 (1) is apomorphic and derived indepen-dently four times in Psocomorpha60 Apex of ventral valve of gonapophyses (0) tapered(Fig 61) (1) with dorsal lobe (Fig 66) (2) swollen(Fig 67) L = 2 CI = 1 RI = 1 (1) is an autapomorphyof Pseudocaeciliidae + Calopscidae and (2) is an auta-pomorphy of Mesopsocidae Remarks some exemplarslacked the ventral valve of the gonapophyles and werescored lsquorsquo61 Dorsal valve of gonapophyses (0) well developed(Fig 67) (1) reduced to membranous lobe (Fig 68) (2)

Figures 41ndash48 (41ndash43) male genitalia of Idatenopsocus orientalis (41) and Trichadenotecnum sexpunctatum (42) lateralview and Heterocaecilius fuscus (43) posterior view (44ndash48) phallosome of Hemipsocus chroloticus (44) Psocus sp (45)Het fuscus (46) Haplophallus sp (47) and Peripsocus quercicola (48) Abbreviations a = aedeagus cl = clunium ep =endophallus hy = hypandrium pp = paraproct pr = paramere

384 K YOSHIZAWA


absent L = 4 CI = 050 RI = 050 (0) is regarded asthe most primitive condition (1) is an autapomorphyof Lachesillidae (excluding Eolachesilla) and Asiopso-cidae respectively (2) was derived from (1) in somespecies of Lachesilla and from (0) in Archipsocus62 Dorsal region of dorsal valve of gonapophyses(0) not swollen (Fig 61) (1) swollen (Fig 66) L = 1CI = 1 RI = 1 (1) is an autapomorphy of Homilopso-cidea Remarks See next character63 Ventral region of dorsal valve of gonapophyses (0)without swelling (Fig 61) (1) with swelling (Fig 62)L = 1 CI = 1 RI = 1 (1) is an autapomorphy of Psoce-tae Remarks the gonapophyses of Homilopsocideaand Psocetae both have a broad dorsal valve and aresomewhat similar to each other However they can beclearly distinguished by comparing them from the

lateral aspect (Figs 62 66 67) In Psocidae the distalprocess of the dorsal valve is projected from the dorsalpart of the valve (Fig 62) whereas it is projected fromthe ventral part in Homilopsocidea (Figs 66 67) Thisillustrates the nonhomologous condition of characters62 and 63 In Lachesillidae dorsal and ventral valvesof the gonapophyses are reduced and thus characters62 and 63 are scored as unknown64 Dorsal swelling of dorsal valve of gonapophyses(0) membranous (Fig 65) (1) sclerotized (Fig 66)L = 1 CI = 1 RI = 1 (1) is a synapomorphy of Trichop-socidae Pseudocaeciliidae Calopsocidae Bryopso-cidae Peripsocidae Philotarsidae and Mesopsocidae65 Dorsal swelling of dorsal valve of gonapophyses(0) lobe-like (Fig 65) (1) forming rounded plate(Fig 66) (2) forming square-shaped plate (Fig 67)

Figures 49ndash60 (49ndash50) female epiproct of Elipsocus abdominalis (49) and Haplophallus sp (50) dorsal view (51ndash55)female genitalia of E abdominalis (51) Psocidus sp (52) and Stenopsocus sp (53) lateral view Lichenomima muscosa (54)and P sp (55) ventral view (56ndash59) apex of subgenital plate of E abdominalis (56) Ectopsocus sp (57) Heterocaecilius fus-cus (58) and Mesopsocus unipunctatus (59) (60) spermatheca of Stenopsocus sp Abbreviations epr = epiproct dv = dorsalvalve ev = external valve pp = paraproct sg = subgenital plate vv = ventral valve



L = 2 CI = 1 RI = 1 (1) is uniquely observed inPseudocaecilioidea (= Trichopsocidae + Calopsocidae +Pseudocaeciliidae) and (2) is observed in Peripsocoidea(= Bryopsocidae + Peripsocidae + Philotarsidae +Mesopsocidae) Remarks these superfamilies com-prise a monophyletic group and thus the state of thischaracter in their common ancestor is uncertain Incomparison with Elipsocidae (1) could be regarded asmore primitive than (2) By treating this character asordered (1) can be assigned to their common ancestor(1) thus provides further evidence for the monophylyof Pseudocaecilioidea + Peripsocoidea and (2) isregarded as an autapomorphy of Peripsocoidea66 External valve of gonapophyses (0) broad(Fig 62) (1) narrowed (Fig 63) (2) reduced (Fig 64)L = 3 CI = 067 RI = 095 (1) is regarded as anautapomorphy of Lachesilloidea (= Ectopsocidae +Lachesillidae) and also observed in Epipsocetae (2) isobserved in Caeciliusetae Remarks Epipsocetae andCaeciliusetae comprise a clade and thus the state ofthis character in their common ancestor is ambiguousMorphologically the narrowed external valve can beregarded as an intermediate condition between broadand reduced external valves By optimizing this char-acter as ordered on the resultant cladogram (1) can beassigned to the basal node of Epipsocetae + Caeciliuse-tae (1) thus provides additional support for the

Epipsocetae + Caeciliusetae clade and (2) is con-sidered as an autapomorphy of Caeciliusetae67 Dorsal and external valves of gonapophyses (0)separated (Fig 66) (1) partly fused (Fig 63) (2) com-pletely united L = 2 CI = 1 RI = 1 (1) is an autapo-morphy of Epipsocetae and (2) is considered as derivedfrom (1) supporting a clade containing EpipsocidaeDolabellopsocidae and Cladiopsocus68 Posterior lobe of external valve (0) absent(Fig 54) (1) present (Fig 55) L = 1 CI = 1 RI = 1 (1)is an autapomorphy of the family Psocidae

RESULTS

The cladistic analysis including all exemplars yielded1108 equally most parsimonious trees (L = 191) Astrict consensus of the trees (Fig 69) shows that phy-logenetic relationships near the basal node are unre-solved Exemplars include two unexamined species(Bryopsocus townsendi and Asiopsocus sonorensisFig 69 arrows) and the poor resolution is possiblycaused by the large number of unknown charactersscored in the matrix for these unexamined species(Platnick et al 1991 Novacek 1992 Kitching et al1998) Judging from the consensus the phylogeneticposition of Asiopsocus is stable (Fig 69) and thusthe exemplar used does not contribute to the poor

Figures 61ndash68 Gonapophyses of Hemipsocus chloroticus (61) Psocus sp (62) Triplocania spinosa (63) Amphipsocusmangifera (64) Elipsocus abdominalis (65) Heterocaecilius fuscus (66) Mesopsocus unipunctatus (67) and Lachesilla sp(68) ventral view Abbreviations dv = dorsal valve ev = external valve vv = ventral valve

386 K YOSHIZAWA


Figures 69ndash73 (69) generated from original data set strict consensus of 1108 trees (70ndash72) generated from new data setstrict consensus of (70) 72 trees of set 1 (71) 56 trees of set 2 (72) two trees generated using successive approximations andimplied weights techniques family names are abbreviated to first three letters (73) 14 trees generated from original dataset using successive approximations and implied weights techniques (part)

HO

MILO

PS

OC

IDE

A



resolution Although Asiopsocus contains manyunknown characters the exemplar can be consideredto have enough information to decide its phylogeneticposition The lack of resolution and large number ofequal length trees may be due therefore to the influ-ence of Bryopsocus To determine the extent of thisinfluence a new data set excluding Bryopsocus wasprepared and reanalysed

The revised data set reduced the number of equallymost parsimonious trees to 128 the topology of thestrict consensus tree does not differ from that of theoriginal data set although Ectopsocidae and Lachesil-lidae now constitute a monophyletic group Trees fromthe new data set are assigned in two sets of trees ofequal length (= islands sensu Maddison 1991) con-taining 72 (set 1 Fig 70) and 56 trees (set 2 Fig 71)Set 2 differs from set 1 in making Psocetae and Hom-ilopsocidea sister groups (Fig 71) This relationship issupported by only one extremely homoplastic charac-ter state a three-segmented tarsus Alternatively thestrict consensus tree of set 1 (Fig 70) supports a cladecontaining Homilopsocidea Epipsocetae and Caecil-iusetae using a reliable nonhomoplastic character themesothoracic dorso-ventral flight muscle Trees in set1 are therefore regarded as more stable than those inset 2

As discussed above poor resolution may beexplained by a few highly homoplastic charactersThus in order to assign a heavier weight to reliablecharacters the characters were weighted using thesuccessive approximations technique based on themaximum value of RC calculated from 128 treesinferred from the new data set (Farris 1969Carpenter 1988) The implied weights technique(Goloboff 1993) was also performed with k = 2 and 10(different k-values do not alter the result) Both analy-ses yielded two equally parsimonious trees Thecladograms produced are identical with two of theoriginal 72 cladograms in set 1 and are thus regardedas stable Topologies of these two trees differ only inthe relationships of outgroups

The techniques were also applied to the originaldata set reducing the number of cladograms to 14These trees are compatible with the two cladogramsestimated from the new data set Bryopsocus wasplaced as sister group to the clade containing Peripso-cidae Philotarsidae and Mesopsocidae (Fig 73)

Within the 14 trees phylogenetic relationshipsamong Peripsocidae Philotarsidae and Mesopsocidaeare unstable although a sister-group relationshipbetween Philotarsidae and Mesopsocidae is morelikely Badonnel amp Lienhard (1988) produced a phy-logeny of Mesopsocidae and regarded state 0 ofcharacters 38 and 48 observed in Mesopsocus as auta-pomorphies of the genus In the present analysis thestates of these characters for the common ancestor of

Mesopsocidae could not be decided because the twoexemplars have different states for them According toBadonnel amp Lienhard (1988) the basalmost taxon ofthe family has state 1 for both characters This can beregarded as a ground plan for Mesopsocidae and thusthey can be considered as synapomorphic for Philotar-sidae and Mesopsocidae

Polytomy is preferred for some nodes (Fig 74arrows) because of lack of information to resolve theirrelationships Different interpretations of outgrouprelationships do not alter the ingroup topology There-fore among 14 equal-length trees one (Fig 74L = 191 CI = 043 RI = 081) was selected as provid-ing the best estimation of the phylogenetic relation-ships of the psocomorphan families Based on themonophyletic groups of families identified by the cla-dogram (Fig 75) the family and higher level classifi-cation of Psocomorpha is revised (Table 1)

DISCUSSION

MONOPHYLY OF PSOCOMORPHA AND RELATIONSHIPS OF INFRAORDERS AND SUPERFAMILIES

Mockford (1967) proposed three autapomorphies sup-porting the monophyly of suborder Psocomorpha ahooked nodulus formed by truncated spines fused attheir base (Fig 34) thickened pterostigma enlarge-ment of mesothorax (Fig 17) The present analysisstrongly supports this monophyly However the thirdis not regarded as autapomorphic at the suborderlevel The following additional characters were alsofound to support monophyly rounded vertex (Fig 2)absence of stipito-galeal muscle (Fig 11) presence ofproximal minute process at apex of first axillary scler-ite (Figs 29 30) presence of nodus (Fig 38) By mostparsimonious optimization of the character states twoadditional character states are regarded as autapo-morphies of Psocomorpha two-segmented tarsus andbroad pulvillus However these characters are quitevariable within the suborder

Six monophyletic infraorders are recognized in thepresent analysis (Table 1) four of which have beenwidely accepted since Pearman (1936) first establishedthe higher taxonomic system Two new infraordersArchipsocetae and Hemipsocetae are proposed Bothare represented by only one family which was formerlyincluded in Homilopsocidea and Psocetae respectively(Table 1)

Archipsocidae is regarded as the basalmost clade ofPsocomorpha (Fig 75) It has long been assigned toHomilopsocidea However it possesses the followingplesiomorphic character states that exclude it fromthis infraorder prothorax strongly bulged dorsallypterothorax weakly bulged dorsally narrow membra-nous region of metaepisternum posteroproximal

388 K YOSHIZAWA


corner of forewing angled proximal median platemobilized not fused with second axillary sclerite Onlyone derived character state broad precoxal bridge ofmesothorax (Fig 23) is shared between Archipsocidaeand families of Homilopsocidea and possibly indicatescloser relationships between them However the mus-culature associated with the precoxal bridge is com-pletely different (Figs 20 21) and similarity of theprecoxal bridge should be regarded as convergence

A narrow internal ridge of the epistomal suture anda strongly bulged pterothorax support the monophylyof Psocomorpha excluding Archipsocidae The formercharacter is reversed in some families of the suborder(Elipsocidae Mesopsocidae and all families ofCaeciliusoidea) Among the families of Psocomorpha(excluding Archipsocidae) Hemipsocidae is placed atthe basalmost clade It was regarded as a member ofPsocetae based on the following character states maleparaproct with distal process female paraproct withdistal cylindrical projection (Fig 54) narrow mesotho-racic precoxal bridge (Fig 22) Although the third is

plesiomorphic the first two are apomorphic and pos-sibly indicate a close relationship between Hemipso-cidae and Psocetae However as discussed in the nextparagraph monophyly of the clade containing Psoce-tae (excluding Hemipsocidae) Homilopsocidea Epip-socetae and Caeciliusetae is well supported by stableor nonhomoplastic autapomorphies Consequently thesimilarities observed between Hemipsocidae and Pso-cetae are regarded as symplesiomorphies or homopla-sies (ambiguous although the latter is more likelybecause similar structures are independently derivedin some other taxa such as some species of Lachesilla)

The remaining infraorders minus Psocetae Homilopso-cidea Epipsocetae and Caeciliusetae minus comprise amonophyletic group and autapomorphies supportingthis clade include (1) the broad membranous region ofthe metaepisternum (Fig 17) (2) fusion of the proxi-mal median plate to the second axillary sclerite(Fig 32) (3) posteroproximal margins of folded forew-ings separated from each other (Fig 32) These threecharacter states are hypothesized to have been

Figure 74 Preferred cladogram of Psocomorpha and most parsimonious reconstruction of character states Outgroups areomitted



derived only once and no convergence or reversalswere detected by the present analysis This is espe-cially significant where (2) is a functionally veryimportant character state for wing folding and

regarded as a reliable autapomorphy strongly sup-porting the monophyly of Psocetae + Homilopsocidea+ Epipsocetae + Caeciliusetae

Homilopsocidea + Epipsocetae + Caeciliusetae is

Figure 75 Phylogenetic relationships among families of Psocomorpha Monophyly of shaded bar is not supported

390 K YOSHIZAWA


supported by only one autapomorphy divided dors-oventral flight muscle of the mesothorax (Fig 20) InPsocetae Hemipsocetae and outgroups one dorsoven-tral flight muscle is inserted into the base of thetrochantin (Fig 19) In contrast dorsoventral flightmuscles of Homilopsocidea Epipsocetae and Caecil-iusetae are divided into two one of which is insertedinto the precoxal bridge and the other into the tro-chantin (Fig 20) As already discussed above only onehighly homoplastic character may possibly contradictthis clade

Monophyly of Caeciliusetae + Epipsocetae is sup-ported by an elongate and posteriorly hollowed man-dible (Fig 7) ball-shaped galea (Fig 9) and narrowedor reduced external valve of the gonapophyses Asomewhat triangular labial palpus is also widelyobserved possibly providing further evidence ofmonophyly Since most synapomorphies supportingCaeciliusetae + Epipsocetae are presumably stronglyassociated with feeding behaviour similar structuresmight easily have occurred independently Howeverthe members of Caeciliusetae are mostly living-foliagedwellers whereas those of Epipsocetae are found inleaf litter or on stone or bark surfaces Their foodsources are considered to be different making it diffi-cult to establish that the similarities of mouthpartsare convergences associated with function Conse-quently similarities of mouthparts observed betweenCaeciliusetae and Epipsocetae are regarded here assynapomorphic

Infraorder ArchipsocetaeArchipsocidae is the only representative of thisinfraorder distributed in tropical regions Monophylyis strongly supported by the following autapomor-phies broad precoxal bridge and related unique mus-culature (Fig 21) forewing veins and membranecovered with long setae (Fig 35) forewing marginalcrossing setae (Fig 35) reduction of venation(Fig 35) absence of preapical tooth on pretarsal claw(Fig 27) lack of parameres absence of ventral valveof gonapophyses (Fig 68) Monophyly is also well sup-ported by their behaviour All known species of thisfamily live on bark or dead leaves beneath webbingsheets in groups and subsociality is also known (New1987) External appearance is very similar

Infraorder HemipsocetaeThe family Hemipsocidae is the only representative ofthis infraorder It is a small family recorded fromall zoogeographical regions but its distribution isrestricted to tropical or warm areas All species aredead-foliage dwellers Monophyly is strongly sup-ported by absence of the epistomal suture apically

broadened first axillary sclerite two-branched M vein(not used for analysis) and CuA1ndashM crossvein(Fig 40) Although absence of aedeagus (Fig 44) doesnot unambiguously support this clade this characterstate is possibly an additional autapomorphy

Infraorder PsocetaeIncludes three families Psilopsocidae Psocidae andMyopsocidae all of which are bark or stone surfacedwellers Monophyly is well supported by thectenidia-based broad setae on the hind tibiae (Fig 25)bare fore- and hindwings (Fig 36) and single-lobedegg guide extended from the ventral margin of thesubgenital plate (Figs 52 55)

Mockford (1961) assigned Psilopsocidae one of theleast known and controversial families to PsocetaePsilopsocidae shares the above-mentioned apomorphiccharacter states with Psocidae and Myopsocidae andcan apparently be included in this infraorder

The areola postica of Psilopsocidae is always sepa-rate from the M vein whereas it is connected with it inPsocidae and Myopsocidae (Fig 36) The latter state isapomorphic and possibly supports the monophyly ofPsocidae + Myopsocidae However a fossil species ofMyopsocidae in which the areola postica is separate isknown (Mockford 1996) if the venation of the fossilspecimen represents the ancestral condition mono-phyly is not justified However three apomorphiccharacter states minus paramere absent (Fig 45) dorsalvalve of gonapophyses with ventral swelling (Fig 62)gonapophyses and egg guide forming the ovipositor minusare shared by Psocidae and Psilopsocidae and supporttheir sister-group relationship

Psilopsocidae contains only one genus and sevennamed species Smithers (1995a b 1997) reporteda unique and characteristic nymphal wood-boringbehaviour in final instar nymphs of Psilopsocus mim-ulus In addition all psilopsocid species of which thenymphal stage is known possess a more or less scle-rotized abdominal tip and thus all are probably wood-borers (Smithers 1995a) If this behaviour is widelyobserved throughout the family monophyly is stronglysupported by this highly derived behaviour andassociated morphological characters In the presentanalysis a rounded aedeagus was considered to bean autapomorphy of this family

Psocidae is one of the largest families of Psocopteradistributed in all zoogeographical regions Monophylyis supported by the following autapomorphic characterstates presence of an articulation between hypan-drium and clunium (Fig 42) presence of the posteriorlobe of the external valve of the gonapophyses(Fig 55) Position of anterior tentorial pit and shape ofpulvillus may possibly support the monophyly butthey are highly homoplastic



Myopsocidae is distributed in all zoogeographicalregions The very long dorsal valve of the gonapophy-ses (Fig 54) supports monophyly Although forewingmarkings were not used for this analysis those of thisfamily are unique and characteristic (Fig 36) andpossibly provide further evidence supporting mono-phyly Male genital characters show great diversityamong genera and as mentioned by Mockford (1961)male genital structures of some myopsocid genera aresimilar to those of Psilopsocidae and may possiblysupport their sister-group relationship To confirmMockfordrsquos findings a transformation series ofmale genital characters within the family must beundertaken

Infraorder HomilopsocideaThis infraorder contains ten families assigned to foursuperfamilies When Pearman (1936) first proposedHomilopsocidea he expressed some doubts about itssoundness Badonnel (1951) also found the infraorderto be heterogeneous Mockford (1976) transferredHemipsocidae to Psocetae Later Mockford (1993)stated that ldquowith the hemipsocids excluded it seemsto consist of a series of related families except that theperipsocid stand apart from the others especially instructure of male genitaliardquo (Fig 48)

The present cladistic analysis shows that thisinfraorder comprises related families except Archipso-cidae which is here assigned to its own infraorder andstands apart from Homilopsocidea Peripsocidae ishere assigned to Homilopsocidea the specialized malephallosome of the family is apparently an autapomor-phic modification and thus provides no informationabout its phylogenetic position

Monophyly is supported by the following twoapomorphic character states presence of egg guideextending from dorsodistal margin of the subgenitalplate (Fig 51) and the dorsal swelling of the dorsalvalve of the gonapophyses (Figs 65minus67)

Elipsocidae is regarded here as the sister groupof all remaining homilopsocid families Badonnel ampLienhard (1988) suggested a sister-group relationshipbetween Elipsocidae and Mesopsocidae based on thereduction of wing setae Specifically hindwing mar-gins of both families are bare except between R2+3 andR4+5 (Fig 40) This condition is apomorphic but highlyhomoplastic and similar conditions are observed alsoin Stenopsocidae and Dasydemellidae (Caeciliusetae)Moreover when the entire hindwing margin is setosemarginal setae between R2+3 and R4+5 are usuallystronger than those in other sections (eg Trichopso-cidae) Forewing ciliation of Elipsocidae is plesiomor-phic Monophyly of the clade containing the remaininghomilopsocid families (including Mesopsocidae) issupported by an apomorphic character state which is

more reliable than wing ciliation presence of rod-likesclerites on the endophallus (Fig 46) Thus the simi-lar condition of hindwing marginal setae observed inElipsocidae and Mesopsocidae should be regarded ashomoplasy

The remaining families are divided into two cladesLachesilloidea and Trichopsocidae + Pseudocaeciliidae+ Calopsocidae + Bryopsocidae + Peripsocidae +Philotarsidae + Mesopsocidae Monophyly of the latteris supported by a sclerotized dorsal valve and ventraland dorsal valves of the gonapophyses tightly unitedto form the ovipositor (Fig 55) These character statesare unique and observed throughout all taxa withinthe clade The latter clade also comprises two mono-phyletic groups Pseudocaecilioidea and Peripso-coidea Monophyly of each superfamily is discussedbelow

Superfamily ElipsocoideaThe family Elipsocidae is the only representative ofthis superfamily distributed in all zoogeographicalregions and all members are bark or stone surfacedwellers

Elipsocidae mostly retains the plesiomorphic condi-tions of the Homilopsocidea Monophyly is supportedby three character states broad internal ridge of theepistomal suture (Fig 3) hindwing marginal setaerestricted between R2+3 and R4+5 and three-segmentedtarsus The latter character state is extremelyhomoplastic Male and female genital structures arevery similar throughout Elipsocidae but some are ple-siomorphic and the polarity of others remains uncer-tain Further study is therefore required to confirmthe monophyly

Superfamily LachesilloideaMonophyly is supported by three apomorphic charac-ter states elongate preepisternum of prothorax(Fig 16) bare hindwing margin and narrowed exter-nal valve of gonapophyses (Fig 68) The systematicposition assigned for Eolachesilla by previous authorshas been controversial Lachesillidae (Badonnel 1967Mockford amp Sullivan 1986) or Elipsocidae (New ampThornton 1981) The present analysis suggests acloser relationship between the genus and Lachesil-lidae rather than Elipsocidae

Ectopsocidae is distributed in all zoogeographicalregions Monophyly is strongly supported by the fol-lowing autapomorphies absence of areola postica inforewing (Fig 38) hindwing Rs and M connected bycrossvein (Fig 38) presence of ball-shaped lobe onmeta-epimeron (not used in the analysis) absence ofpreapical tooth of pretarsal craws

392 K YOSHIZAWA


Lachesillidae is also distributed in all zoogeogra-phical regions Monophyly of the family excludingEolachesilla is strongly supported by the position ofanterior tentorial pit the absence of the ventral valveof the gonapophyses and reduction of the dorsal valveof the gonapophyses (Fig 68) Eolachesilla lacks theseapomorphic character states and monophyly includingEolachesilla is therefore uncertain In this paper thisgenus is declared incertae sedis and it should be notedthat Eolachesilla may represent its own family

Superfamily PseudocaecilioideaMonophyly of Trichopsocidae + Pseudocaeciliidae+ Calopsocidae is supported by apically broadenedfirst axillary sclerite (Fig 15) position of the anteriortentorial pit forewing veins with two rows of setaeand absence of preapical tooth The latter three char-acters are highly homoplastic Presence of eversiblevesicles on the abdomen may also provide further sup-port for this superfamily although the state of thischaracter is ambiguous at the basal node of the cladeThe male phallosome and female gonapophyses ofthese three families are similar but they are symple-siomorphic Trichopsocidae Pseudocaeciliidae andCalopsocidae are all living-foliage dwellers This fea-ture is apomorphic and provides further evidence forthe monophyly of the Pseudocaecilioidea

The present analysis posits that within Pseudo-caecilioidea Pseudocaeciliidae and Calopsocidaecomprise a monophyletic group Smithers (1967)suggested close affinity of these families but later(Thornton amp Smithers 1984) doubted this mentioningthat similarities between them were largely due totheir retention of primitive features the two synapo-morphies being bilobed subgenital plate and two-segmented tarsi Most of shared characters suggestedin the earlier paper are plesiomorphic or homoplasticincluding the two character states that Thornton ampSmithers (1984) suggested as apomorphic Monophylyis well supported by the following character statescrossing marginal setae of forewing (Fig 37) hind-wing veins with one row of setae presence of an artic-ulation between clunium and hypandrium (Fig 43)presence of hypandrial lateral bristles (Fig 43)ventral valve of gonapophyses with dorsal lobe(Fig 66)

Trichopsocidae is a small family containing onegenus and eight species distributed in all zoogeo-graphical regions except the Orient Monophyly issupported by secondary loss of the distal extension ofthe subgenital plate An additional apomorphic fea-ture hindwing Rs and M + Cu separate basally(Fig 37) is observed in one exemplar but it is uncer-tain whether this character state is widely observedthroughout this family

Pseudocaeciliidae is the largest family among Hom-ilopsocidea and is distributed in all zoogeographicalregions No autapomorphy supporting monophyly wasfound in the present study Thus the family isregarded here as a paraphyletic group Calopsocidae isregarded as a highly specialized clade within it Ideclare the family Pseudocaediliidae incertae sedisand further phylogenetic study of Pseudocaeciliidae+ Calopsocidae is required to confirm theirrelationships

Calopsocidae is known only from the Oriental andMelanesian regions Monophyly is well supported by asharply angled vertex that is deeply emarginatedmedially and presence of numerous secondary veinsin the forewing (not used in the analysis) A somewhatelongate mandible ball-shaped galea and presence ofa preapical tooth on the pretarsal claw also supportmonophyly

Superfamily PeripsocoideaMonophyly is supported by the following apomorphiccharacter states single-lobed egg guide (Fig 59)strongly sclerotized and square-shaped dorsal valve ofthe gonapophyses (Fig 67)

The family Bryopsocidae is represented by one spe-cies Bryopsocus townsendi (Smithers 1969) knownfrom New Zealand Smithers (1969) originallydescribed the species under the genus Austropsocus ofthe family Philotarsidae Thornton Wong amp Smithers(1977) subsequently erected a new genus Bryopsocusin the family Philotarsidae designating A townsendias type species of the genus Mockford (1984) exten-sively studied philotarsid and pseudocaeciliid generaand concluded that Bryopsocus is intermediatebetween Philotarsidae and Pseudocaeciliidae Thushe founded the monotypic family Bryopsocidae for thegenus and considered it to be the sister group toPseudocaeciliidae + Calopsocidae This conclusion wasbased on lsquoextent of character sharingrsquo (Mockford 1984)but he did not discuss their polarity

Judging from published descriptions and illustra-tions Bryopsocidae possesses all the apomorphiessupporting the monophyly of Homilopsocidea Onecharacter state dorsal extension of the subgenitalplate (character 55) is difficult to assess from pub-lished illustrations but a transverse line was clearlydrawn near the base of the egg guide (Smithers 1969fig 192) which most probably indicates that the eggguide extends from the dorsal surface Thus thefamily is regarded as a member of Homilopsocidea

Bryopsocidae also appears to share the apomorphiesof Peripsocoidea listed above These character statesare not observed in Pseudocaecilioidea However Bry-opsocidae has long rod-like sclerites on the endophal-lus that are similar to those of Pseudocaeciliidae and



Calopsocidae This may suggest closer affinities assuggested by Mockford (1984) However similar struc-tures are also observed in some taxa within Lachesil-loidea and Peripsocidae and consequently similarityof the endophallic sclerites observed in Bryopsocidaeand Pseudocaeciliidae should be regarded as a sym-plesiomorphy Thus Bryopsocidae is regarded as amember of this superfamily

Within the Peripsocoidea Bryopsocidae occupies thebasalmost clade Monophyly of Peripsocidae + Philo-tarsidae + Mesopsocidae is well supported by a square-shaped female epiproct (Fig 50) Bryopsocidae sharesthree-segmented tarsi with Philotarsidae and Mesop-socidae although this character state is highlyhomoplastic

In the present analysis phylogenetic relationshipsamong Peripsocidae Philotarsidae and Mesopsocidaewere not resolved because no decisive autapomorphysupporting the clade comprising two of the three fam-ilies was detected However as discussed above state1 of character 39 and state 1 of character 48 can beregarded as autapomorphies supporting the mono-phyly of Philotarsidae + Mesopsocidae

Peripsocidae is distributed in all zoogeographicalregions Members are mostly bark or stone surfacedwellers but some are collected from dead foliageMonophyly is well supported by absence of areola pos-tica in the forewing (Fig 38) and strap-like paramere(Fig 48) Because of the first character Peripsocidaewas once classified with Ectopsocidae but is only dis-tantly related to it

Philotarsidae is distributed in all zoogeographicalregions Members are all bark or stone surface dwell-ers Monophyly is supported by only one decisive apo-morphic character presence of ventral setae on theforewing Position of the anterior tentorial pit cross-ing marginal setae of forewing more than one row ofsetae on forewing veins and apically rounded aedea-gus possibly provide further evidence of monophylybut they are highly homoplastic or their states at thebasal node of the family are ambiguous

Mesopsocidae is widely distributed in all zoogeo-graphical regions and is particularly abundant in theAfrotropical and Palearctic regions Members aremostly bark or stone surface dwellers although onespecies is known to be associated with termites Mono-phyly is supported by the following apomorphic char-acter states broad internal ridge of the epistomalsuture glabrous fore- and hindwings apically broad-ened ventral valve of the gonapophyses (Fig 67)

Infraorder EpipsocetaeMonophyly is well supported by the following apomor-phic character states anterior tentorial pit separatedfrom ventral margin of cranium (Fig 5) labrum with

a pair of longitudinal sclerotized lines (Fig 5) forew-ing veins with more than one row of setae (Fig 39reversed in Epipsocidae) presence of A2 vein (Fig 39reversed in Epipsocidae) hindwing veins with tworows of setae (Fig 39) dorsal and external valves ofgonapophyses (partly) fused (Fig 63) A long gena andbroad lacinial tip possibly support monophyly butthese continuous quantitative characters were notused for the analysis

Spurostigma of the family Cladiopsocidae isregarded as the basalmost clade of the infraorder andmonophyly of Epipsocetae excluding Spurostigma issupported by one stable autapomorphy forewing veinsRs and M connected by a crossvein

Ptiloneuridae is regarded as the second basalmostclade of Epipsocetae The family is known from theAfrotropical Nearctic and Neotropical regions Mono-phyly is supported by three highly homoplasticcharacter states epistomal suture absent dorsallypresence of ventral setae of forewing and three-segmented tarsus Although not used for the cladisticanalysis highly modified male genital structuresappear to be autapomorphic However they areextremely variable within the family and thus it isvery difficult to decide their homology I have exam-ined only two species of this family and further mor-phological study of the male genitalia will providefurther support for the validity of the autapomorphy

The remaining exemplars Epipsocidae Dolabellop-socidae and Cladiopsocus of the family Cladiopsocidaecomprise a monophyletic group a stable autapomor-phy complete fusion of external and dorsal valves ofthe gonapophyses supports its monophyly Within thisclade Epipsocidae and Dolabellopsocidae are consid-ered to comprise a subclade supported by only oneautapomorphy absence of ventral valve of gonapophy-ses This feature is considered as derived indepen-dently three times in Psocomorpha

Dolabellopsocidae is known only from the Neotropi-cal region Monophyly is supported by only one highlyhomoplastic character state broad pulvillus Diag-nostic characters proposed by Eertmoed (1973) aremostly plesiomorphic highly homoplastic or not con-sistent within the family Further studies are requiredto confirm its validity

Epipsocidae is distributed in all zoogeographicalregions Monophyly is supported by three autapomor-phies epistomal suture absent dorsally absence of A2

vein in forewing and presence of ventral setae offorewing

Cladiopsocidae also is known only from theNeotropical region As discussed above the family isregarded as polyphyletic in the present analysis andcharacter states suggesting nonmonophyly can beregarded as stable In contrast diagnostic charactersproposed by Eertmoed (1973) are mostly plesiomor-

394 K YOSHIZAWA


phic highly homoplastic or not consistent within thefamily As the present analysis strongly suggests poly-phyly I postpone division and declare it incertae sedisbecause I have examined only two species Cladiopso-cidae probably will be divided into at least two inde-pendent families in a future study

Infraorder CaeciliusetaeMonophyly is supported by the following apomorphiccharacter states elongate preepisternum of prothorax(Fig 16) campaniform sensilla on the radius dividedinto two groups absence of preapical tooth of the pre-tarsal claw reduction of external valve of thegonapophyses

Superfamily AsiopsocoideaThis superfamily is represented by only one barkdwelling family Asiopsocidae known from the Nearc-tic Neotropical and Palearctic regions Although itdoes not have to be established to translate the cla-dogram to the Linneaean system I herein maintainthis widely accepted superfamily Monophyly is sup-ported by two unique apomorphic character statesbroadened subapical region of the lacinia (Fig 12)and reduced and membranous dorsal valve of thegonapophyses

Superfamily CaeciliusoideaThis superfamily comprises three families Stenopso-cidae Amphipsocidae and Caeciliusidae A well-developed internal ridge of the epistomal suture andpresence of abdominal eversible vesicles supportmonophyly The derived male and female externalgenitalia are remarkably uniform throughout andmost possibly indicate closer relationships among cae-ciliusioid families Members are living-foliage dwell-ers providings further evidence of monophyly

Phylogenetic relationships are very difficult toestablish because of uniformity of external charactersthroughout the superfamily As mentioned byMockford (1978) the external genitalia are remark-ably uniform However the external valve of thegonapophyses of Dasydemellinae is rather well devel-oped whereas it is greatly reduced in the other taxaThe former character state is regarded as autapomor-phic in the present analysis although it does notunfortunately provide information about interfamilialphylogenetic relationships Only one highly homoplas-tic character state presence of ventral setae on theforewing supports the monophyly of Amphipsocidaesl + Caeciliusidae

Stenopsocidae is distributed in the AfrotropicalPalearctic Oriental and Australian regions Mono-phyly is well supported by the presence of the R1ndashR2+3

crossvein (Fig 40) MndashCuA1 crossvein (Fig 40) andlateral pouch of the spermathecal sac (Fig 40)

Mockford (1993) raised the status of Dasydemellidaefrom subfamilial level within the AmphipsocidaeMonophyly of Amphipsocidae sl is supported by apo-morphic ciliation of the forewing and a pair of concav-ities on the vertex Autapomorphy was found forDasydemellinae and monophyly of Amphipsocinae isalso supported by apomorphic ciliation of the hindwingIn the present analysis Kodamaius which has beenassigned at various times to Amphipsocidae (Smithers1990) Stenopsocidae (Smithers 1972 Badonnel 1981)and Caeciliusidae (Mockford 1993 1999 Yoshizawa1997) is assigned to the clade comprising Dasydemel-linae and Amphipsocinae but included in neither Thusthere are two options for the taxonomic treatment ofthese taxa either all three should be placed within asingle family or treated as independent A morpholog-ical gap is observed between Dasydemellinae and theother taxa especially in the shape of the external valveof the gonapophyses As mentioned above the taxo-nomic position of Kodamaius was ambiguous due to itsunique forewing venation However these morpholog-ical gaps cannot be considered as evidence of familylevel hierarchy because the external valve similar tothat of Dasydemellinae is also observed in some spe-cies of Stenopsocidae and forewing venation is variablewithin Amphipsocinae Therefore it is more practicalto treat the taxa as a single family Amphipsocidae sl(Yoshizawa 2001) Members are widely distributed inall zoogeographical regions

Caeciliusidae is the largest family of Psocomorphaand is widely distributed in all zoogeographicalregions Monophyly was not supported by any autapo-morphy Mockford (1978 1999) proposed some apo-morphic character states but they were not confirmedin the present study or are too homoplasious The fam-ily is possibly paraphyletic I declare it incertae sedisuntil the phylogenetic relationships within it can beestablished

DIAGNOSES OF PSOCOMORPHA INCLUDING INFRAORDERS AND SUPERFAMILIES

SUBORDER PSOCOMORPHA

Head usually with rounded vertex antennae 13-segmented flagellomeres never annulated maxillawithout stipito-galeal muscle hypopharyngeal fila-ments fused for most of their length labial palpus1-segmented Forewing with nodus and thickenedpterostigma nodulus hook-shaped formed by trun-cated spines fused at their base CuP ending togetherwith A1 at wing margin Apex of first axillary scleriteusually with minute proximal process Subgenitalplate lacking median sclerite



ARCHIPSOCETAE INFRAORDER NOV

Small to medium in size about 1ndash3 mm in length(from head to tips of closed wings) Brown to reddish-brown in colour Almost all body surfaces densely cov-ered with long hyaline setae Members mostly barkdwellers living within dense silk nests

Head with rounded vertex postclypeus stronglyconvex epistomal suture with broad internal ridgeanteclypeus weakly sclerotized eyes small no sexualsize dimorphism ocelli usually present not clusteredon small tubercle antennae short mandible shortwith rounded outer margin lacinial tip bicuspid galeaflattened labial palpus rounded Prothorax well devel-oped bulged dorsally Pterothorax not strongly bulgeddorsally dorsal margin almost same level with vertexmedian part of mesoscutellum triangular precoxalbridge of mesothorax broad meso-trochantin narrowsecond phragma strongly swollen posteriorly mem-branous region of metaepisternum narrow Wing poly-morphism present macropterous micropterous orapterous Forewing almost uniformly brown postero-proximal corner strongly angled veins membraneand wing margin densely clothed with long hyalinesetae veins faint pterostigma shallow weakly thick-ened M unbranched areola postica flattened Hind-wing hyaline narrow in shape posteroproximalcorner slightly extended posteriorly veins and mem-brane covered with minute setae distal and posteriormargins with longer setae veins faint and reduced tounbranched Rs M CuA and CuP Sc R1 and Aalmost indistinguishable Tarsi 2-segmented preapi-cal tooth of pretarsal claw absent pulvillus broadAbdomen lacking eversible vesicles ventrally Malegenitalia hypandrium simply rounded posteriorlyPhallosome aedeagus rounded or pointed apicallyparameres weakly sclerotized or more often com-pletely absent aedeagus and phallobase sometimesdetached phallobase closed or opened apicallyendophallus without sclerite Female genitalia Sub-genital plate simply rounded or slightly flattened dis-tally Gonapophyses modified ventral valve absentdorsal valve present or absent narrow and short ifpresent external valve large with long marginal bris-tles All valves absent in viviparous species

Included family Archipsocidae

HEMIPSOCETAE INFRAORDER NOVMedium-sized about 25ndash35 mm Usually whitish topale brown in coloration with blackish bristles Dead-foliage dwellers

Head with rounded vertex clypeus not stronglybulged epistomal suture reduced anteclypeus sclero-tized eyes rather small no sexual size dimorphismocelli complete not clustered on small tubercle

mandible short with smoothly rounded outer marginlacinial tip bicuspid galea flattened labial palpusrounded Prothorax rather well developed Pterotho-rax strongly bulged dorsally its dorsal margin higherthan vertex mesoscutellum somewhat pentagonalmedially precoxal bridge of mesothorax narrowmesotrochantin broad basally membranous region ofmetaepisternum narrow Forewing hyaline to verypale brown in ground colour bearing marginal setaeposteroproximal corner strongly angled pterostigmashallow veins and wing margin setose veins with onerow of setae except CuP bare Rs and M joined atpoint M 2-branched areola postica low CuA1 con-nected with stem of M by crossvein A2 absent Hind-wing glabrous posteroproximal corner angled Scstrong basally but faint distally Rs and M +Cu fusedfor long distance basally Rs and M fused for short dis-tance Tarsi 2-segmented preapical tooth of pretarsalclaw present pulvillus broad Abdomen lacking ever-sible vesicles ventrally Male genitalia paraproct withdistal process directed posteriorly to dorsally Hypan-drium simple Phallosome triangular consisting ofthin phallobase and paramere aedeagus absentparameres strongly bent internally supporting poste-rior margin of endophallus endophallus with or with-out sclerotized portion Female genitalia subgenitalplate simple posterior margin smoothly rounded orslightly hollowed at middle Gonapophyses completeventral valve and dorsal valve simple narrow pointedapically external valve large

Included family Hemipsocidae

INFRAORDER PSOCETAE

Medium to large in size about 3ndash10 mm Colorationvariable but usually whitish in ground colour andwith dark brown markings All bark or stone surfacedwellers

Head with rounded vertex postclypeus not stronglybulged internal ridge of epistomal suture narrowanteclypeus sclerotized male eyes usually muchlarger than femalersquos ocelli complete not clustered onsmall tubercle mandible short outer marginsmoothly rounded posterolateral margin not hol-lowed lacinial tip variable galea flattened labialpalpus rounded Prothorax less bulged dorsallyPterothorax greatly bulged dorsally median part ofmesoscutellum pentagonal mesothorax with narrowprecoxal bridge and broad trochantin membranousregion of metaepisternum broad Forewing oftenwith dark brown markings posteroproximal cornersmoothly rounded veins and wing margin glabroustop of CuA1 usually connected with M except free inPsilopsocidae A2 never present Hindwing glabrousposteroproximal corner smoothly rounded Sc clear Rs

396 K YOSHIZAWA


and M + Cu fused for long distance basally Rs and Mfused or connected by crossvein (some genera in Myo-psocidae) Hind tarsus with row of broad ctenidiabased setae (condition of this character in Psilopso-cidae not examined or documented) tarsi 2- or 3-segmented claws with preapical tooth pulvillusbroad or narrow (Psocidae) Abdomen without eversi-ble vesicles ventrally Male genitalia paraproct withprocess on distal margin Hypandrium usually withcomplex distal processes or lobes Phallosome vari-able internal valve present or absent external valveoften absent endophallus lacking sclerotized portionFemale genitalia paraproct with cylindrical distalprojection Subgenital plate with 1-lobed egg guideextending from ventral margin apex of egg guide usu-ally bearing setae Gonapophyses complete and vari-able in structure external valve internally expanded

Included families Psilopsocidae MyopsocidaePsocidae

INFRAORDER HOMILOPSOCIDEA

This infraorder contains a great diversity of taxa bothmorphologically and biologically

Head usually with rounded vertex except sharplypointed in Calopsocidae postclypeus usually wellbulged epistomal suture with or without well-developed internal ridge anteclypeus variable maleeyes usually much larger than femalersquos ocelli com-plete usually clustered on weakly developed tuberclemandible short with rounded outer margin exceptCalopsocidae with somewhat angled outer marginlacinial tip usually bicuspid galea usually flattenedlabial palpus rounded Prothorax less bulged dorsallypterothorax strongly bulged dorsally mesothoraxwith broad precoxal bridge and narrow trochantinmetaepisternum with broad membranous regionForewing with various coloration and markings pos-teroproximal corner smoothly rounded venationgenerally of Caecilius-type Hindwing with roundedposteroproximal corner Sc faint Male genitalia vari-able phallosome complete Female genitalia sub-genital plate usually with distal projection extendedfrom dorsal margin Gonapophyses usually completedorsal valve swollen dorsally usually with subapicalventral process

ELIPSOCOIDEA SUPERFAM NOVSmall to medium in size about 1ndash4 mm in lengthBody generally brown to blackish brown Bark orstone surface dwellers

Head with rounded vertex postclypeus well bulgedepistomal suture with broad internal ridge ante-clypeus membranous or sclerotized narrowly Wing

polymorphism present macropterous or apterousMacropterous forewing venation of Caecilius-typeveins and wing margin with one row of setae CuPsometimes bare Hindwing scarcely setose marginalsetae present between R2+3 and R4+5 only Rs andM + Cu separate or fused for short distance basally Rsand M fused Tarsi 3-segmented claws with preapicaltooth pulvillus narrow Abdomen without eversiblevesicles ventrally Male genitalia Hypandrium simplyrounded posteriorly Phallosome simple aedeaguspointed apically parameres extended distally farbeyond aedeagus endophallus without rod-like scler-ite Female genitalia Subgenital plate with less devel-oped pair of tubercles on distal margin each bearingsome apical setae Gonapophyses complete ventralvalve simple dorsal valve narrowed apically with orwithout subapical process external valve enlarged

Type genus Elipsocus Hagen 1866

Included family Elipsocidae

LACHESILLOIDEA SUPERFAM NOVLachesillidae Badonnel (1951) emended fromPterodelidae Pearman 1936 is the oldest familygroup name within the superfamily

Small in size about 15ndash4 mm in length Body gen-erally pale brown to blackish-brown Mostly collectedfrom dead-foliage ground litter birdrsquos nests and frombark

Head with rounded vertex postclypeus less bulgedepistomal suture sometimes reduced always withoutbroad internal ridge anteclypeus usually not sclero-tized Pre-episternum of prothorax elongate Meta-epimeron of Ectopsocidae with ball-shaped lobelaterally Forewing hyaline sometimes with blackish-brown markings pterostigma usually shallow areolapostica if present usually free from M absent inEctopsocidae and fused with M in some LachesiliidaeHindwing veins and margin glabrous Rs and M + Cufused for long distance basally Rs and M fused(Lachesillidae) or connected by crossvein (Ectopso-cidae) Tarsi 2- or 3- (Eolachesilla) segmented clawswith or without (Ectopsocidae) preapical tooth pulvil-lus broad or narrow Abdomen without eversiblevesicles ventrally Male genitalia variable cluniumepiproct paraproct and hypandrium often with vari-ous processes Phallosome variable endophallususually with sclerites but absent in some genera ofLachesillidae Female genitalia subgenital plate with(Ectopsocidae and some genera of Lachesillidae) orwithout distal projection extended from dorsal mar-gin Dorsal and ventral valves of gonapophyses oftenreduced external valve narrowed bearing long setae

Type genus Lachesilla Westwood 1840



Included families Lachesillidae Ectopsocidae

PSEUDOCAECILIOIDEA SUPERFAM NOVAll three included families of this superfamily weresimultaneously established by Pearman (1936) and aname could not be decided by the principle of priorityI propose Pseudocaecilioidea since Pseudocaecilius hasthe widest distributional range among the type generaof the three families

Small to middle in size about 2ndash4 mm Colorationvariable Usually living-foliage dwellers except forsome genera of Pseudocaeciliidae (eg Ophiodopelma)

Vertex generally rounded but strongly angled inCalopsocidae postclypeus not strongly bulged epis-tomal suture faint dorsally without broad internalridge anteclypeus sclerotized Forewing veins withmore than one row of setae marginal setae crossing onanterior margin or not (Trichopsocidae) Apex of firstaxillary sclerite broadened Hindwing veins and mar-gins setose Rs and M + Cu separate or fused for shortdistance basally Rs and M fused Tarsi usually 2-segmented claws with or without preapical toothpluvillus broad Abdomen with eversible vesiclesventrally except some dead-foliage dwelling taxa inPseudocaeciliidae Male genitalia hypandrium artic-ulated laterally with clunium or not (Trichopsocidae)with or without (Trichopsocidae) lateral projection andlateral bristle Phallosome aedeagus pointed apicallyparameres long extending far beyond aedeagusendophallus with or without (Trichopsocidae and somegenera of Pseudocaeciliidae) rod-like sclerites Femalegenitalia subgenital plate with well-developed eggguide extending from dorsal margin egg guide 2-lobedeach bearing 1 or 2 apical setae Gonapophyses com-plete ventral valve usually with dorsal lobe apicallydorsal valve with dorsal lobe and subapical processexternal valve variable bearing many long bristles

Type genus Pseudocaecilius Enderlein 1903

Included families Trichopsocidae CalopsocidaePseudocaeciliidae

PERIPSOCOIDEA SUPERFAM NOVPeripsocinae Kolbe 1880 is the oldest family groupname within the superfamily

Small to large in size about 2ndash5 mm in length Bodywhitish in ground colour with blackish-brown mark-ings or wholly blackish-brown All bark dwellers

Head with rounded vertex postclypeus well bulgedepicranial suture with (Mesopsocidae) or withoutbroad internal ridge anteclypeus sclerotized or not(Mesopsocidae) Forewing coloration venation andciliation variable Hindwing ciliation variable Rs andM + Cu separate (most Mesopsocidae) or fused for

short (Philotarsidae) or long (Peripsocidae) distancebasally Rs and M fused Tarsi 2- (Peripsocidae) or 3-segmented claws with preapical tooth pulvillus nar-row Abdomen without eversible vesicles ventrallyMale genitalia hypandrium usually simple exceptPhilotarsus Phallosome variable aedeagus rounded(most of Mesopsocidae and Philotarsidae) or pointedapically phallobase united and rounded apicallyendophallus with (Bryopsocidae Peripsocidae andsome Philotarsidae) or without sclerites Female gen-italia epiproct usually rectangular in dorsal aspectexcept rounded in Bryopsocidae Subgenital plate with1-lobed egg guide extended from dorsal marginGonapophyses complete ventral valve and dorsalvalve strongly united forming ovipositor dorsal valvestrongly sclerotized somewhat rectangular in lateralaspect with or without (Peripsocidae) subapical pro-cess external valve variable

Type genus Peripsocus Kolbe 1866

Included families Bryopsocidae Peripsocidae Philo-tarsidae Mesopsocidae

INFRAORDER EPIPSOCETAE

Medium to large in size about 3ndash7 mm in length Col-oration variable Collected from dead foliage leaf lit-ter or bark and stone surfaces

Head with rounded vertex gena elongate post-clypeus less bulged epistomal suture often reducedalways without well-developed internal ridge ante-clypeus broadly sclerotized anterior tentorial pit sep-arate from ventral margin of cranium male eyes muchlarger than femalersquos ocelli complete or absent dor-sally clustered on small tubercle labrum with pair oflongitudinal sclerotized lines mandible elongate itsouter margin strongly angled and posterolateral mar-gin deeply hollowed maxilla with ball-shaped galealabial palpus somewhat triangular Prothorax lessbulged dorsally Pterothorax well bulged dorsallymesoscutellum pentagonal medially mesothorax withnarrow precoxal bridge and trochantin metaepister-num with broad membranous region Forewing setoseveins often with more than 1 row of setae postero-proximal corner smoothly rounded Rs and M usuallyfused by crossvein A2 usually present but absent inEpipsocidae Hindwing with smoothly rounded poster-oproximal corner veins setose often with more than 1row of setae in distal half margins wholly setoseexcept anterior margin proximal to marginal end ofR1 Rs and M + Cu fused for long distance basally Rsand M fused Tarsi 2- or 3-segmented claws withpreapical tooth pulvillus narrow Abdomen withouteversible vesicles ventrally Male genitalia phallo-some variable phallobase usually open apicallyFemale genitalia subgenital plate usually simple

398 K YOSHIZAWA


Gonapophyses ventral valve present or absent dorsalvalve narrowed to pointed apex external valve nar-rowed dorsal and external valve fused with eachother

Included families Cladiopsocidae PtiloneuridaeDolabellapsocidae Epipsocidae

INFRAORDER CAECILIUSETAE

Small to large in size about 2ndash10 mm in length Sex-ual dimorphism present

Head with rounded vertex eyes variable ocelli com-plete clustered on small tubercle mandible more orless elongate outer margin angled galea ball-shapedlabial palpus somewhat triangular Prothorax lessbulged dorsally preepisternum of prothorax elongatePterothorax greatly bulged dorsally in winged formmesothorax with broad precoxal bridge and narrowtrochantin metathorax with broad membranousregion Forewing variable in venation size shape andcolour posteroproximal corner smoothly roundedHindwing with smoothly rounded posteroproximalcorner ciliation variable Sc clear Rs and M + Cufused for long distance basally Rs and M fused Tarsi2-segmented claws without preapical tooth Male gen-italia Hypandrium simple posterior margin smoothlyrounded Phallosome aedeagus pointed apicallystrongly arched in lateral aspect paramere extendedalong aedeagus strongly arched in lateral aspectphallobase simple and narrow anterior margin notdivided endophallus usually with weakly sclerotizedregion but never with rod-like sclerites Female geni-talia Subgenital plate simply rounded or slightly hol-lowed posteriorly Gonapophyses variable externalvalve more or less reduced

SUPERFAMILY ASIOPSOCOIDEA

Small to medium in size about 2ndash3 mm in lengthGenerally brownish in colour Bark dwellers Sexualdimorphism present male (Notiopsocus) or female(Asiopsocus) often apterous

Head clypeus less bulged epistomal suture withoutbroad internal ridge anteclypeus desclerotized distalmargin of labrum with small projections laterallymandible variable its outer margin more or lessangled lacinia with subapical broadened region lacin-ial tip broad with denticle Forewing as for Caeciliuse-tae pterostigma shallow areola postica present orabsent (Notiopsocus) Hindwing veins glabrous wingmargin glabrous or setose from apex to posterior mar-gin Pulvillus variable sometimes absent Abdomenwithout eversible vesicles on ventral surface Malegenitalia as for Caeciliusetae Female genitaliagonapophyses reduced ventral valve more or less

reduced dorsal valve reduced to membranous lobeexternal valve reduced but somewhat narrowed

Type genus Asiopsocus Guumlnther 1968

Included family Asiopsocidae

SUPERFAMILY CAECILIUSOIDEA

Small to large in size about 3ndash10 mm in length Col-oration of body and wings greatly diverse All living-foliage dwellers

Head capsule somewhat elongate parallel sidedvertex usually rounded anteclypeus narrowly sclero-tized male eyes always much larger than femalersquosocelli complete clustered on well-developed smalltubercle mandible elongate external margin stronglyangled posterior margin deeply hollowed Maxillawith ball-shaped galea lacinial tip usually narrow notdenticulated labium with well-developed salivaryduct labial palpus triangular Forewing venation col-oration and ciliation variable veins and wing marginalways setose Hindwing variable Pulvillus broadAbdomen with eversible vesicles on ventral surfaceMale genitalia as in Caeciliusetae Female genitaliaSubgenital plate simple its posterior margin smoothlyrounded or slightly hollowed medially Gonapophysessimple ventral and dorsal valves narrow and pointedapically external valve reduced sometimes almostindistinguishable with or without few setae

Type genus Caecilius Curtis 1837

Included families Stenopsocidae AmphipsocidaeCaeciliusidae

ACKNOWLEDGEMENTS

I thank T Saigusa H Shima O Yata J Yukawa andK Ogata for their critical reviews and valuable sug-gestions B J Sinclair for critical review correctingthe English version and discussions concerning thecladistic analysis C N Smithers E L Mockford CLienhard and A N Garciacutea Aldrete for their commentson earlier drafts E L Mockford A N Garciacutea AldreteC Lienhard K Konishi T Matsumura K Yasuda NTakahashi and R Matsumoto for providing materialsfor this project E L Mockford and N V Golub for pro-viding their opinions about the phylogenetic positionof Hemipsocidae This study was supported by a fel-lowship from the Japan Society for the Promotion ofSciences and a Grant-in-Aid from the Ministry of Edu-cation Science Sports and Culture Japan

REFERENCES

Badonnel A 1951 Psocoptegraveres In Grasseacute PP ed Traiteacute deZoologie (10) Paris 1301ndash1340



Badonnel A 1967 Psocoptegraveres eacutedaphiques du Chili (2e note)Biologie de lrsquoAmeacuterique Australe 3 541ndash585

Badonnel A 1981 Psocoptegraveres (Insecta Psocoptera) delrsquoInde Mission Besuchet-Loumlbl (1978) et voyage ento-mologique Loumlbl 1979 Revue Suisse de Zoologie 88 381ndash411

Badonnel A Lienhard C 1988 Reacutevision de la famille desMesopsocidae (Insecta Psocoptera) Bulletin du Museacuteumnational drsquoHistoire naturelle 4 10 (A) 2 375ndash412

Betz BW 1983 The biology of Trichadenotecnum alexanderaeSommerman (Psocoptera Psocidae) IV Mechanism of geni-talic coupling Journal of the Kansas Entomological Society56 427ndash433

Brodsky AK 1994 The Evolution of Insect Flight New YorkOxford University Press

Carpenter JM 1988 Choosing among equally parsimoniouscladograms Cladistics 4 291ndash296

Eertmoed GE 1973 The phenetic relationships of the Epip-socetae (Psocoptera) the higher taxa and the species of twonew families Transactions of the American EntomologicalSociety 99 373ndash414

Enderlein G 1903 Die Copeognathen des indo-australischenFaunengebietes Annales Historico-Naturales Musei Nation-alis Hungarici 1 179ndash344

Farris JS 1969 A successive approximations approach tocharacter weighting Systematic Zoology 26 269ndash276

Garciacutea Aldrete AN 1996 Richness of psocid species at theRio Tambopata Reserve Peruvian Amazonia Presented atthe Second International Workshop on Psocoptera GenevaSwitzerland

Goloboff PA 1993 Estimating character weights during treesearch Cladistics 9 83ndash91

Hills DM 1998 Taxonomic sampling phylogenetic accuracyand investigator bias Systematic Biology 47 3ndash8

Kitching IJ Forey PL Humphries CJ Williams DM1998 Cladistics The Theory and Practice of ParsimonyAnalysis 2nd edn London Oxford University Press

Lienhard C 1998 Psocoptegraveres euro-meacutediterraneacuteens Faunede France 83 xx+1ndash517

Maddison DR 1991 The discovery and importance of multi-ple islands of most-parsimonious trees Systematic Zoology40 315ndash328

Maddison WP Donoghue MJ Maddison DR 1984 Out-group analysis and parsimony Systematic Zoology 33 83ndash103

Maddison WP Maddison DR 2000 MacClade Version 4Computer Software and Userrsquos Manual SunderlandSinauer Associates

Matsuda R 1965 Morphology and evolution of the insecthead Memoirs of the American Entomological Institute 41ndash334

Matsuda R 1970 Morphology and evolution of the insectthorax Memoirs of the Canadian Entomological Society 761ndash483

Matsuda R 1976 Morphology and Evolution of the InsectAbdomen Oxford Pergamon Press

Mockford EL 1961 The rediscovery and probable phyloge-netic position of Psilopsocus (Psocoptera) Psyche 68 38ndash44

Mockford EL 1967 The electrentomoid psocids (Psocoptera)Psyche 74 118ndash165

Mockford EL 1972 Anomopsocus and Antipsocus synonymyspecies and taxonomic position (Psocoptera Lachesillidae)Annals of the Entomological Society of America 65 1359ndash1364

Mockford EL 1976 The taxonomic position of the Hemipso-cidae (Psocoptera) Presented at the XV International Con-gress of Entomology Washington DC USA

Mockford EL 1977 Asiopsocus sonorensis (PsocopteraAsiopsocidae) a new record augmented description andnotes on reproductive biology Southwestern Naturalists 2221ndash29

Mockford EL 1978 A generic classification of familyAmphipsocidae (Psocoptera Caecilietae) Transactions of theAmerican Entomological Society 104 139ndash190

Mockford EL 1984 Relationships among philotarsid andpseudocaeciliid genera and a proposed new family Bryopso-cidae (Psocoptera) Psyche 91 (3ndash4) 309ndash318

Mockford EL 1993 North American Psocoptera (Insecta)Fauna and Flora Handbook no 10 Netherland SandhillCrane Press Inc

Mockford EL 1996 The Psocoptera in amber from theDominican Republic Presented at the Second InternationalWorkshop on Psocoptera Geneva Switzerland

Mockford EL 1999 A classification of the psocopteranfamily Caeciliusidae (Caeciliidae auct) Transactions of theAmerican Entomological Society 125 325ndash417

Mockford EL Garciacutea Aldrete AN 1976 A new species andnotes on the taxonomic position of Asiopsocus Guumlnther(Psocoptera) Southwestern Naturalists 20 335ndash346

Mockford EL Sullivan DM 1986 Systematics of the gra-phocaeciliine psocids with a proposed higher classification ofthe family Lachesillidae (Psocoptera) Transactions of theAmerican Entomological Society 112 1ndash80

New TR 1987 Biology of the Psocoptera Oriental Insects 211ndash109

New TR Thornton IWB 1981 Psocoptera from centraland southern Chile Pacific Insects Monograph 37 136ndash178

Novacek MJ 1992 Fossils as critical data for phylogeny InWheeler QD Novacek MJ eds Extinction and PhylogenyNew York Columbia University Press 46ndash88

Pearman JV 1936 The taxonomy of the Psocoptera prelim-inary sketch Proceedings of the Royal Entomological SocietyLondon (B) 5 58ndash62

Platnick NI Griswold CE Coddington JA 1991 Onmissing entries in cladistic analysis Cladistics 7 337ndash343

Roesler R 1944 Die Gattungen der Copeognathen StettinerEntomologische Zeitung 105 117ndash166

Smithers CN 1967 On the relationships of the Calopsocidae(Psocoptera) Journal of the Australian EntomologicalSociety 6 61ndash64

Smithers CN 1969 The Psocoptera of New Zealand Recordsof the Canterbury Museum 84 259ndash344

Smithers CN 1972 The classification and phylogeny of thePsocoptera Memoirs Australian Museum 14 1ndash349

400 K YOSHIZAWA


Smithers CN 1990 Keys to families and genera of Psocoptera(Arthropoda Insecta) Technical Reports of the AustralianMuseum 2 1ndash82

Smithers CN 1991 Psocoptera (Psocids booklice) InNaumann ID Carne PB Lawrence JF Nielsen ESSpradbery JP Taylor RW Whitten MJ Littlejohn MJ edsThe Insects of Australia a Textbook for Students andResearch Workers 2nd edn Melbourne CSIRO MelbourneUniversity Press 412ndash420

Smithers CN 1995a Final instar nymph of Psilopsocus neb-ulosus Mockford (Psocoptera Psilopsocidae) redescribedand compared with two wood-boring species of the genusBeitraumlge zur Entomologie 45 375ndash381

Smithers CN 1995b Psilopsocus mimulus Smithers (Psocop-tera Psilopsocidae) the first known wood-boring PsocopteraJournal of the Australian Entomological Society 34 117ndash120

Smithers CN 1996 Psocoptera In Walls A ed ZoologicalCatalogue of Australia 26 Psocoptera Phthiraptera Thys-anoptera Melbourne CSIRO 1ndash79

Smithers CN 1997 Notes on the biology and annual cycle ofthe wood boring psocopteran Psilopsocus mimulus Smithers(Psocoptera Psilopsocidae) Australian Entomologist 24131ndash136

Swofford DL 2001 PAUP Phylogenetic Analysis using Par-simony (and other methods) Version 4 MassachusettsSinauer Associates

Thornton IWB Smithers CN 1984 Systematics of theCalopsocidae an Oriental and Melanesian family of Psocop-tera Systematic Entomology 9 183ndash244

Thornton IWB Wong SK Smithers CN 1977 The Philo-tarsidae (Psocoptera) of New Zealand and the islands of theNew Zealand Plateau Pacific Insects 12 197ndash228

Wiley EO 1981 Phylogenetics The Principles and Practice ofPhylogenetic Systematics New York John Wiley amp Sons

Yeates DK 1995 Groundplans and exemplars paths to thetree of life Cladistics 11 343ndash357

Yoshizawa K 1997 Psocoptera of Bangladesh collected byProf Emer Katsura Morimoto with descriptions of four newspecies Esakia 37 15ndash24

Yoshizawa K 2001 Systematic study of Amphipsocidae inJapan (Psocodea lsquoPsocopterarsquo) with comments on higherclassification within the family Insecta Matsumurana NewSeries 58 1ndash25

Yoshizawa K Saigusa T 2001 Phylogenetic analysis ofparaneopteran orders (Insecta Neoptera) based on forewingbase structure with comments on monophyly of Auchenor-rhyncha (Hemiptera) Systematic Entomology 26 1ndash13

372

K YOSHIZAWA



2002

136

371ndash400

tae) Homilopsocidea and Psocetae

minus

are presentlyassigned within Psocomorpha (Table 1)

Roesler (1944) proposed three suborders in Psocop-tera corresponding to the presently acceptedTrogiomorpha Troctomorpha and Psocomorpha(

=

Eupsocida

sensu

Roesler) In Psocomorpha he rec-ognized three family groups Epipsocetae Psocetaeand Caecilietae (

=

Caeciliusetae) Psocetae andHomilopsocidea of Pearmanrsquos system were included inPsocetae (

sensu

Roesler) He also grouped or splitsome of Pearmanrsquos families as shown in Table 1

The taxonomic system proposed by Badonnel (1951)is a combination of those of Pearman and Roesler Heretained all Pearmanrsquos families and family groupsand arranged them into Roeslerrsquos subordersBadonnelrsquos system has been widely accepted with onlytwo modifications Smithers (1967) transferred Calop-socidae from Caeciliusetae to Homilopsocidea andMockford (1976) transferred Hemipsocidae fromHomilopsocidea to Psocetae

The monograph by Smithers (1972) is the mostextensive study of the higher classification of Psocop-tera He also investigated the phylogenetic relation-ships among all families and genera of PsocopteraHowever as mentioned above some lineages in his

dendrogram were defined by symplesiomorphic orhomoplastic characters and his phylogenetic classifi-cation has not been accepted

The classifications adopted by Smithers (1996) andLienhard (1998) are based on Badonnel (1951) andinclude the previous updates A few years prior to thisMockford (1993) raised the status of Dasydemellidaeformerly a subfamily of Amphipsocidae but this wasnot followed by Smithers and Lienhard

Within the family groups of Psocomorpha the fol-lowing phylogenetic hypotheses have been proposed

(1) Pseudocaeciliidae and Calopsocidae comprise amonophyletic group (Smithers 1967 Thornton ampSmithers 1984)(2) Elipsocidae and Mesopsocidae comprise a mono-phyletic group with Philotarsidae the sister group(Badonnel amp Lienhard 1988)(3) Caeciliusidae Stenopsocidae and Amphipsocidaecomprise a monophyletic superfamily Caeciliusoideawith Asiopsocidae the sister group (Mockford amp GarciacuteaAldrete 1976)(4) Bryopsocidae is the sister group of the clade com-prising Pseudocaeciliidae and Calopsocidae (Mockford1984)

Table 1

History of the higher classification of Psocomorpha


373



2002

136

371ndash400


Ectopsocus


Peripsocus

(Mockford 1972)


minus





deg






PAUP


CLADISTIC ANALYSIS

E

XEMPLARS






and

Tineomorpha


et al


Tapinella


(1) Archipsocidae

Archipsocus




Hemipsocus

and

Anopistoscena


Hemipsocus




Myopsocus

sp (from Japan) and

Lichenomima muscosa


Psilopso-cus

one exemplar

Psilopsocus


Psocus

sp (nr

bipunc-tatus

from Japan) and

Psocidus



and

Hem-ineura dispar

374

K YOSHIZAWA



2002

136

371ndash400

Table 2


=

missing data)

10

20

30

40

50

60

Echmepteryx

0000000000 0000000000 020000000 0000000000 000000010 0000000110 00000000

Tapinella

0000000000 000000000 000000000 0000200000 0000000100 0000000000 00000000

Troctopsocidae

0000000000 0000000000 100000000 0000200000 0000000100 0000000000 00000000

Tineomorpha

0000000000 0000000000 020000000 0000000000 000000010 0000000000 00000000

Paramphientomum

0000000000 0000000000 020000000 1000000000 000000010 0000000000 00000000

Archipsocus

1000000010 0002100100 021110000 1111000001 1100000102 000000001 20000000

Pararchipsocus

1000000010 0002100100 021110000 1111000001 1100000102 000000001 00000000

Hemipsocus

1011000010 0010000200 0000100001 1011000001 100100010 0010000000 00000000

Psilopsocus

1010000010 0010010111 1000100000 1011200010 1001000102 001001100 00100000

Psocus

1010100010 0010010111 1000100002 1011200011 0001100002 0010010100 00100001

Sigmatoneura

1010100010 0010010111 1000100002 1011200011 0001100002 0010010100 00100001

Myopsocus

1010000010 0010010111 1000100002 1011200010 100100010 0011010000 00000000

Lichenomima

1010000010 0010010111 1000100002 1011201010 1001000000 0011010000 00000000

Elipsocus

1000000010 0011110111 0000100000 1011100000 0000000000 0000100000 01000000

Hemineura

1000000010 0011110111 0000100000 1011100000 0000000000 0000100000 01000000

Ectopsocus

1010000010 0111110111 0000100010 1011201001 1100000000 1000200000 01000100

Eolachesilla

1010000010 0111110111 0000100000 1011200000 1000000000 1000000000 01000100

Nanolachesilla

1011100010 0111110111 0000100000 1011200001 1000000000 100030001 100100

Lachesilla

(Vietnam)

1010100010 0111110111 0000100000 1011200001 000000012 000000001 100100

Lachesilla

1010100010 0111110111 0000100000 1011200001 100000012 000000001 200100

Trichopsocus

1010100010 0011110211 0001100000 1011000101 1110000000 1000000100 01011000

Pseudocaecilius

1010100010 0011110211 0011100000 1011010001 1110111000 0000200101 01011000

Ophiodopelma

1010100010 0011110211 0011100000 1011010001 1100110000 1000200101 01011000

Heterocaecilius

1010100010 0011110211 0011100000 1011010001 1110111000 1000200101 01011000

Calopsocus

0010101110 0011110211 0111100000 1011010001 1010111000 1000200101 01011000

Bryopsocus

1 0100100000 1100 00000000 100030100 0102000

Kaestneriella

1011000010 0011110111 0000100010 1011200001 0000000001 1100300100 01012000

Peripsocus

1010000010 0011111111 1000100010 1011000001 0000000001 1100300100 01012000

Philotarsus

1010100010 0011110111 0011100000 1111010100 0000100100 0100300100 01012000

Aaroniella

1010100010 0011110111 0011100000 1111010100 0000000100 1100300100 01012000

Idatenopsocus






CHARACTERS




376 K YOSHIZAWA









378 K YOSHIZAWA










380 K YOSHIZAWA









382 K YOSHIZAWA









384 K YOSHIZAWA









RESULTS



386 K YOSHIZAWA



HO

MILO

PS

OC

IDE

A










DISCUSSION





388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA

















373



2002

136

371ndash400


Ectopsocus


Peripsocus

(Mockford 1972)


minus





deg






PAUP


CLADISTIC ANALYSIS

E

XEMPLARS






and

Tineomorpha


et al


Tapinella


(1) Archipsocidae

Archipsocus




Hemipsocus

and

Anopistoscena


Hemipsocus




Myopsocus

sp (from Japan) and

Lichenomima muscosa


Psilopso-cus

one exemplar

Psilopsocus


Psocus

sp (nr

bipunc-tatus

from Japan) and

Psocidus



and

Hem-ineura dispar

374

K YOSHIZAWA



2002

136

371ndash400

Table 2


=

missing data)

10

20

30

40

50

60

Echmepteryx

0000000000 0000000000 020000000 0000000000 000000010 0000000110 00000000

Tapinella

0000000000 000000000 000000000 0000200000 0000000100 0000000000 00000000

Troctopsocidae

0000000000 0000000000 100000000 0000200000 0000000100 0000000000 00000000

Tineomorpha

0000000000 0000000000 020000000 0000000000 000000010 0000000000 00000000

Paramphientomum

0000000000 0000000000 020000000 1000000000 000000010 0000000000 00000000

Archipsocus

1000000010 0002100100 021110000 1111000001 1100000102 000000001 20000000

Pararchipsocus

1000000010 0002100100 021110000 1111000001 1100000102 000000001 00000000

Hemipsocus

1011000010 0010000200 0000100001 1011000001 100100010 0010000000 00000000

Psilopsocus

1010000010 0010010111 1000100000 1011200010 1001000102 001001100 00100000

Psocus

1010100010 0010010111 1000100002 1011200011 0001100002 0010010100 00100001

Sigmatoneura

1010100010 0010010111 1000100002 1011200011 0001100002 0010010100 00100001

Myopsocus

1010000010 0010010111 1000100002 1011200010 100100010 0011010000 00000000

Lichenomima

1010000010 0010010111 1000100002 1011201010 1001000000 0011010000 00000000

Elipsocus

1000000010 0011110111 0000100000 1011100000 0000000000 0000100000 01000000

Hemineura

1000000010 0011110111 0000100000 1011100000 0000000000 0000100000 01000000

Ectopsocus

1010000010 0111110111 0000100010 1011201001 1100000000 1000200000 01000100

Eolachesilla

1010000010 0111110111 0000100000 1011200000 1000000000 1000000000 01000100

Nanolachesilla

1011100010 0111110111 0000100000 1011200001 1000000000 100030001 100100

Lachesilla

(Vietnam)

1010100010 0111110111 0000100000 1011200001 000000012 000000001 100100

Lachesilla

1010100010 0111110111 0000100000 1011200001 100000012 000000001 200100

Trichopsocus

1010100010 0011110211 0001100000 1011000101 1110000000 1000000100 01011000

Pseudocaecilius

1010100010 0011110211 0011100000 1011010001 1110111000 0000200101 01011000

Ophiodopelma

1010100010 0011110211 0011100000 1011010001 1100110000 1000200101 01011000

Heterocaecilius

1010100010 0011110211 0011100000 1011010001 1110111000 1000200101 01011000

Calopsocus

0010101110 0011110211 0111100000 1011010001 1010111000 1000200101 01011000

Bryopsocus

1 0100100000 1100 00000000 100030100 0102000

Kaestneriella

1011000010 0011110111 0000100010 1011200001 0000000001 1100300100 01012000

Peripsocus

1010000010 0011111111 1000100010 1011000001 0000000001 1100300100 01012000

Philotarsus

1010100010 0011110111 0011100000 1111010100 0000100100 0100300100 01012000

Aaroniella

1010100010 0011110111 0011100000 1111010100 0000000100 1100300100 01012000

Idatenopsocus






CHARACTERS




376 K YOSHIZAWA









378 K YOSHIZAWA










380 K YOSHIZAWA









382 K YOSHIZAWA









384 K YOSHIZAWA









RESULTS



386 K YOSHIZAWA



HO

MILO

PS

OC

IDE

A










DISCUSSION





388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA
















374

K YOSHIZAWA



2002

136

371ndash400

Table 2


=

missing data)

10

20

30

40

50

60

Echmepteryx

0000000000 0000000000 020000000 0000000000 000000010 0000000110 00000000

Tapinella

0000000000 000000000 000000000 0000200000 0000000100 0000000000 00000000

Troctopsocidae

0000000000 0000000000 100000000 0000200000 0000000100 0000000000 00000000

Tineomorpha

0000000000 0000000000 020000000 0000000000 000000010 0000000000 00000000

Paramphientomum

0000000000 0000000000 020000000 1000000000 000000010 0000000000 00000000

Archipsocus

1000000010 0002100100 021110000 1111000001 1100000102 000000001 20000000

Pararchipsocus

1000000010 0002100100 021110000 1111000001 1100000102 000000001 00000000

Hemipsocus

1011000010 0010000200 0000100001 1011000001 100100010 0010000000 00000000

Psilopsocus

1010000010 0010010111 1000100000 1011200010 1001000102 001001100 00100000

Psocus

1010100010 0010010111 1000100002 1011200011 0001100002 0010010100 00100001

Sigmatoneura

1010100010 0010010111 1000100002 1011200011 0001100002 0010010100 00100001

Myopsocus

1010000010 0010010111 1000100002 1011200010 100100010 0011010000 00000000

Lichenomima

1010000010 0010010111 1000100002 1011201010 1001000000 0011010000 00000000

Elipsocus

1000000010 0011110111 0000100000 1011100000 0000000000 0000100000 01000000

Hemineura

1000000010 0011110111 0000100000 1011100000 0000000000 0000100000 01000000

Ectopsocus

1010000010 0111110111 0000100010 1011201001 1100000000 1000200000 01000100

Eolachesilla

1010000010 0111110111 0000100000 1011200000 1000000000 1000000000 01000100

Nanolachesilla

1011100010 0111110111 0000100000 1011200001 1000000000 100030001 100100

Lachesilla

(Vietnam)

1010100010 0111110111 0000100000 1011200001 000000012 000000001 100100

Lachesilla

1010100010 0111110111 0000100000 1011200001 100000012 000000001 200100

Trichopsocus

1010100010 0011110211 0001100000 1011000101 1110000000 1000000100 01011000

Pseudocaecilius

1010100010 0011110211 0011100000 1011010001 1110111000 0000200101 01011000

Ophiodopelma

1010100010 0011110211 0011100000 1011010001 1100110000 1000200101 01011000

Heterocaecilius

1010100010 0011110211 0011100000 1011010001 1110111000 1000200101 01011000

Calopsocus

0010101110 0011110211 0111100000 1011010001 1010111000 1000200101 01011000

Bryopsocus

1 0100100000 1100 00000000 100030100 0102000

Kaestneriella

1011000010 0011110111 0000100010 1011200001 0000000001 1100300100 01012000

Peripsocus

1010000010 0011111111 1000100010 1011000001 0000000001 1100300100 01012000

Philotarsus

1010100010 0011110111 0011100000 1111010100 0000100100 0100300100 01012000

Aaroniella

1010100010 0011110111 0011100000 1111010100 0000000100 1100300100 01012000

Idatenopsocus






CHARACTERS




376 K YOSHIZAWA









378 K YOSHIZAWA










380 K YOSHIZAWA









382 K YOSHIZAWA









384 K YOSHIZAWA









RESULTS



386 K YOSHIZAWA



HO

MILO

PS

OC

IDE

A










DISCUSSION





388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA




















CHARACTERS




376 K YOSHIZAWA









378 K YOSHIZAWA










380 K YOSHIZAWA









382 K YOSHIZAWA









384 K YOSHIZAWA









RESULTS



386 K YOSHIZAWA



HO

MILO

PS

OC

IDE

A










DISCUSSION





388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA
















376 K YOSHIZAWA









378 K YOSHIZAWA










380 K YOSHIZAWA









382 K YOSHIZAWA









384 K YOSHIZAWA









RESULTS



386 K YOSHIZAWA



HO

MILO

PS

OC

IDE

A










DISCUSSION





388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA




















378 K YOSHIZAWA










380 K YOSHIZAWA









382 K YOSHIZAWA









384 K YOSHIZAWA









RESULTS



386 K YOSHIZAWA



HO

MILO

PS

OC

IDE

A










DISCUSSION





388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA





















380 K YOSHIZAWA









382 K YOSHIZAWA









384 K YOSHIZAWA









RESULTS



386 K YOSHIZAWA



HO

MILO

PS

OC

IDE

A










DISCUSSION





388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA




















382 K YOSHIZAWA









384 K YOSHIZAWA









RESULTS



386 K YOSHIZAWA



HO

MILO

PS

OC

IDE

A










DISCUSSION





388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA





















384 K YOSHIZAWA









RESULTS



386 K YOSHIZAWA



HO

MILO

PS

OC

IDE

A










DISCUSSION





388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA




















RESULTS



386 K YOSHIZAWA



HO

MILO

PS

OC

IDE

A










DISCUSSION





388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA
















386 K YOSHIZAWA



HO

MILO

PS

OC

IDE

A










DISCUSSION





388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA

























DISCUSSION





388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA
















388 K YOSHIZAWA













390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA






















390 K YOSHIZAWA

























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA





























392 K YOSHIZAWA





























394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA

































394 K YOSHIZAWA
























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA


























INFRAORDER PSOCETAE



396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA
















396 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA

































398 K YOSHIZAWA


















ACKNOWLEDGEMENTS


REFERENCES













































400 K YOSHIZAWA



























































400 K YOSHIZAWA
















phylogeny and higher classiﬁcation of suborder...

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