Cladistics (1996) 12:41–64

THE HIGHER CLASSIFICATION OF THE ORDER EMBIOPTERA:A CLADISTIC ANALYSIS

Claudia A. Szumik

Department of Entomology, American Museum of Natural History, Central Park West at79th Street, New York, 10024, U.S.A. and CONICET, Facultad de Ciencias Naturales e

Instituto Miguel Lillo, Miguel Lillo 205, 4000 S.M. de Tucuman, Argentina

Received for publication 24 April 1994; accepted 24 July 1995

Abstract — A matrix of 41 Embiid taxa (representing the 8 formally recognized families of theOrder) and 36 characters were cladistically analysed as a first attempt for understanding thehigher classification of the Order Embioptera. The resulting trees were rooted withClothodidae as the sister group of the other Embioptera. The results suggest that the currentclassification contains several artificial groups. With the rooting used, only Anisembiidae andAustralembiidae are monophyletic. Embiidae is polyphyletic, as Australembiidae+Notoligotomidae, Enveja (incertae sedis) and Oligotomidae+Teratembiidae appear withinEmbiidae, and the “embiid” Microembia appears within Notoligotomidae. Oligotomidae isparaphyletic in terms of Teratembiidae. Four of the genera included in the analysis areparaphyletic: Mesembia, Chelicerca (in terms of Dactylocerca and Pelorembia), Aposthonia (in termsof Oligotoma), and Metoligotoma (in terms of Australembia). Pelorembia and Dactylocerca aresynonymized with Chelicerca.

1996 The Willi Hennig Society

Introduction

The insect order Embioptera is a monophyletic group defined by severalmorphological and behavioral characters. The most striking apomorphy of thesesubsocial insects is their ability to produce silk in all instars from unicellular glandsin the swollen first basitarsus (Barth, 1954; Alberti et al., 1976; Nagashima et al.,1991) and to use it to construct tunnels under bark, stones or soil. Other synapo-morphies of the order are the prognathous head, presence of a gula, absence ofocelli, poorly developed compound eyes, three segmented tarsi, thickened hindfemora, bisegmented cerci, secondary intromittent organs, ovipositor absent andwingless females (Hennig, 1981). In a cladistic analysis of insect orders (Wheeler etal., in prep.), Embioptera appeared as sister group of the Plecoptera by virtue ofsharing tarsal plantulae, reduced phalomeres, a trocantin-episternal sulcus, separ-ate coxopleuron and premental lobes.

Most publications on the systematics of the order Embioptera are descriptions ofnew species and new records. Of all published research on this group, only threepapers (Davis, 1938, 1940b; Ross, 1970) refer to their higher classification.

Two of these contributions (Davis, 1938, 1940b) were based on systematic analy-ses, which, interestingly, used methods closely resembling parsimony methodsused today. Davis (1938) chose 21 terminal taxa as representatives of the mainlineages of Embioptera. He used seven multistate characters, coding the characterswith numbers and the states with letters. Based on the distribution of characterstates observed in his representative taxa, he proposed a scheme (Fig. 1) that had

0748-3007/96/010041+24/$18.00/0 1996 The Willi Hennig Society

42 C. A. SZUMIK

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43ORDER EMBIOPTERA: A CLADISTIC ANALYSIS

“been drawn up in the simplest possible way; convergence has not been invokedunless the opposite course does violence to any known facts.” (1938:266). Davisindicated on his cladogram all the changes (even the homoplastic ones), therebyallowing a complete reconstruction of his data set (Appendix 3). Although Davisincluded some terminal taxa at the nodes of the tree, he pointed out that that wasnot to be interpreted as a proposal of actual ancestry, but instead, “implied thatthe hypothetical ancestor resembled the existing form named—at least for thosecharacters here dealt with—and that the existing type has changed little in thesecharacters from this ancestor.” (1938:262). In a subsequent analysis, the sameauthor (Davis, 1940b), using 55 taxa and 15 characters, presented four trees (eachfor a different geographic area), but this time specifying only the changes betweenhypothetical ancestors and terminal taxa.

Ross (1970) proposed an entirely new classification. He divided the order into 4subgroups and 14 families and rejected all previous classifications on the followinggrounds (1970:164): “No attempts will be made at this time to discuss the history ofEmbioptera classification. . . The great number of new species and higher categor-ies known to the writer make all old classifications obsolete”. It is true that, afterDavis’ work, the number of described genera and species of embiids greatlyincreased (mostly thanks to Ross’ excellent contributions). However, the discoveryof new taxa is certainly not in itself a reason to refute any previous classification,particularly when the evidence cited in support for some of the groups in the newclassification is weak. This lack of justification is extreme in several cases. Forexample, all information given by Ross for his new “Suborder B” (said to includeEnveja bequaerti and undescribed species from Africa) is that it includes “. . . verylarge, often colorful embiids with distinctive features in almost every structure”(1970:169). A new “Family C” is proposed, but no information for the constituentgenera and species is given, except that they are undescribed, they live in Peru andColombia, and they have been discovered recently (1970:168).

Previous work by Davis and Ross on the higher classification of Embiopterainvites, for different reasons, a reanalysis of the problem. In the case of Davis’work, it is tempting to examine to what extent results obtained in one of the firstpapers using cladistic techniques would hold 50 years later. In the case of Ross’work, it is interesting to see to what extent a deep general knowledge of the groupis sufficient in itself to erect groups which, given the lack of explicit justificationand methodology, appear to be based on no more than intuition.

Methods and Materials

CLADISTIC ANALYSIS

Hennig86 (Farris, 1988) was used to calculate most parsimonious tree(s) underequal weights for all characters, with the commands mh* bb*. However, as arguedby Farris (1969, 1983) and most recently by Goloboff (1993a), cladistic analysisdepends on the weights assigned to the characters. Therefore, the programPee-Wee (Goloboff, 1993b) was used to weight the characters and calculate tree(s)that give the highest weight for the characters. The quantity to be maximized isreferred to as the “fit” of the characters, as the weight of a character is a function

44 C. A. SZUMIK

of its homoplasy or fit to a tree; the fit for each character is measured as a concavefunction of its number of extra steps (see Goloboff, 1993a, for justification anddiscussion). In this analysis, the concavity value of Pee-Wee was set at 5 and allcharacters were given a prior weight of 10. Character fits were then calculated as50/5+R+ES, where R is the number of steps implied by polymorphisms in terminaltaxa and ES is the number of extra steps on the tree. The total fit of all the charac-ters is reported as rescaled fit (expressed as percentage) and was calculated usingthe formula 1−(max.fit−fit)/(max.fit−min.fit) (this formula is analogous to that ofthe retention index; see Goloboff’s 1993b documentation). Among all possibletrees, the tree(s) with the highest total fit are chosen.

The command mult* was used to search trees of highest fit, performing 50 repli-cations of a random addition sequence Wagner tree each followed by tree bisec-tion reconnection branch-swapping. The command jump* was used in an attemptto find additional fittest trees, swapping in trees suboptimal by a fit difference ofup to 0·2.

To compare the results with previous classifications the commands force, max/and cmp of Pee-Wee were used. Force was used to define groups to be constrainedfor monophyly and max/ to search the fittest trees containing those groups. Then,cmp was used to check the difference in steps and fit of each character betweenthe constrained and the best-fitting tree(s). To give some notion of how strong thesupport is for each clade in the tree, the commands swap, mv and cmp were used;these commands used in combination find the differences in fit and length when aclade or taxon is moved to another part of the cladogram.

The program Clados (Nixon, 1992) was used to produce tree diagrams. Ambigu-ous optimizations were not considered as support for any clade (i.e. the ambigu-ous- option of Pee-Wee was used). The consensus tree does not show a single mostparsimonious optimization, but instead those synapomorphies shared by 500dichotomous parsimonious resolutions. This was accomplished with the commandapo of Pee-Wee (Goloboff, 1993b), which can be used to find change common tosets of trees. Many more dichotomous parsimonious resolutions exist; these 500were arbitrarily chosen.

TAXA EXAMINED

The trees were rooted with Clothodidae as the sister group to all other Embiop-tera, following the concepts of rooting and polarity most recently described inNixon and Carpenter (1993). Clothodidae has been ‘’defined” (Davis, 1938,1940b; Ross, 1970, 1984) as the family comprising the “simplest” Embioptera. Allother embiids differ from clothodids and are united into a monophyletic group bybeing medium to small in size, having weak and incomplete wing venation, modi-fied head and thoracic structures and asymmetrical male terminalia. The treeswere rooted arbitrarily on Clothoda with Clothodidae as paraphyletic, but the rootmight as well have been placed on Antipaluria or Clothodidae.

To make an analysis of the higher classification, one should include representa-tives of the different groups used by Davis and Ross. However, six of the 14 familiesmentioned by Ross (1970) are made up exclusively of undescribed genera andspecies. Ross claimed (1970:158) that the “. . . species and other details [would] betreated in a series of regional monographs . . .”, but more than 20 years later those

45ORDER EMBIOPTERA: A CLADISTIC ANALYSIS

species still remain undescribed. Obviously, those presumed new families couldnot be included in this analysis. A matrix with 41 species in 32 genera and eightfamilies was assembled (Table 1; see Appendix 1 for a complete list of specimensand vouchers). Each species used was chosen because it has a character combi-nation corresponding to a taxon used in the classifications of Davis and Ross. Toinsure that no significant character combinations were excluded from the analysis,51 other species were examined (see Appendix 2).

CHARACTERS

Characters 0, 4, 5, 7, 9, 23 and 24 were used by Davis (1938 and 1940b), andcharacters 2, 11, 13, 14, 15, 20 and 22 by Davis (1940b).

The following abbreviations are used in the description of the characters. Wings:Rs, radial section; R1, first radial; Ma, anterior medial vein; Mp, posterior medialvein; Cu, cubittal vein. Terminalia: Ep, epiproct; Lpp, left paraproct; H, hypand-rium or ninth abdominal sternite; Hp, process of the hypandrium; LC1, basal seg-ment of the left cercus; LC2, apical segment of the left cercus; RC1, basal segmentof the right cercus; RC2, apical segment of the right cercus; 10T, tenth abdominaltergite; 10L, left hemitergite of the tenth abdominal tergite; 10R, right hemitergiteof the tenth abdominal tergite.

(0) Middle bladder of the hind basitarsus: 0, present; 1, absent.(1) Shape and chaetotaxy of the hind basitarsus: 0, broad and long with many

setae on the ventral surface; 1, narrow and long with few setae on the ventralsurface; 2, broad and short with few setae on the ventral surface.

(2) Size of the male hind basitarsus middle bladder: 0, large; 1, small.(3) Female genital plate (Fig. 2): 0, central plate separated from lateral plates

(Fig. 2a); 1, central plate partially fused to the lateral plates (Fig. 2b); 2, centralplate fused to the lateral plates and differentiated from them on the posteriormargin (Fig. 2c). Stefani (1953a) first described the subgenital plate (8th sternite)and made dissections in some species of Embia and Haploembia (Stefani, 1953a,b,c,1955). Because his work suggested that this character could differentiate taxa, thefemale genitalia was examined for all taxa included here (if females wereavailable).

(4) Wings, Ma vein: 0, forked; 1, unforked.(5) Wings, Cu vein: 0, forked: 1, unforked.(6) Unions on the wing base between Rs+Ma and Mp with R1 and Cu veins (Fig.

3). 0, Rs+Ma and Mp fork together from Cu, a cross-vein present between Rs+Maand R1 (Fig. 3a); 1, Rs+Ma and Mp start separately from R1 and Cu (Fig. 3b); 2,Rs+Ma and Mp start fused from R1 and Cu (Fig. 3c).

(7) Wings: 0, present; 1, absent. Apterism occurs in all families of the order; itcan occur within a genus, a species, or even a population (Davis, 1938; Ross, 1984).Those species with both winged and wingless males (Anisembia texana, Chelicercawheeleri and Notoligotoma nitens) were scored here as winged.

(8) Mandibles (Fig. 4): 0, incisor and molar areas robust with many teeth with-out any space between them (Fig. 4a); 1, incisor area conspicuous, three teeth onthe left mandible, two on the right, molar area with two or more teeth with a spacebetween these areas (Fig. 4b); 2, incisor area with the same number of teeth as in1, but sharper and located on the tip of the mandibles, molar area with few

46 C. A. SZUMIK

Tab

le1

Dat

am

atri

x.C

hara

cter

num

bers

and

codi

ngco

rres

pond

toth

ose

indi

cate

din

the

text

12

30

12

34

56

78

90

12

34

56

78

90

12

34

56

78

90

12

34

5

Clo

thod

a0

00

20

00

01

00

0—

—0

00

00

00

0—

——

00

00

00

00

00

0A

ntip

alur

ia0

00

20

01

01

00

0—

—0

00

00

01

0—

00

00

00

00

00

00

0A

rche

mbi

a0

11

10

11

02

00

11

11

00

00

04

10

22

00

01

00

00

00

0C

onic

erem

bia

00

01

01

20

10

00

——

10

00

00

41

02

40

00

00

00

00

00

Din

embi

a0

01

10

11

01

00

10

01

10

00

03

11

52

00

00

10

00

00

0D

onac

onet

his

10

——

01

—0

10

01

00

11

00

00

41

15

20

00

00

01

00

00

Em

bia

10

—1

01

00

10

01

11

10

00

10

31

12

20

00

01

01

00

00

Mac

hado

embi

a1

0—

10

10

01

00

11

11

00

01

04

11

22

00

00

10

00

00

0M

etem

bia

00

01

01

00

10

01

00

11

00

00

31

12

20

00

00

01

00

00

Mic

roem

bia

11

——

01

20

31

11

11

10

00

00

41

01

20

00

00

00

00

00

Neo

rhag

adoc

hir

00

12

01

10

20

00

——

00

01

00

41

02

40

00

01

10

00

00

Pac

hyle

mbi

a0

00

1—

——

12

00

0—

—0

00

10

04

10

21

00

00

11

00

00

0P

arar

haga

doch

ir0

01

20

11

02

00

11

11

00

00

04

10

54

00

01

10

00

00

0P

arem

bia

00

0—

01

10

10

01

21

10

00

00

41

12

20

00

00

01

00

00

Pse

udem

bia

00

11

01

00

10

01

01

11

00

10

31

15

20

00

01

00

00

00

Scel

embi

a0

01

10

11

02

00

11

11

00

00

04

10

52

00

01

00

00

00

0E

nvej

a0

00

—0

11

06

00

12

10

10

00

04

10

13

00

00

00

00

00

0A

ustr

alem

bia

10

—0

——

—1

03

11

01

10

00

01

31

21

11

00

00

00

00

00

M.c

onve

rgen

s0

00

——

——

10

31

10

11

00

00

13

12

52

10

00

00

00

00

0M

.ing

ens

00

00

——

—1

03

11

01

10

00

01

31

22

11

00

00

00

00

00

A.b

orne

ensi

s1

1—

21

12

03

00

0—

—1

00

00

05

10

33

00

0—

——

——

11

0A

.gla

uert

i1

1—

21

12

03

00

0—

—1

00

00

05

10

43

00

00

00

00

01

0H

aplo

embi

a0

00

——

——

16

00

0—

—1

00

00

05

10

32

00

00

00

00

01

0O

ligo

tom

a1

1—

21

12

03

00

0—

—1

01

00

05

10

33

00

0—

——

——

11

0T

erat

embi

a1

1—

20

12

03

00

0—

—1

01

00

05

10

43

00

00

00

00

01

0A

nise

mbi

a1

2—

21

11

04

21

11

11

00

00

01

0—

11

00

00

00

00

00

0B

ulbo

cerc

a1

1—

——

——

14

21

11

11

00

00

02

0—

11

00

00

00

00

00

0C

.dam

pfi

11

—2

11

10

53

11

11

10

00

00

20

—4

30

01

00

00

00

01

C.d

avis

i1

1—

21

11

05

11

11

11

00

00

02

0—

43

00

10

00

00

00

1C

.jal

isco

a1

1—

2—

——

14

31

11

11

00

00

02

0—

43

00

10

00

00

00

1C

.max

ima

11

—2

11

10

52

11

11

10

00

00

20

—4

30

10

00

00

00

00

C.w

heel

eri

11

—2

11

10

53

11

11

10

00

00

20

—4

30

10

00

00

00

00

Dac

tylo

cerc

a1

1—

21

11

04

31

11

11

00

00

02

0—

43

00

10

00

01

00

1M

.cat

emac

oa1

2—

21

11

05

21

11

11

00

00

01

0—

11

00

00

00

00

00

0M

.cha

mul

ae1

2—

21

11

05

00

11

11

00

00

01

0—

11

00

00

00

00

00

0M

.ven

osa

12

—2

11

10

53

11

11

10

00

00

10

—2

10

00

00

00

00

00

Pel

orem

bia

10

—2

——

—1

01

11

11

10

00

00

10

—4

30

10

00

00

00

00

Sten

embi

a1

1—

21

11

05

00

0—

—0

00

00

01

0—

11

00

00

00

00

00

0N

.har

dyi

00

0—

11

10

12

11

11

10

00

00

41

02

20

00

00

00

10

00

N.n

iten

s0

00

—1

11

03

21

10

11

00

00

04

10

22

00

00

00

01

00

0P

tilo

cere

mbi

a1

0—

—0

11

03

21

11

11

00

00

04

10

42

00

00

00

00

00

0

47ORDER EMBIOPTERA: A CLADISTIC ANALYSIS

2a 2b

2c

Fig. 2. Female genital plate. (a) Australembia incompta; (b) Archembia sp.; (c) Antipaluria caribbeana.

R1

3c3a 3b

Rs+Ma

Mp

Cu

Fig. 3. Wing base unions. (a) Pseudembia truncata; (b) Mesembia venosa; (c) Oligotoma saundersii.

rounded teeth (Fig. 4c); 3, incisor area same as 2, but molar area with a conspicu-ous, sharp tooth (Fig. 4d); 4, ncisor area with 2 teeth on the left mandible, and oneon the right, molar area same as 3 (Fig. 4e); 5, incisor area with one tooth on eachmandible, molar area same as 3 (Fig. 4f); 6, mandibles curves, on each tip manyrounded teeth incisor and molar area not differentiated (Fig. 4g). A great varietyof shapes are exhibited by the mandibles. Some of those shapes are clearly distinctand easily identifiable (for example, states 3, 4 and 5). However, the distinctionbetween some other states (e.g. 0, 1 and 2) is more difficult and some taxa couldperhaps be scored differently for this character. This is the case for Parembia per-sica, Metembia ferox and Machadoembia angolica which were scored as having state 1,but might be scored state 2 as well.

(9) Apical left cercus (LC2) (Fig. 5): 0, normal (Fig. 5a); 1, reduced (Fig. 5b); 2,fused to the basal segment but conspicuous (Fig. 5c); 3, fused and not distinguish-able from basal segment (Fig. 5d). The form of the LC2 is variable in Anisembii-dae. Chelicerca has species with states 1, 2 or 3, and the genus Mesembia includesspecies with states 0, 2 or 3. For both of these genera, several species (representingthe variation in this character) were included in the matrix (Table 1).

(10) Apical right cercus (RC2): 0, normal; 1, reduced.(11) Basal left cercus (LC1), macrosetae: 0, absent; 1, present.(12) LC1, macrosetae distribution: 0, on apical and basal area; 1, on apical area

48 C. A. SZUMIK

4b 4c4a

4f 4g4d 4e

IM

Fig. 4. Mandibles. (a) Australembia incompta; (b) Dinembia ferruginea; (c) Machadoembia angolica; (d)Microembia rugosifrons; (e) Anisembia texana; (f) Chelicerca wheeleri; (g) Haploembia.

5a 5b 5c 5d

Fig. 5. Left cercus. (a) Aposthonia borneensis; (b) Microembia rugosifrons; (c) Mesembia catemacoa; (d)Chelicerca jaliscoa.

only; 2, on basal area only. Dactylocerca flavicollis and Chelicerca jaliscoa have somemiddle-basal macrosetae (Fig. 5d), and therefore could have state 0 (instead of 1,as scored here). Other taxa with state 0, however, have many more macrosetae inthe basal area.

(13) LC1, macrosetae size: 0, large; 1, small.(14) LC1, apical nodule: 0, absent; 1, present.(15) LC1, basal nodule: 0, absent; 1, present.(16) LC1, basal and lateral process: 0, absent; 1, present.(17) LC1, base: 0, narrow; 1, broad.(18) LC1, inner side: 0, not depressed; 1, depressed.(19) RC1, shape: 0, normal; 1, reduced to a broad base.(20) Tenth abdominal tergite (10T) (Fig. 6): 0, only one sclerotized plate (Fig.

6a); 1, two plates (10L and 10R) more or less fused to each other on the basal half(Fig. 6b); 2, 10L and 10R with irregular inner margins and with a central membra-nous area between both plates (Fig. 6c); 3, 10L and 10R with well-differentiated

49ORDER EMBIOPTERA: A CLADISTIC ANALYSIS

6d 6e 6f

6a 6b 6c

Fig. 6. Tenth abdominal tergire. (a) Clothoda longicauda; (b) Stenembia perenensis; (c) Chelicerca jaliscoa;(d) Metembia ferox; (c) Scelembia malkini; (f) Oligotoma saundersii.

7a 7b 7c 7d 7e 7f

Fig. 7. Process of the left hemitergite of the tenth abdominal tergite. (a) Antipaluria caribbeana; (b)Australembia incompta; (c) Machadoembia angolica; (d) Aposthonia sp; (e) Chelicerca jaliscoa; (f)Pararhagadochir trachelia.

inner margins and fused on the base with a sclerotized band (Fig. 6d); 4, twoplates, 10R with a membranous basal area and sclerotized basal band present (Fig.6e); 5, 10R apical half not fused with the rest of the tergite (Fig. 6f).

(21) 10R, second process: 0, absent; 1, present.(22) 10R, second process form: 0, rod-like, very sclerotized; 1, lateral flap not

extended over the 10L, more or less sclerotized; 2, lateral flap extended over the10L, sclerotized.

(23) 10L, process (Fig. 7): unsclerotized lobe (Fig. 7a); 1, more or less sclerot-ized lobe (Fig. 7b); 2, sclerotized flat hook (Fig. 7c); 3, more or less sclerotizedtwisted oval leaf (Fig. 7d); 4, inner and outer margins differentiated in hook andmembranous portions, fused to each other (Fig. 7e); 5, same as 4 but partially ortotally separated (Fig. 7f). Some species of Oligotoma (not included in the matrix)have state 4; the only species of Oligotoma included (O. saundersii) has state 3. Anextra step was therefore added to this character.

50 C. A. SZUMIK

(24) 10R, first process: 0, unsclerotized lobe; 1, sclerotized lobe; 2, sclerotizedsharp hook; 3, sclerotized lobe with a small thorn on the dorsal surface; 4, same as3, but with a long spine instead of the small thorn.

(25) H and Hp: 0, Hp fused to H; 1, Hp separated of H.(26) Sclerotized hook on the posterior area of the Hp: 0, absent; 1, present.(27) Hp, denticles: 0, absent; 1, present.(28) Posterior side of the Lpp turned on the right as a flat hook: 0, absent; 1,

present. The genus Pararhagadochir is polymorphic for this character because somespecies do not have a flat hook in the Lpp. Therefore, 1 extra step was added tothis character.

(29) Lpp, nodule: 0, absent; 1, present. In Embia and Pseudembia the nodule canbe present or absent; two extra steps were added.

(30) Lpp, denticles: 0, absent; 1, present.(31) Lpp, sclerotized spine: 0, absent; 1, present. Species of Embia may either

have or lack this spine; 1 extra step was added.(32) Ep denticles: 0, absent; 1, present.(33) Lpp reduced to a long sclerotized spine: 0, absent; 1, present.(34) Length of the Hp and H: 0, Hp short, minor than H; 1, Hp longer (longer

than H).(35) Hp: 0, rectangular; 1, circular.

Results

Pee-Wee produced three trees of maximum fit (378·84 [58%]), 132 steps long.Two of those trees differ only in the partial resolution of the group formed byChelicerca davisi+C. dampfi+C. jaliscoa+Dactylocerca; the third is the same as the strictconsensus of the three. All of the characters require the same number of steps onthe three trees; the different resolutions result only from ambiguities in optimiza-tions. The discussion will use as reference the strict consensus (Fig. 8). The blackboxes indicate only unambiguous synapomorphies, common to 500 dichotomousparsimonious resolutions (see Methods section), and the white boxes indicate syn-apomorphies present in only some of the resolutions. Only those changes occur-ring in all trees are considered to be synapomorphies of the groups, but thoseoccurring in some of the trees might become unambiguous if future evidenceallows further resolution of the cladogram (i.e. forbid some of the possible resol-utions!).

The three trees of highest fit found with Pee-Wee are not among the 2275 (bb*overflow; length=131, CI=45, RI=74) found by Hennig86 with all characters equallyweighted and non-additive (they are one step longer). The strict consensus ofthose 2275 trees is almost totally unresolved, but each of its groups is also presentin the consensus of the three Pee-Wee trees. The groups in common are: Neorhaga-dochir+Pachylembia, Aposthonia borneensis+Oligotoma, Australembia+Metoligotoma con-vergens+M. ingens, Archembia+(Pararhagadochir+Scelembia), Chelicerca davisi+C.dampfi+(C. jaliscoa+Dactylocerca).

Discussion

Anisembiidae—This is one of only two families which are monophyletic in thepresent analysis. The group is characterized by the absence of middle bladder

51ORDER EMBIOPTERA: A CLADISTIC ANALYSIS

Fig. 8. Consensus of the three fittest trees. CLO, Clothodidae; ANI, Anisembiidae; EMB, Embiidae;OLI, Oligotomidae; TER, Teratembiidae; NOT, Notoligotomidae; AUS, Australembiidae; INC, incertaesedis. Black hashes indicate unambiguous synapomorphies occurring in all of 500 dichotomousparsimonious resolutions and white hashes indicate synapomorphies occurring in only some.

(char. 0), Ma vein unforked (char. 4), and mandibles with incisor and molar areawith only one tooth (char. 8).

Two of the three trees obtained in the present analysis differ in the partial resol-ution the group formed by Chelicerca davisi+C. dampfi+C. jaliscoa+Dactylocerca, the

52 C. A. SZUMIK

third tree is identical to the consensus tree of the other two. These two partial res-olutions are based on two possible optimizations of the highly homoplastic charac-ter 8 (mandibles). One of the trees groups C. davisi and C. dampfi by having anincisor and molar area with one tooth (state 5), the other groups C. jaliscoa andDactylocerca by having one and two teeth on the right and left incisor (state 4). Ifchar. 12, of dubious scoring for Dactylocerca flavicollis and C. jaliscoa was scored ashaving state 0 in those taxa, the results would remain unchanged (with state 0appearing as a synapomorphy of those two taxa).

The genus Chelicerca appears as paraphyletic; some of its species are more closelyrelated to Pelorembia (sharing a hook on the Hp [char. 26]) and others to Dac-tylocerca (sharing the presence of denticles on the Hp [char. 27] and the hypandr-ial shape [char. 35]). Two species of Chelicerca not included in the present analysis(C. grandis and C. callani) have plesiomorphic states for those three characters;given their character combinations it is likely that those species, if included, wouldappear as sister group of the other species of Chelicera+Dactylocerca+Pelorembia.

Table 2Character statistics, according to the trees with fit 278.84

Character Steps (E.S.) Weight CI RI

0 6 (5) 5·00 16 701 7 (5) 5·00 28 682 3 (2) 7·14 33 603 4 (2) 7·14 50 774 3 (2) 7·14 33 855 1 (0) 10·00 100 1006 6 (4) 5·55 33 557 6 (5) 5·00 16 288 13 (7) 4·16 40 709 9 (6) 4·54 33 60

10 2 (1) 8·33 50 9411 2 (1) 8·33 50 9012 5 (3) 6·25 40 6213 2 (1) 8·33 50 5014 3 (2) 7·14 33 6015 3 (2) 7·14 33 5016 2 (1) 8·33 50 017 1 (0) 10·00 100 10018 1 (0) 10·00 100 10019 1 (0) 10·00 100 10020 8 (3) 6·25 62 8621 1 (0) 10·00 100 10022 2 (0) 10·00 100 10023 16 (11) 2·94 31 5224 9 (5) 5·00 44 7625 1 (0) 10·00 100 10026 1 (0) 10·00 100 10027 1 (0) 10·00 100 10028 1 (0) 10·00 100 10029 3 (2) 5·55 33 6630 1 (0) 10·00 100 10031 3 (2) 6·25 33 3332 2 (1) 8·33 50 5033 1 (0) 10·00 100 10034 1 (0) 10·00 100 10035 1 (0) 10·00 100 100

53ORDER EMBIOPTERA: A CLADISTIC ANALYSIS

Tab

le3

Fitd

iffe

renc

ebe

twee

nth

efi

ttest

tree

and

tree

sha

ving

the

grou

psin

dica

ted

asm

onop

hyle

tic.

For

the

diff

eren

cein

fit

inin

divi

dual

char

acte

rs,

the

num

bers

inpa

rent

hese

sin

dica

te(s

tep

diff

eren

ce,f

itdi

ffer

ence

)

Tre

eFi

tdif

f.L

engt

hdi

ff.

Cha

ract

ers

with

bette

rfi

tC

hara

cter

sw

ithw

orse

fit

Ani

sem

biid

ae−8

·18

61(

2,1·

25);

8(2,

0·84

);13

(1,1

·67)

;2(

1,0·

89);

9(1,

0·38

);10

(1,1

·19)

;11(

2,2·

08);

mon

ophy

letic

14(1

,1·1

9);2

3(1,

0·18

)12

(1,0

·70)

;21(

1,1·

67);

24(2

,0·8

4)26

(1,1

·67)

;27(

1,1·

67);

29(1

,0·5

5);3

5(1,

1·67

)

Em

biid

ae−6

·21

423

(3,0

·39)

;31(

1,0·

89)

0(2,

0·89

);4(

1,0·

89);

10(1

,1·1

9);1

1(1,

1·19

);m

onop

hyle

tic12

(1,0

·70)

;18(

1,1·

67);

20(1

,0·7

0);2

9(1,

0·55

)

Not

olig

otom

idae

−1·1

33

13(1

,1·6

7);1

4(1,

1·19

);23

(1,0

·18)

0(2,

0·84

);2(

1,0·

89);

11(1

,1·1

9);1

2(1,

0·70

);m

onop

hyle

tic29

(1,0

·55)

Olig

otom

idae

−3·4

46

13(1

,1·6

7);1

4(1,

1·19

)0(

2,0·

84);

1(1,

0·46

);2(

1,0·

89);

11(1

,1·1

9);

mon

ophy

letic

12(1

,0·7

0);2

9(1,

0·55

);33

(1,1

·67)

Ros

s,19

70−1

9·77

181(

1,0·

55);

23(1

,0·1

8)0(

2,0·

84);

2(1,

0·89

);8(

1,0·

24);

9(1,

0·38

);31

(1,0

·89)

10(3

,2·7

8);1

1(1,

1·10

);12

(1,0

·70)

;14(

1,0·

89);

18(1

,1·6

7);2

0(1,

0·7)

;21(

1,1·

67);

24(1

,0·4

6);

26(1

,1·6

7);2

7(1,

1·67

);28

(1,1

·67)

;29(

1,0·

55);

33(1

,1·6

7);3

5(1,

1·67

)

54 C. A. SZUMIK

Pelorembia was created for a single species with highly modified head, mandiblesand terminalia, but those characters are autapomorphies and this species and Dac-tylocerca are both within the Chelicerca clade. Ross was aware of the paraphyly ofChelicerca in terms of Pelorembia and Dactylocerca: “In spite of the simplicity of theterminalia, the affinities of the genus [Pelorembia] appear to be with the wheelerigroup of Chelicerca . . . Dactylocerca appears to be derived from the davisi group ofChelicerca. Some species of these genera could almost be assigned to either, but thebulk of Dactylocerca are clearly generically distinct.” (1984:41, 38). Chelicerca mustnow be considered to be the senior synonym of Pelorembia and Dactylocerca.

Constraining the monophyly of all anisembiid genera implies an additional sixsteps, saving steps in 5 characters and sacrificing them in 11 (Table 3).

Oligotomidae+Teratembiidae—Both families share the 10R apical half not fusedwith the rest of the tergite (char. 20, state 5) and the Hp prolonged and curved(char. 34).

Haploembia is the sister group of the other Oligotomidae and Teratembiidae(here represented only by Teratembia), which share the absence of a bladder (char.0) and a narrow and long basitarsus with few setae (char. 1). Both characters arequite homoplastic; the loss of the middle bladder, in particular, occurs repeatedlyin the order. A third shared character is the lobe with a sharp thorn on the dorsalsurface of 10R (char. 24), a state also found in some anisembiids.

Davis (1940b:536) listed a series of characters shared by Oligotomidae andTeratembiidae but was still of the opinion that they were not closely related. Ross(1970) grouped Oligotomidae+Teratembiidae in his “suborder C”, although hedid not explain the basis for this action.

Aposthonia was synonymized with Oligotoma by Davis (1940b). In 1951, Rosstreated Aposthonia as a subgenus of Oligotoma, but later as a genus (1963; discussingsome Australian species of Aposthonia). The present results suggest that Aposthoniais paraphyletic. The character linking it to Oligotoma is the Ma unforked (char. 4),a state found also in Anisembiidae and Notoligotoma. A. borneensis and O. saundersiishare the shape of the Lpp (char. 34), but this character is actually absent in someAposthonia. The generic limits made by Ross for these genera seem to have beenbased, not on morphological characters, but only on the assumption that the spec-ies of India would fit in Oligotoma, and those of Australia and the Pacific in Apos-thonia. The actual limits between the two genera (which include, in total, 34species) seem more complex than suggested by this simple geographic delimi-tation and constitute a problem exceeding the scope of this paper.

Constraining the monophyly of Oligotomidae and its genera implies a totalincrease of 8 steps (shorter for 2 characters and longer for 7; Table 3).

Notoligotomidae+Australembiide—Notoligotomidae was one of the familiesmost studied by Davis (1936a,b, 1938, 1940a, 1942a,b, 1944a,b) and he includedNotoligotoma, Ptilocerembia, Embonycha, Burmitembia and Metoligotoma in it. In 1963,Ross made several significant changes to the composition of Notoligotomidae. Heremoved Embonycha and gave it familial status, providing a typically gradistic justifi-cation: “Although Embonycha appears to have developed from the same stock asNotoligotoma, it seems sufficiently differentiated to justify a separate family”(1963:123). Burmitembia was also removed from the family, but it was not explicitlyplaced elsewhere, so it has been treated as incertae sedis. Metoligotoma was alsoremoved and included with Australembia in the newly erected family Australembii-

55ORDER EMBIOPTERA: A CLADISTIC ANALYSIS

dae. As a result of the changes made by Ross, the family Notoligotomidae eventu-ally contained only three species and two genera, Ptilocerembia roepkei, Notoligotomahardyi and Notoligotoma nitens.

In his 1963 paper, Ross listed the characters supporting his view that Australem-biidae is not related to any other group of Embioptera. It is obvious that Austra-lembiidae is a monophyletic group (since it is supported by numerous, almost non-homoplastic characters), but it is no less obvious that (unless it is the sister groupof all other Embioptera, a hypothesis easily rejected by numerous characters) itmust be more closely related to some Embioptera than to others. The problem iswhich one. According to Ross (1970) Australembiidae is the only family in his“suborder B”, and Notoligotomidae belongs to the suborder “Embioptera”,together with Embiidae, Clothodidae and Anisembiidae. In the best fit trees pre-sented here, however, Notoligotomidae and Australembiidae are grouped by hav-ing the LC2 fused to the LC1 but conspicuous (char. 9) and by the reduced RC2(char. 10). Both characters are also found in some anisembiids.

Notoligotomidae, as defined by Ross, is paraphyletic because Ptilocerembia ismore closely related to Microembia Ross (Embiidae) than to Notoligotoma (Fig. 8).Ptilocerembia and Microembia are grouped by having no middle bladder (char. 0).Constraining the monophyly of Notoligotomidae increases the tree length by threesteps. The fit decreases only slightly, but only three characters are favored (whilefive have a worse fit).

Microembia (Embiidae) is a genus with a complex character combination andimportant autapomorphies. Ross (1944) did not make his reasons for includingthe genus in Embiidae explicit, but it can be assumed that it was because of its for-ked Ma, because that is the only character shared with embiids (and teratembiids).The absence of the middle bladder (char. 0), the type of mandibles (char. 8) andthe form of the 10T (char. 20) relate this genus to Teratembiidae. The absence ofthe middle bladder and the type of genitalia (char. 10, 23, 24) relate it to someAnisembiidae. Other possible placements of Microembia included as sister group ofeither Notoligotoma or Australembiidae, or Australembiidae+Notoligotomidae, butall of these imply a decrease in fit as small as 0·46 (adding a step in char. 0).

Embiidae—As currently delimited, the Embiidae is one of the largest families inthe order, with representatives in all continents except Australia. In the trees ofmaximum fit, the family appears polyphyletic. Forcing the monophyly of Embiidae(and Ross’ subfamilies) implies eight additional steps and a decrease in fit of 5·56,improving the fit for 4 characters and decreasing it for 10 (Table 3).

The Neorhagadochir+Pachylembia clade diverges first from the tree because itsmembers lack a process and denticles in the LC1 separate it from the other Neo-tropical embiids. The monophyly of Neorhagadochir+Pachylembia is supported by thetype of mandible (state 2, char. 8), the exclusive basal broadening of the LC1(char. 17) and the presence of a nodule and denticles in the Lpp (chars. 29 and30, present in other embiid genera as well).

Conicerembia has some modifications that obscure its possible relationships withother Neotropical embiids, particularly in the shape of the LC1. For example,making Conicerembia the sister group of Teratembia+Oligotomidae implies adecrease in fit of only 0·19, saving one step in char. 6 and adding another in char.3 (as char. 6 is more homoplastic than 3, the trees that save a step in char. 3 areslightly preferable, even if having the same absolute number of steps).

56 C. A. SZUMIK

AustralembiaM. convergensM. ingensTeratembiaHaploembiaOligotomaA. borneensisA. glavertiEnvejaAnisembiaBulbocercaDactylocercaPelorembiaStenembiaM. catemacoaM. chamulaeM. venosaC. davisiC. dampfiC. jaliscoaC. maximaC. wheeleriPtilocerembiaN. hardyiN. nitensDonaconethisEmbiaMachadoembiaMetembiaParembiaDinembiaPseudembiaConicerembiaMicroembiaNeorhagadochirPachylembiaParahagadochirScelembiaArchembiaClothodaAntipaluria

Australembiidae

Teratembiidae

Oligotomidae

Unnamed family

Anisembiidae

Notoligotomidae

Embiinae

Subfamily D

Subfamily B

Subfamily A

Clotodidae

Em

biid

ae

Em

biid

ina

Suborder A

Suborder C

Suborder B

Embioptera

Fig. 9. Tree containing all the groups in the classification proposed by Ross (1970).

Two groups of embiid genera appear as sister taxa to Conicerembia, each sup-ported by several synapomorphies (Fig. 8). The relationship between these twogroups is not resolved and they are part of a trichotomy, together withAustralembiidae+Notoligotomidae. This group is supported by the presence ofmacrosetae in the LC1 (char. 11), with parallelism in some anisembiids). Theresulting trees imply a lot of homoplasy in the mandibles (char. 8). Therefore, thischaracter exerts little influence in the results. Using an alternative scoring fordoubtful taxa (the case of Metembia ferox, Machadoembia angolica and Parembia per-sica, see section on Characters) or even deactivating the character, does notchange the results (the alternative scoring would only add to tree length).

According to Ross (1970), Enveja belongs in the “Suborder B” (monotypic).Enveja appears here grouped with other African and Asian embiids sharing thepresence of macrosetae in the basal part of the LC1 (char. 12). An alternative, onlyslightly inferior placement for Enveja would be as sister group of Embiidae+Teratembiidae+Oligotomidae+Australembiidae+Notoligotomidae, with a decreasein fit of only 0·52 (saving one step in characters 14 and 13, and adding it in 11 and12).

57ORDER EMBIOPTERA: A CLADISTIC ANALYSIS

The present analysis includes roughly half of the described embiid genera (mostof the unrepresented genera are small, highly autapomorphic and containing onlyone or two species). It is evident that the family Embiidae should be redelimited(as currently saying Embiidae is almost equivalent to saying “non-Anisembiidaeand non-Clothodidae”), but such redelimitation requires a more detailed analysis.

COMMENTS ON PREVIOUS CLASSIFICATIONS

Classification of Ross (1970)Given that Ross did not provide either a matrix or explicit character information

to support his groupings, his classification can only be judged by its fit to the pre-sent data set. The best-fit trees which contain all of Ross’ groups (Fig. 9) are 22steps longer than the best-fit unconstrained trees. These trees gives a better for 4characters, worse for 20, and implies an overall decrease in fit of 23·36 (Table 3).Those results show that Ross’ classification is far from being the most explanatoryfor the current set of data.

Classification of Davis (1938, 1940b)Two types of comparisons are relevant here: first, to what extent Davis was suc-

cessful in finding the trees which best explain his own data, compared to modernmethods of tree-searching with computer programs, and second, to what extentDavis’ results are stable to the addition of new evidence.

In his two papers, Davis used 7 (in 1938) and 15 characters (in 1940b) to sup-port his schemes of relationships. All the characters used by Davis in those twoanalyses were included in the matrix of Table 1. On the basis of the informationprovided in the tree of Davis (1938, reproduced here in Fig. 1), it is possible tounequivocally assign states for all characters for each of the terminals (see Appen-dix 3; some of Davis’ 7 characters were recoded in binary form and therefore thematrix contains 11 characters). On the basis of those data, as scored by Davis(1938), Pee-Wee produces eight trees (29 steps long). The tree proposed by Davisis only 3 steps longer, and it agrees with the best fit trees in showing as monophy-letic Metoligotoma, Notoligotoma and the species now included in Aposthonia(included by Davis in Oligotoma). In the global analysis based on the complete dataset (which includes all of Davis’ characters; Table 1), Aposthonia appears par-aphyletic because some species are more closely related to Oligotoma (as alreadydiscussed under Oligotomidae). However, it must be noted that Davis did notinclude any species of Oligotoma sensu Ross in his analysis. Davis’ tree and the globalanalysis both support the monophyly of Metoligotoma and Notoligotoma, as well as thegroup formed by Bulbocerca+Anisembia.

In 1940b, Davis presented four separate trees, indicating only the putative syna-pomorphies of the clades. Because not all the homoplastic changes are indicatedon the tree it is not possible to reconstruct Davis’ original data. In those trees, thefamily Anisembiidae, the genera Metoligotoma and Notoligotoma and Anisembia+Chelicerca (present in the trees of best fit for the complete data set) appear as mon-ophyletic; in most cases, the characters proposed by Davis (1940b) as synapo-morphies for those taxa do indeed appear as their synapomorphies in the globalanalysis.

In conclusion, the most interesting aspect of Davis’ classification is that he pro-

58 C. A. SZUMIK

posed that Metoligotoma belonged in Notoligotomidae (whereas Ross placed thatgenus in the Australembiidae) and that Embiidae (considered as a family by Ross)was a polyphyletic group. The evidence presented here clearly supports Davis’point of view.

Acknowledgements

The material used in this analysis was available as a courtesy of Abraham Willink(IFML), Axel O. Bachmann (MACN), Alcide Costa and Ricardo P. da Rocha(MZSP), G. B. Monteith and Robert J. Raven (QM), John E. Rawlins (CMNH),David G. Furth (MCZ), David A. Nickle (USNM), Randall T. Schuh (AMNH), andEdward S. Ross (CAS). James K. Liebherr (Cornell University, Ithaca), Norman I.Platnick (AMNH), Axel O. Bachmann (MACN), Elisa Angrisano (Universidad deBuenos Aires) and Abraham Willink (IFML) provided working space and help atdifferent stages of my research. Axel O. Bachmann, James M. Carpenter, Pablo A.Goloboff, Diana Lipscomb, Arturo Roig Alsina and two anonymous reviewers pro-vided useful comments on different versions of the manuscript. The encourage-ment by James M. Carpenter was greatly appreciated.

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Oligotomidae). Atti. Accad. naz. Lincei Rc., Rome, 15: 211–216.STEFANI, R. 1953b. Un particolare modo de accopiamento negli Insetti Embiotteri. Atti.

Accad. naz. Lincei Rc., Rome 15: 544–549.STEFANI, R. 1953c. Nuovi Embiotteri della Sardegna. Boll. Soc. Entomol. Ital., Genoa 83:

84–98.STEFANI, R. 1955. Revisione del genere Haploembia Verth. e descrizione di una nuova species

(Haploembia palaui n. sp.) (Embioptera: Oligotomidae). Boll. Soc. Entomol. Ital.,Genoa 85: 110–120.

Appendix 1

Species used for the cladistic analysis (including material examined, where cor-responding, using the following acronyms for Institutions: USNM, NationalMuseum of Natural History, Washington; AMNH, The American Museum ofNatural History, New York; CMNH, Carnegie Museum of Natural History,Pittsburgh; MCZ, Museum of Comparative Zoology, Cambridge; CAS, CaliforniaAcademy of Sciences, San Francisco; MACN, Museo Argentino de Ciencias Natu-rales “Bernardino Rivadavia”, Buenos Aires; IFML, Instituto y Fundacion MiguelLillo, Tucuman; MZSP, Museu de Zoologia, Sao Paulo; QM, Queensland Museum,South Brisbane). Those species for which no specimens were seen were scored onthe basis of published data.

CLOTHODIDAEAntipaluria caribbeana Ross, 1987

Venezuela; Topoparatypes USNM.Clothoda longicauda Ross, 1987

Peru; Topoparatypes USNM.

EMBIIDAEArchembia sp.

Peru; AMNH.Conicerembia tepicensis Ross, 1984

Mexico; Paratypes USNM.

60 C. A. SZUMIK

Dinembia ferruginea Davis, 1939Congo; Holotype MCZ.

Donaconethis abyssinica Enderlein, 1909Africa (no specimens seen).

Embia savignyi Westwood, 1837Egypt; USNM.

Machadoembia angolica Ross, 1952Angola; Paratypes MCZ, USNM.

Metembia ferrox Davis, 1939India; Holotype MCZ, specimens USNM.

Microembia rugosifrons Ross, 1944Peru; Holotype and Paratypes USNM.

Neorhagadochir salvini (MacLachlan, 1877)El Salvador and Honduras; USNM.

Pachylembia unicincta Ross, 1984Mexico; Paratypes USNM.

Pararhagadochir balteata Ross, 1972Brazil; Paratypes USNM.

Parembia persica (McLachlan, 1877)India (no specimens seen).

Pseudembia truncata Davis, 1939India; Holotype MCZ.

Scelembia malkini (Ross, 1952)Angola; Paratype MCZ.

INCERTAE SEDISEnveja bequaerti Navas, 1914

Congo; CAS.

AUSTRALEMBIIDAEAustralembia incompta Ross, 1963

Queensland, Australia; Topoparatypes USNM.Metoligotoma convergens Davis, 1938

New South Wales, Australia; QM.M. ingens Davis, 1936

ACT, Australia; MCZ.

OLIGOTOMIDAEAposthonia borneensis (Hagen, 1885)

Java and Thailand; MCZ, USNM.A. glauerti (Davis, 1936)

West Australia (no specimens seen).Haploembia sp.

Turkey; USNM.Oligotoma saundersii (Westwood, 1837)

India; AMNH, CMNH, MZSP, MCZ, USNM.

TERATEMBIIDAETeratembia geniculata Krauss, 1911

Argentina; IFML, MACN.

61ORDER EMBIOPTERA: A CLADISTIC ANALYSIS

ANISEMBIIDAEAnisembia texana (Melander, 1902)

Mexico; USA, AMNH, MCZ, USNM.Bulbocerca sini (Chamberlin, 1923)

Mexico (no specimens seen).Chelicerca dampfi Ross, 1944

Mexico (no specimens seen).C. davisi (Ross, 1940)

USA; USNM.C. jaliscoa Ross, 1984

Mexico; Paratypes USNM.C. maxima Ross, 1984

Mexico (no specimens seen).C. wheeleri (Melander, 1902)

Mexico; Holotype MCZ.Dactylocerca flavicollis Ross, 1984

Mexico; Paratypes USNM.Mesembia catemacoa Ross, 1984

Mexico; Paratypes USNM.M. chamulae Ross, 1984

Mexico; Paratypes USNM.M. venosa (Banks, 1924)

Cuba; Holotype MCZ.Pelorembia tumidiceps Ross, 1984

Mexico, Paratypes USNM.Stenembia perenensis Ross, 1972

Peru, Paratypes USNM.

NOTOLIGOTOMIDAENotoligotoma hardyi (Friederichs, 1914)

West Australia (no specimens seen).N. nitens Davis, 1936

New South Wales, Australia; MCZ.Ptilocerembia sp.

Java; CAS.

Appendix 2

Additional material examined.

CLOTHODIDAEAntipaluria aequicercata (Enderlein, 1912)

Colombia; USNM.A. marginata Ross, 1987

Colombia; Paratype USNM.A. panamensis Ross, 1987

Panama; Paratypes USNM.A. silvestris Ross, 1987

Venezuela; USNM.

62 C. A. SZUMIK

A. urichi (de Saussure, 1896)Trinidad; USNM.

Chromatoclothoda aurata Ross, 1987Peru; Paratypes USNM.

C. elegantula Ross, 1987Brazil; Paratypes USNM.

Clothoda nobilis (Gerstaecker, 1888)Brazil; USNM.

EMBIIDAEArchembia lacombea Ross, 1971

Brazil; Paratypes USNM.Archembia sp.

Brazil; MZSP, USNM.Brachypterembia moreliensis Ross, 1984

Mexico; Paratypes USNM.Embia mauritanica Lucas, 1849

Iraq; USNM.E. ramburi Rimsky Korsacov, 1905

France; USNM.Embia sp.

Spain; MCZ, USNM.Pachylembia chapalae Ross, 1984

Mexico; Paratypes USNM.P. taxcoensis Ross, 1984

Mexico; Paratypes USNM.Pararhagadochir christae Ross, 1972

Brazil; Paratypes USNM.P. confusa Ross, 1944

Paraguay; Paratypes and specimens MCZ.P. trinitatis (de Saussure, 1896)

Trinidad; MCZ, USNM.P. birabeni (Navas, 1918)

Argentina; IFML, MACN.P. trachelia (Navas, 1915)

Argentina; IFML, MACN.Pseudembia flava (Ross, 1943)

India; Holotype MCZ.P. immsi (Davis, 1939)

India; Holotype MCZ.

AUSTRALEMBIIDAEMetoligotoma reducta subtropica Davis, 1942

New South Wales, Australia; QM.Metoligotoma sp.

New South Wales, Australia; QM.

OLIGOTOMIDAEAposthonia ceylonica (Enderlein, 1912)

63ORDER EMBIOPTERA: A CLADISTIC ANALYSIS

Thailand, Sri Lanka, India; MCZ, USNM.A. indica (Davis, 1940)

India; MCZ, USNM.Aposthonia sp.

New South Wales, Australia; QM.Oligotoma falcis Ross, 1943

India; Paratype MCZ.O. humbertiana (de Saussure, 1896)

Sri Lanka; USNM.O. japonica Okajima, 1926

Japan; MCZ.O. nigra Hagen, 1866

Mexico, USA; AMNH, USNM.Oligotoma sp.

Queensland, Australia; QM.

TERATEMBIIDAEDiradius erba Szumik, 1991

Argentina; Holotype and Paratypes MACN.D. lobatus (Ross, 1944)

USA; Paratypes USNM.D. pallidus Ross, 1984

Mexico; Paratypes USNM.D. plaumanni (Ross, 1944)

Brazil; Paratypes MZSP, USNM.Diradius sp.

Panama, AMNH.Oligembia capensis Ross, 1984

Mexico; Paratype USNM.O. hubbardi (Hagen, 1885)

USA; AMNH, USNM.O. melanura Ross, 1944

USA; Paratypes USNM, specimens AMNH.O. mini Szumik, 1991

Argentina; Holotype and Paratypes MACN.O. versicolor Ross, 1972

Brazil; Paratype USNM.Oligembia sp.

Peru; AMNH.Teratembia banksi (Davis, 1939)

Paraguay; MCZ.

ANISEMBIIDAEChelicerca galapagensis Ross, 1966

Galapagos; Paratypes USNM.C. grandis (Ross, 1944)

Colombia; USNM.Dactylocerca ashworthi Ross, 1984

USA; Paratypes USNM.

64 C. A. SZUMIK

D. multispiculata Ross, 1984Mexico; Paratypes USNM.

D. rubra (Ross, 1940)USA; Paratypes USNM, specimens AMNH.

Mesembia aequalis Ross, 1944Brazil; Paratypes MZSP.

M. hospes (Myers, 1928)Cuba; Holotype and specimens MCZ, USNM.

Stenembia exigua Ross, 1972Brazil; Paratype USNM.

Appendix 3Matrix according to Davis’ tree (1938:265, Fig. 120), 22 taxa and 11 characters:

0 1 2 3 4 5 6 7 8 9 0

Ancestral type 0 0 0 0 0 0 0 0 0 0 0Clothoda nobilis 1 0 0 0 0 0 0 0 0 0 0Donaconethis abyssinica 0 0 0 0 1 0 1 0 1 0 0Embia sabulosa 1 0 0 0 1 0 1 0 1 0 0E. major 1 0 1 0 1 0 1 0 1 0 0Rhagadochir flavicollis 1 0 0 0 1 0 2 0 1 0 0Ptilocerembia roepkei 1 0 0 0 2 0 1 0 1 1 0Haploembia solieri 3 0 1 0 1 0 1 0 1 0 0Monotylota ramburi 3 0 0 0 1 0 1 0 1 0 0Anisembia texana 4 1 0 0 2 0 1 0 1 0 0A. sini 3 1 0 0 2 0 1 0 1 0 0A. heymonsi 2 1 0 1 1 0 1 0 1 0 0Notoligotoma hardyi 2 1 1 0 2 0 1 0 1 1 0N. nitens 4 1 1 0 2 0 1 0 1 1 0Metoligotoma convergens 3 1 1 1 1 1 2 0 1 0 1M. pugionifer 3 1 1 1 1 1 1 1 1 0 1M. pentanesiana 3 1 1 1 1 1 2 0 1 0 1Oligotoma vosselieri 2 1 0 0 1 0 1 0 2 0 0O. gurneyi subclavata 2 1 0 0 1 0 1 1 2 0 0O. g. gurneyi 4 1 0 0 1 0 1 1 2 0 0O. tillyardi 2 1 0 0 1 0 2 0 2 0 0O. glauerti 2 1 0 0 1 0 1 1 2 0 0