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ACTA GEOLOGICA HISPANICA, v. 35 (2000), nº 1,p. 149-164

AWinged Zorotypusin Miocene from the Dominican Republic (: Zorotypidae), with Discussion on Relationships of and within the Order

M.S. ENGEL(1) and D.A. GRIMALDI(2)

(1) Division of Entomology, Natural History Museum and Biodiversity Research Center, and Department of Ecology and Evolutionary Biology, 1460 Jayhawk Boulevard, Snow Hall, University of Kansas, Lawrence, Kansas 66045-7523, U.S.A. E-mail: [email protected] (2) Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192, U.S.A. E-mail: [email protected]

ABSTRACT

A new fossil zorapteran is described and figured in Miocene Dominican amber. The specimen is the first winged Zo ro t y p u s fo s s i l , and is described as Zo r otypus goe l e t i n.sp. The species is distinguished from the only other fossil zorapteran, Z. palaeus also in Do- minican amber, as well as an extant species to which it appears most similar, Z. snyd e r i . The new fossil is significant in the possession of segmented cerci, a plesiomorphic character unique for the order. The classification of the order is briefly summarized and genera pro- posed by Kuk a l ov á - P eck and Peck (1993) and Chao and Chen (2000) are newl y synonymized under Zo ro t y p u s . Phyl o genetic affin i t i e s within Zoraptera and of the order among other lower are briefly discussed. The order is considered to be most closely allied to the webspinners, order Embiidina.

Keywo rd s : Am b e r . Embiidina. Pal e o n t o l og y . Phyl og e n y. Pol yneoptera. Zoraptera.

IN T RO D U C T I O N Adults occur in two morphs within a species - eyed , winged forms (Fig. 1) or eyeless, apterous forms (Fig. 3). The order Zoraptera is a small group presently Much remains to be discovered of Zoraptera biology . Th e consisting of 32 extant and one extinct species with an es- most detailed biological accounts have been prepared for se n t i a l l y pantropical distribution (Tab le I). The order was the we s t e rn hemisphere species Z o rotypus hubbard i one of the last living insect orders to be recogn i z e d , hav- CA UDELL (1918) (Crampton, 1920; Gurne y, 1938; ing been established in 1913 by the great Italian entomol- Ri e gel, 1963; Shetlar, 1978; Rasnitsyn, 1998), Z. barberi ogist Filippo Silvestri. Zoraptera are tiny, polyn e o p t e r o u s GURNEY (1938) (Choe, 1992, 1995, 1997), and Z. gur- superfic i a l l y resembling Psocoptera. Species are ney i CHOE (1989) (Choe, 1992, 1994a,b, 1997). No gre garious, typically living in nests and under the common name has been widely applied to these insects bark of decaying logs, where they feed on fungal myc e l i a . outside of zorapterans; although several early German ar-

149 Figure 1. Dorsal habitus of holotype of Zorotypus goeleti ENGEL AND GRIMALDI n.sp.

150 ticles refer to them with the unpleasant and not ver y de- tooth, maxillary palpus five- s e gmented (Figs. 4, 7), distal sc r i p t i ve name of “Bodenläuse” (soil lice). palpal segment of labial and maxillary palpi larger than preceding palpal segments (Figs. 4, 7); prementum divi d - The phyl o genetic affinities of the order have been ed , labial palpus three-segmented; antennae nine-seg- co n t r o versial and hypotheses have shifted dramatically me n t e d , moniliform (Fig. 1); lateropleurite and lateroster- since its discover y. Numerous authors have addressed the nite differentiated in alates; six Malpighian tubules (whe r e question of Zoraptera’s position, placing it as sister gro u p ob s e r ved); wings membranous with reduced venation or to Isoptera (Boudreaux, 1979; Caudell, 1918; Crampton, wings frequently absent, forewings narro w and paddle- 1920; Wei d n e r , 1969, 1970), sister group to Isoptera + shaped due to reduced anal lobe, forewing with radial, Blattaria (Silvestri, 1913), sister group to Embiidina median, and cubital veins fused at base; hind wings small- (Minet and Bourgoin, 1986), basal to (Hen- er than forewings; all wings shed by a basal fracture; cox- nig, 1953, 1969, 1981; Kristensen, 1975), in a basal poly- ae large; metafemora stoutly exp a n d e d , with stiff spines to m y of seven neopterous orders (Kristensen, 1991, running along ventral surface (Figs. 5, 8); tarsi two- s e g- 1995), sister group to Holometabola (Rasnitsyn, 1998), me n t e d , first minute, second elongate; claws simple; ab- basal to Th ysanoptera (Karny, 1922), a suborder of Pso- domen 11-segmented; two abdominal ganglia (where ob- coptera (Karny, 1932), intermediate between Isoptera and se r ved); cerci present, short, unsegmented in living forms Psocoptera (Til ly a r d , 1926), sister group to a Derma p t e r a (F ig. 9), two- s e gmented in the fossil Z. goe l e t i (F ig. 6); + clade (Kuk a l ov á - P eck and Peck, 1993), un- ovipositor absent; female with 4-6 panoistic ovar i o l e s r e s o l ved with Orthoptera, Phasmida, and Embiidina (w here observed); male genitalia asymmetrical. (Ku k a l ov á - P eck, 1991), and near the base of Dictyop t e r a in the Pol yneoptera (Carpenter and Wh e e l e r , 1999). Al - Co m m e n t s : Pr e s e n t l y consisting of 34 species (Tabl e though Til l yard (1926: p. 125) asserted that Silvestri orig- 1) distributed pantropically, except for Z. hubbardi , whi c h in a l l y placed Zoraptera in the Ap t e r ygota owing to the ab- extends well into the Nearctic. sence of wings in the series before him, this is incorre c t since Silvestri (1913: p. 205) clearly states his belief that Zoraptera is near Isoptera + Blattaria. Fam i ly : Zorotypidae SILVESTRI, 1913 Zorotypidae SILVESTRI, 1913: 196. Herein we present the description of a second fossil species (Figs. 1-2), also in amber from the Miocene of Type genus: Zorot y p u s SILVESTRI, 1913. the Dominican Republic (Iturralde-Vinent and MacPhee, 1996, 1999). Poinar (1988) indicated this amber to be Di ag n o s i s : As for the order (see above). Lower Miocene to Upper Eocene but as discussed by Grimaldi (1995) and stratigraphically presented by Itur- GENUS: Zo ro t y p u s SI L VESTRI, 1913 ralde-Vinent and MacPhee (1996, 1999), there is no ba- sis for such an old age as Eocene or even Lower Type species: Zorotypus guineensis SI LV E S T R I , Oligocene. Unlike Z. palaeus P O I NAR (1988) the 1913, original designation. species presented below is represented by an alate fe- male. The new species is differentiated from Z. palaeus * Fl o r i d a z o ro s KU K A L O VÁ-PECK AND PECK, 1993: 340. Typ e as well as extant members of Zorotypidae. We also pre- species: Zo r otypus snyd e r i CA UDELL, 1920, monobasic and sent a brief summary of affinities among taxa within the original designation. New synonym y. order as well as the position of zorapterans among other hemimetabolous insects. * Us a z o ro s KU K A L O VÁ-PECK AND PECK, 1993: 340. Typ e species: Zo r otypus hubbardi CA UDELL, 1918, monobasic and original designation. New synonym y. SY S T E M AT I C S * Me r i d o z o ro s KU K A L O VÁ-PECK AND PECK, 1993: 341. Typ e Ord e r : Zoraptera SILVESTRI, 1913 species: Zo r otypus leleupi WEIDNER, 1976, monobasic and Zoraptera SILVESTRI, 1913: 195. original designation. New synonym y.

D i ag n o s i s : Adults minute (around 3 mm), * Bra z i l o z o ro s KU K A L O VÁ-PECK AND PECK, 1993: 342. Typ e hemimetabolous Neoptera; mouthparts mandibu l a t e ; species: Zo r otypus bras i l i e n s i s SI L VESTRI, 1946, monobasic lacinia fused to stipes, lacinia with strong inner, apical and original designation. New synonym y.

151 Figure 2. Photomicrograph of holotype of Zorotypus goeleti ENGEL AND GRIMALDI n.sp.

152 Figure 3. Scanning electron micrograph of Zorotypus juninensis ENGEL, dorsal habitus (from Engel, 2000).

* Centrozoros KUKALOVÁ-PECK AND PECK, 1993: 342. Type example, the breadth of the petiole in the forewing var i e s species: Zorotypus gurneyi CHOE, 1989, monobasic and origi- co n t i n u o u s l y across species, with no discrete separation nal designation. New synonymy. be t w een “broad” and “narro w”. Similar continuous var i a - tion occurs for their characters “breadth of C-R+RA * La t i n o z o ro s KU K A L O VÁ-PECK AND PECK, 1993: 342. Typ e area”, “length of ScP”, “length of MP+Cu”, “length of species: Zo r otypus barberi GU R N E Y , 1938, monobasic and orig- MP+CuA”, “length of “CuA1+ 2 ”, “height of CuA1+ 2 inal designation. New synonym y. cu r ve”, and “length of CuA3+ 4 ” (terms in accordance with Kuk a l ov á - Pe c k ’ s wing vein termi n o l o gy). The degree to * For m o s o z o ro s CH A O AND CHEN, 2000: 24. Type species: For - which the pterostigma is developed is continuous both mo s o z o r os newi CH A O AND CHEN, 2000, monobasic and orig- across taxa and individuals of a single species. The ven a - inal designation. New synonym y. tional homologies proposed by these authors are dubious.

Di ag n o s i s : As for the fam i l y (see above). The classification of Kuk a l ov á - P eck and Peck (1993) is based solely on wing venation and owing to the gre a t Co m m e n t s : We have taken a conservat i ve stance with variability in these characters, the classification is proba- zorapteran classification. The generic characters pro- bly unstable; additionally so given that most species are posed by Kuk a l ov á - P eck and Peck (1993) are either con- kn o wn on the basis of apterous individuals only. Th u s , tinuous across taxa or var i a b le within a given species. For owing to the extreme morph o l o gical homogeneity among

153 Figures 4-6. Characters of Zorotypus goeleti ENGEL AND GRIMALDI n.sp. 4. Ventral surface of head. 5. Metafemur. 6. Abdominal apex detailing cercal structure and setae. Scale bar = 1 mm.

154 the species and the low specific diversity of the order, the mur more pronouncedly swollen than the distal half, R1 re c o gnition of multiple genera is unneeded, parti c u l a r l y reaching to the wing apex in the forewing, and ovoid cer- since Zo ro t y p u s s. s t r . is rendered paraphyletic. Until a ci. Perhaps the most distinctive feature of the new species system can be developed and taxa diagnosed on a var i e t y is the presence of segmentation in the cerci which readily of character systems, including metafemoral spination, separates it from all other members of the order. genitalia, wing venation, structure of the cerci, chaetotaxy of the vert e x, etc. (e.g., see below “Internal Af fin i t i e s . . .” ) , De s c r i p t i o n : Alate female.- Body length (exc l u s i ve of it is most prudent not to present a new, formal classific a - antennae) 1.98 mm; antennal length 1.25 mm; forewi n g tion. All of the specific combinations proposed by length 2.43 mm; hind wing length 1.95 mm; head length Kuk a l ov á - P eck and Peck (1993) are, therefore, here re- 0.5 mm, anterior width (just anterior to compound eyes ) tu r ned to Zo ro t y p u s . 0.42 mm, posterior width 0.51 mm (from outer margi n s of the compound eyes); distance between lateral ocelli Most recently Chao and Chen (2000) proposed anoth- 0.26 mm; distance from median ocellus to lateral ocellus er genus in the fam i l y based on a single apomorph i c 0.18 mm; distance from lateral ocellus to species from Tai w an. All of the characters in their gener- 0.03 mm; pronotal length 0.38 mm, anterior width 0.4 ic description except for two are universal for the order mm, posterior width 0.29 mm; mesonotal length 0.33 (i.e., found in all zorapterans). The remaining two charac- mm, width 0.38 mm; metanotal length 0.21 mm, width ters, relativel y long first tarsomere and eloncate cerci, 0.37 mm; metafemoral width 0.23 mm, length 0.65 mm; while notewo rt hy and apomorphic are not generically dis- metatibial length 0.71 mm; metatarsomere II length 0.3 ti n c t i ve. The first feature serves to separate the Tai wa n e s e mm; abdominal length 0.86 mm; cercus length 0.14 mm. species from all other Asian species but is an isolated apo- In t e gument generally light brown to brown and smooth m o rp hy that certa i n l y renders Zo ro t y p u s pa r a p hy l e t i c , except pterothorax dark brown. pa rt i c u l a r l y when the following character is considered with a global view. The elongate cerci are by no means He a d : Head subtriangular, portion of head anterior to di s t i n c t i ve of the species from Tai w an and are identical in compound eyes slightly longer than compound eye, of shape and overall structure to those of Z. longicerca t u s equal width to anterior margin of pronotum; posterior CA UDELL from Jamaica. Zo r otypus longicerca t u s ha s border of head slightly broader than pronotum. Median the same elongate cerci (perhaps even slightly longer than ocellus set just apical to anterior tangent of compound Chao and Chen’s species) that lack a terminal spine. eyes; distance between lateral ocelli slightly greater than While certa i n l y an important new species and a val u a bl e length of first antennomere. Second antennomere wea k l y co n t r i b ution, the recognition of a monotypic genus is not cu r ved outwar d , narro w, width one-half that of first an- war ranted or advisable and For m o s o z o r os newi is trans- tennomere; combined length of second and third anten- fe r red to the genus Zo ro t y p u s [Zo r otypus newi (C H A O nomeres slightly less than that of first, about as long as AND CHEN), new combination]. that of fourth; second antennomere about as long as third. Inner margins of compound eyes emarginate above, two facets deep by five facets long, emargination on posterior Zo r otypus goe l e t i n. s p . half of eye. Integument just posterior to compound eyes Figures 1-2, 4-6 wea k l y bulging, creating an exc e e d i n g l y short posterior co l l a r . Setae of ve rt ex mostly restricted to posterior bor- Di ag n o s i s : The new fossil would apparently belong to ders of head, a few fine, scattered setae between lateral a group of wes t e r n hemisphere species in which the row ocelli. Epicranial suture exc e e d i n g l y faint. Gena with a of spines on the ventral surface of the metafemur is inter- fe w long setae just below compound eyes. Setal pattern of rupted (Fig. 5); alterna t i vel y this species could be sister to head depicted in figure 1. Mandible apparently simple all living species (see discussion of relationships below) . without subapical teeth, apex pointed; lacinia with point- Zo r otypus goe l e t i can be readily distinguished from the ed apex, and possibly with subapical tooth (difficult to on l y other fossil zorapteran, Z. palaeus, which has a sin- see); galea with fringed apex as in living species; maxil- gle, uninterrupted row of such spines, more numerous la r y palpal segments 2, 3, and 5 elongate, segments 1 and spines closely spaced along the distal third of the metafe- 4 about as long as apical width, segment 2 not widened at mu r , and a basal-most spine which is not signific a n t l y ap e x and about as long as segment 3, segment 3 slightly longer than the others. Zo r otypus goeleti most closely re- wider at apex than at base, segment 5 slightly longer than se m b les Z. snyd e r i CA UDELL (1920), which differs in pr o ximal segments, gently swollen on ventral margi n , ha ving a quadrate pronotum, the basal half of the metafe- more densely setose than preceding segments; glossa

155 Figure 7. Ventral surface of head of Z o rotypus juninensis ENGEL, scanning electron micrograph (from Engel, 2000); note that the right labial palp (left in the micrograph) is broken and missing its terminal segment while the right palpus (left in the micrograph and with its terminal segment labeled) is oddly appressed to the ventral side of the head and slightly broken between the terminal seg- ment and the segment immediately preceding it. Ant = , G = glossa, LP = labial palp, Md = mandible, MP = maxillary palp, Mx = maxilla. sh o r t and rounded with numerous, fine scattered setae; cept for middorsal line; setae long and anteriorly directed labial palpal segments 1 and 3 elongate, segment 1 except posterolateral-most setae, which are directed longest, narro w, and not widening towards apex, segm e n t posterad with two minute setae just anterior to this. 2 short about one-third length of segment 1, segment 3 Mesonotum and metanotum each about as broad as ante- sl i g h t l y shorter than segment 1, gently swollen on ven t r a l rior width of pronotum and slightly broader than long. su r f ace, distal segment more densely setose than preced- For e wing venation faint with most veins represented by ing segments. Mouthparts depicted in figure 4. fuscous lines; pterostigma exc e e d i n g l y lightly sclerotized; C+Sc becoming faint by base of pterostigma; R1 disap- Th o ra x : Pronotal length subequal to anterior width (re- pearing by base of pterostigma; Rs reaching nearly to wing fer to metrics above); posterior width slightly less than an- ap e x before becoming slightly diffuse around apex of terior width, therefore pronotum is wea k l y constricted pterostigma; r-m present, ver y short, equal in length to po s t e r i o r l y. Two crescentic ridges running along anterior basal section of Rs between stem of R and juncture with r- third of pronotum (Fig. 1). Longest setae of pronotum on m; M reaching to posterior wing margin and equidistant lateral margins, smaller setae scattered on pronotum ex- from wing apex as is R1; CuA1 present and reaching to

156 Figures 8-9. Characters of Zorotypus juninensis ENGEL, scanning electron micrograph (from Engel, 2000). 8. Metafemur. 9. Abdo- minal apex, showing detail of unsegmented cercus.

157 posterior wing margin just beyond midwing; CuA2 pr e s e n t overlapping slightly (Figs. 1-2). The ventral surface is al- as a short stub in basal third of wing; setae on wing mem- so easily visible except for a few areas obscured by the brane numerous and short, not concentrated in any porti o n le gs. The ventral surface of the head is parti c u l a r l y wel l of the wing. Hind wing with M+R running in anterior half, positioned so that the mouthparts can be examined with both R and M reaching respective wing margins; Cu ab- ease (Fig. 4). sent. Marginal hairs on both fore and hind wings notice- ab ly longer medially on posterior margin; lengths gre a t e r than twice that of setae on wing membrane. Legs with DI S C U S S I O N scattered setae of moderate length intermi x ed with much sh o r ter and more numerous setae. Metafemur about three In t e r nal Af f inities of Zo ro t y p u s times longer than broad, not noticeably more swollen to- wards base than towards apex, gently but rapidly tapering A specific - l e vel phyl og e n y of Zoraptera has never at apex to join metatibia; ventral surface with four short, been attempted and we do not underta k e a cladistic analy- st i f f spines (Fig. 5); proximal spine about one-third along sis of the species here. This is due to the difficulty in length of metafemur, distinctly separated from remainder comparing across all of the presently known taxa for two of spines beginning around midpoint. Proximal spine no- reasons; 1) most species are presently known from only ti c e a b ly longest, apical series of spines of roughly equal one morph (apterous vs. winged) or sex; and 2) a wor l d length except distal spine slightly smaller. Metatibia re vision of the order is needed, since species are mostly sl i g h t l y longer than metafemur. Metatarsomere II long, kn o wn from a few, isolated specimens that are not wel l ne a r l y one-half length of metatibia. described and for which material is difficult to obtain (es- pe c i a l l y for African or southeast Asian species). Despite Ab d o m e n : Posterior margin of each tergite with a sin- these difficulties, some comments can be made on possi- gle, transverse row of setae of moderate length, exc e p t ble affinities of species or species groups within the or- such hairs longer laterally; T9 and T10 with two distinct de r , to provide a context for the new fossil species, Z. apical setae and shorter laterals; T10 broadly rounded api- goe l e t i . This discussion will perhaps indicate characters ca l l y; sterna with scattered short setae; medio-apical mar- that could be explored more fully for understanding gin of distal sternum shallowl y and narrow l y emargi n a t e . Zo ro t y p u s phy l og e n y. Cerci not ovoi d , instead strongly narro wed distad and api- ca l l y pointed (Fig. 6); two- s eg m e n t e d , slight constriction The new species presented above would appear to just beyond midpoint with distinct annular ring; with nu- belong to a group of at least four we s t e rn hemisphere merous long preapical setae, of these outer-m e d i o l a t e r a l species in which the row of metafemoral spines is inter- seta longest; apical seta just longer than cercus, slightly rupted. In some species (e.g., Z. barberi, Z. sny d e r i, Z . sh o r ter than outer-mediolateral seta. s h a n n o n i) the basal-most spine, or set of spines, is c l e a r ly separated from the apical series, whereafter the Male and apterous morphs. Unknown. spines are eve n ly spaced. The altern a t ive type (e.g., Z . g u r n ey i, Z. neotro p i c u s, Z. leleupi, Z. palaeus, Z. we i d- Ho l o t y p e : Alate female; Miocene, Dominican Repub- n e r i, Z. hamiltoni, Z. cra m p t o n i) have all the spines oc- lic, AMNH, DR-14-34; from north e r n mines (specific c u rring at regular intervals. It is unclear whether this mine unknown). Labeled “Holotype, Zo r otypus goe l e t i character is congruent with a species group that can be Engel & Grimaldi”. In amber fossil collection, Depart- d e fined by the coiled intromittent organ positioned be- ment of Entomology , American Museum of Natural His- t ween the arms of the bifurcated basal plate in males to r y. ( N ew, 1978; Choe, 1989); the inclusion of Z. sny d e r i within this group of “coiled” species appears to be the Et y m o l o gy: The specific epithet is a patronymic hon- o n ly exception. With respect to the metafemoral stru c- oring a ver y generous benefactor of the American Muse- ture Z. go e l e t i falls into the first categ o ry and can be um of Natural History, Mr. Robert G. Goelet. r e a d i ly distinguished from the only other fossil zo- rapteran, Z. palaeus, which belongs to the other. T h e Pre s e r v a t i o n : The specimen is completely preserved presence of CuA1 in the forewing resembles the state in a piece of dark yel l o w amber originally 15 x 12 mm, 4 seen in Z. gurney i, Z. bra s i l i e n s i s, Z. leleupi, and Z . mm thick (oval), even t u a l l y embedded in epoxy and hu b b a rd i , but all of these species have ovoid cerci and trimmed to better see details of the inclusion. The entire- metafemoral structures of both categories as do many of ty of the dorsal surface is fully visible, with the wings the other neotropical species. In Z. go e l e t i R1 does not

158 SP E C I E S DI S T R I BU T I O N Zo r otypus barberi GU R N E Y Costa Rica (Cocos), Pan a m á Zo r otypus bras i l i e n s i s SI LV E S T R I Br a z i l Zo r otypus bux t o n i KA R N Y Sa m o a Zo r otypus caudelli KA R N Y Indonesia (Sumatra) Zo r otypus ceyl o n i c u s SI LV E S T R I Sri Lanka Zo r otypus congen s i s RYN - TO U R N E L Za i r e Zo r otypus cram p t o n i GU R N E Y Gu a t e m a l a Zo r otypus delamarei PAUL I A N Ma d a ga s c a r † Zo r otypus goe l e t i ENGEL AND GRIMALDI Dominican Republi c Zo r otypus guineensis SI LV E S T R I Gu i n e a Zo r otypus gurneyi CH O E Pan a m á Zo r otypus hamiltoni NE W Co l o m b i a Zo r otypus hubbardi CAU D E L L United States Zo r otypus huxleyi BO L I V AR Y PIELTAIN AND CORONA D O Brazil, Per u Zo r otypus javanicus SI LV E S T R I Indonesia (Java) Zo r otypus juninensis EN G E L Per u Zo r otypus lawren c e i NE W Christmas Island Zo r otypus leleupi WE I D N E R Ga l a p a g o s Zo r otypus longicerca t u s CAU D E L L Ja m a i c a Zo r otypus manni CAU D E L L Bo l iv i a Zo r otypus medoensis HWA N G Tib e t Zo r otypus mexicanus BO L I V AR Y PIELTAI N Mex i c o Zo r otypus neotrop i c u s SI LV E S T R I Costa Rica Zo r otypus newi (C H A O AND CHEN) Tai wa n † Zo r otypus palaeus PO I NA R Dominican Republi c Zo r otypus philippinensis GU R N E Y Ph i l i p p i n e s Zo r otypus shannoni GU R N E Y Br a z i l Zo r otypus silvestrii KA R N Y Indonesia (Mentawai ) Zo r otypus sinensis HWA N G Tib e t Zo r otypus snyd e r i CAU D E L L Jamaica, United States Zo r otypus swez ey i CAU D E L L Ha wai i Zo r otypus vinsoni PAUL I A N Ma u r i t i u s Zo r otypus wei d n e r i NE W Br a z i l Zo r otypus zimmermani GU R N E Y Fij i

Table 1. List of presently known Zoraptera species (updated from Choe, 1992), alphabetical by species*. Daggers indicate fossil taxa. A species, excluded from this list, is recorded from Angola under the nomen nudum Zorotypus machadoi (Delamare-Deboutteville, 1951). * The names Z. chinensis given in Hwang (1974), Z. hubbardis given in the title of Crampton (1920), and Z. guineenis listed in Choe (1992) are all lapsus calami for the species Z. sinensis, Z. hubbardi, and Z. guineensis, respectively. Abundant material, perhaps of Z. barbieri, is kown from the Dominican Republic (M. Ivie, pers. comm.)

159 the above similarities may also prove to be plesiomor- phic or conve rgent.

Additional characters appear to be useful in picking out species groups but are not known for enough taxa for consideration at this time. The presence/absence of a medio-apical cleft on S8 in females; the shape, posi- tion, and size of the anterior arms on S9 in females; the general structure of the male genitalia; the relative sizes and shapes of the maxillary and labial palpi; and the structure of the mandibles all deserve detailed s t u d y.

Or dinal Ph yl o gen y

As was alluded to in the Introduction, the phyl og e n e t - ic placement of the Zoraptera has been of considerable in- terest and legitimate debate. For our discussion we shall on l y briefly touch on the more significant studies ad- dressing zorapteran affinities.

Hennig (1969, 1981) considered the Zoraptera to be the basal-most Paraneopteran lineage, based on the de- velopment of the “areola postica” in the forewing, the reduced number of tarsomeres, and the reduced number of abdominal ganglia. Later, Minet and Bourgoin (1986) associated the Zoraptera with the order Embiidina based on the development of the metafemur and tibial muscu- lature (both orders share a number of characteristics Figure 10. Preferred hypothesis of phylogenetic relationships which are detailed further below). This is perhaps the within . best supported position for Zoraptera among other in- sects. Most recently, Rasnitsyn (1998) hypothesized that Zoraptera is allied with the extinct order Caloneurodea reach the wing apex as a well defined vein and instead and represents the sister group of the Holometabola. His becomes entirely fuscous and diffuse at the base of an position was based on evo l u t i o n a ry hypotheses of par- ex c e e d i n g ly faint pterostigma, differing from species ticular characters that were ove r l o o ked because of their such as Z. gurney i, Z. sny d e r i, and Z. caudelli in this re- apparent lability, or homoplasy. Most notably, the medi- spect. The new species also lacks vein Cu in the hind al mesocoxal articulation was used along with the pres- wing and has the fine hairs noticeably longer on the ence of the “discrimen” (impressed line indicating ster- posterior margins of the fore and hind wings. T h e nal invagination) to unite Zoraptera with the pronotum is slightly narr owed posteriorly, somewh a t e n d o p t e rygote orders. Although Zoraptera lack a medial similar to the African species Z. conge n s i s, Z. dela- m e s o c oxal articulation, the presence of a medial meso- ma re i , and Z. vinsoni, but the structure of the cerci and c oxal swelling in was argued to be an incipient precur- metafemur can quickly distinguish Z. go e l e t i from these sor to a true articulation and homologous with the art i c- species. The second metatarsomere is elongate in Z . ulation in Holometabola. Rasnitsyn’s own ev i d e n c e , go e l e t i (and in Z. neotro p i c u s), being nearly one-half h oweve r, suggests a gladular function for this stru c t u r e the length of the metatibia. This differs from the United by the presence of secreted substances and numerous se- States species in which this structure is only about one- c r e t o ry cells under the integumental surface. Thus, the third the length of the metatibia. If Z. go e l e t i is sister to h o m o l ogy between the medial mesocoxal articulation in the remainder of Z o ro t y p u s species owing to the ple- Holometabola and the glandular swelling in Zoraptera s i o m o rphic segmentation of the cercus, then some of does not seem well justified. The discrimen of Rasnitsyn

160 (1998), howeve r, seems more well founded and may tru- c h a e ognatha, Ephemeroptera, Coleoptera, some or- ly represent remnants of an invaginated sternum. Take n thopterans, and Plecoptera. This is an ex c e p t i o n a l ly gen- in context with the numerous characters uniting Zo- eralized character based on shape which is ex c e e d i n g ly raptera with lower Neoptera, howeve r, it must be inter- d i fficult to homologize. The Zoraptera were ex c l u d e d preted as conve rgent. Many of the musculature charac- from a clade containing the Orthopteroidea, Plecoptera, ters discussed by Rasnitsyn (1998) are homoplastic and Embiidina on the basis of characters of antennal cir- across all of the insect orders (occurring in groups as di- c u l a t o ry development, which are actually unknown in verse as Ephemeroptera and Coleoptera) while the fossil the Zoraptera, and by the free premental lobes. In their orders with which he attempts to unite major groups are “minimal incongruence” tree Dermaptera and Dicty- u n k n own for these characters. Also, some of the ex t e r- optera are united by the fusion of MA with the radial nal characters he used are ambiguous for the extinct or- system in the forewing (a character adapted from ders, and the position of the Caloneurodea is even of se- Kuk a l ov á - P eck, 1985; Kuk a l ov á - P eck and Peck, 1993). rious question. Sharov (1966) and Rasnitsyn (1980) While this coding is accurate for Dictyoptera and the fu- both considered the order to show affinity with neu- sion of R+MA+Cu is accurate for Zoraptera, the coding ropteroids and the Coleoptera while Kuk a l ov á - Pe c k of this feature for Dermaptera needs revision. This char- (1991) considered it to be allied with the hemipteroid acter was coded as present for Dermaptera even though a s s e m blage. The order is perhaps best classified as a rel- the hemelytra of lack venation of any sort. T h e a t ive of the extinct order (Carp e n t e r, coding was based on Kuk a l ov á - P eck and Pe c k ’s (1993) 1992; Burnham, 1984: This order is itself an ex t r e m e ly a s s e rtion that the extinct order are prim- d i ffuse, cert a i n ly paraphyletic group.) and no characters i t ive relatives of or stem-group dermapterans. This posi- c o n c l u s ive ly support Caloneurodea as basal to the tion is not entirely justifi e d, and has been based on the Holometabola. c o nve rgence of a large anal fan in hemelytrous insects [e.g., Umenocoleidae, placed in Protelytroptera by A recent comprehensive analysis of hexapod ordinal Kuk a l ov á - P eck (1991), but based on abundant Creta- relationships has been presented by Carpenter and ceous material they are actually roaches (personal Wheeler [1999: a summary of Wheeler et al. (in press)], obs.)]. Large, fa n - l i ke anal lobes occur throughout the based on a combination of molecular characters and pre- insects, correlated with tegminous or elytrous forew i n g s v i o u s ly published morp h o l ogical and ethological char- ( O rthoptera, Blattaria, Mantodea, Coleoptera), so bas- acters. In their simultaneous analysis cladogram Zo- ing a protely t r o p t e r a n - d e rmapteran relationship on this raptera is placed in a poly t o my with Gry l l o blattaria and feature is unsupported. There are indeed fossils wh i c h D e rmaptera basal to the Dictyoptera, and the position of can be positive ly assigned to Dermaptera which do pos- the order within the Po lyneoptera corroborates the con- sess some venational details in the hemelytra; these be- clusions of Boudreaux (1979) and Minet and Bourg o i n ing members of the extinct suborder A r c h i d e rm a p t e r a (1986) even though disagreeing in finer details of rela- ( M a rt y n ov, 1925; Bey - B i e n ko, 1936; Vi s h n i a kova , tionship. In 21 of their 25 analyses the Zoraptera are 1980; Carp e n t e r, 1992). In the A r c h i d e rmaptera, howev- s u p p o rted as sister group to the Dermaptera. This nove l e r, MA and R are not fused in the forewing at any point. grouping is not well support e d, (a conclusion eve n D e rmaptera, therefore, should be considered as lacking reached by Carpenter and W h e e l e r, op. cit.) and corr e c- this feature, thereby failing to link this order with Zo- tions to their morp h o l ogical interpretations might ex- raptera or Dictyoptera. plain the result and perhaps revise future analyses. Con- ical coxae [a character extracted from Boudreaux Five characters were proposed that specific a l l y united (1979)] are used to unite Zoraptera, Gry l l o bl a t t a r i a , De r maptera and Zoraptera in some analyses. The charac- D e rmaptera, and some Dictyoptera; yet, the Clothodi- ters optimized at this node, however , are quite homopla- dae, a basal or the most basal group of Embiidina, show sious. Not discussed by Carpenter and Wheeler (1999) is a similar coxal structure to that seen in Zoraptera (per- the presence of maternal brood care (a groundplan feature sonal obs.). This character needs to be more carefully of Embiidina), and the reduction of male gonostyli, whi c h explored across the insect orders since it does seem to is present in Embiidina and Plecoptera. With regard to the appear outside of the aforementioned six groups. The re- remaining three characters, Zoraptera do not share states duced indirect flight musculature, which is incomplete- with Dermaptera nor any of the other closely allied orders ly surveye d, and a discoid pronotum are the only char- (i.e., Dictyoptera and Gryl l o b lattaria). The first of these is acters which appear to unite this purp o rted clade. A the development of the ovipositor which they coded as discoid pronotum, howeve r, also occurs in some A r- present in the Dictyoptera (actually, it is vestigial and in-

161 te r nal in the living Blattodea and Isoptera) and Gryl - found in webspinners and zorapterans since the highly de- lo b lattaria, and vestigial in Dermaptera [where it is actu- veloped metafemoral muscles in Orthoptera are tibial le- al l y more developed than in most Dictyoptera, parti c u l a r - vators, versus tibial depressors in the Embiidina and Zo- ly in the Pygidicranidae (cf. Fig. 23.5, Rentz and Kevan , raptera. 1991)]. The Zoraptera, however , have lost the ovi p o s i t o r , in this way again resembling the Embiidina and Ple- 4. For e and Hind Wings Narrow, Paddle-Shaped (re- coptera. The tarsomere count is autapomorphic for Zo- duction of anal regi o n ) : The anal region of the wing is raptera, which has two tarsomeres; Dermaptera and Em- gre a t l y reduced in both Embiidina and Zoraptera result- biidina have three; and the Dictyopteran orders have five ing in a distinctive “paddle-shaped” wing. This is also tarsomeres each in the groundplan (reduced to four in all so m e what true in Plecoptera but in forewings only, and but the most basal Isoptera). Lastly, the development of for all except the most basal Isoptera. the antennal circulatory organs is completely unknown for Zoraptera and thereby cannot convi n c i n g l y serve to 5. A p t e rous Morphs: Zorapterans occur in both unite it with any other order. winged and apterous morphs while the same condition occurs among some Embiidina males. Embiidina females Ca r penter and Wheeler (1999) did not advocate a are always wingless. [D e r maptera + Zoraptera] clade even though this gro u p - ing appeared in their simultaneous analysis. As we have 6. Dehiscent Win g s : The orders Embiidina, Zoraptera, di s c u s s e d , reconsidering the distribution and polarities of and Isoptera all possess dehiscent wings that are shed by a ma n y of the non-molecular characters they cited may ac- basal fracture, although the basal fracture in Isoptera dif- count for such a novel result. We suggest that the Embi- fers signific a n t l y from that seen in the previous two orders. idina and Zoraptera share numerous synapomorp h i c traits, that Pol yneopteran phyl og e n y more closely resem- 7. Gregarious (maternal behavior): Both Embiidina bles that depicted in figure 10, and that future cladistic and Zoraptera are gre garious. Neither possesses societies a n a lyses will like ly recover such a grouping. Such which can be classified as social (e.g., communal or bet- sy n a p o m o r phies for [Zoraptera + Embiidina] include: ter) but both do exhibit an extended maternal care.

1. Loss of Gonostyli: As noted above, the loss of the 8. Tars o m e r e Reduction: Both orders share a reduced gonostylus in the male genitalia is also a feature of Em- tarsomere count; two in Zoraptera, three in Embiidina. biidina and Plecoptera, as well as Zoraptera and This latter number is homoplasious and shared with De r maptera. No other insect orders have this in their De r maptera. groundplan.

2. Cerci Reduced: The reduced number of cercal seg- CO N C L U S I O N S ments is found in both orders, and two- s e gmented cerci in Z. goe l e t i is parti c u l a r l y tantalizing given the presence of With the sole exception of the interesting and highly two - s e gmented cerci in Embiidina. Embiidina are autapo- si g n i f icant character of segmented cerci, Z. goe l e t i in Do- m o rphic for asymmetrically-shaped cerci (although minican amber is ver y similar to living species. If the seg- some, like Oligotomidae, may plesiomorph i c a l l y have mented cerci are viewed as a retention of the plesiomor- symmetrical cerci). A definite annular ring occurs on the phic condition, and unsegmented cerci as a cerci of Z. goe l e t i be yond their midpoint (refer to the de- sy n a p o m o rp h y linking all modern species, then the other scription above). Examination of some living zorapterans features that the fossil shares with some of the livi n g failed to find any evidence of segmentation and thus Z. species (e.g., pattern of metafemoral spination) would be goe l e t i li ke l y represents the sister group to all other due to conver gence. The new fossil provides little evi - Zo ro t y p u s species (see Conclusions below). dence to address phyl o genetic relationships of the Zo- raptera to other polyneopterous orders, although its 3. Enlarged Metafem o ra : The tibial depressor muscles mo d e r nity probably reflects the antiquity of the order. Al - and metafemora are gre a t l y developed and expanded in so, a reconsideration of the morph o l o gical and behavi o r a l both Embiidina and Zoraptera, a derived feature unique to characters used by Carpenter and Wheeler (1999) and these two orders. A similar but conver gent condition is Wheeler et al. (in press) as well as consideration of addi- seen in saltatorial groups (e.g., Orthoptera). The or- tional characters, indicates a sister-g roup relationship of thopteroid condition is easily distinguished from the state the zorapterans and webspinners.

162 Zorapterans and embiids share several ver y interest- Bu r nham, L., 1984. Les insectes du Carbonifère supérieur de ing biological traits (dehiscent wings with reduced ven a - Montceau-les-Mines. I. L’ordre des Caloneurodea. Ann. Pa- tion, and aptery), which are obvi o u s l y adaptations for a leontol., 70, 167-180. cr yptic way of life; this lifestyle probably fostered devel - Ca rp e n t e r , F.M., 1992. Superclass . In Kaesler, R.L. opment of social behavi o r , analogous to the situation in (ed.). Treatise on Inver tebrate Pal e o n t o l og y , Par t R, A rt h r o- te r mite colonies. Given the rich pre-Cenozoic record of poda 3-4, 1-655, Geol. Soc. Am e r ., Boulder. orders in the Pol yneoptera (e.g., Carpe n t e r , 1992), it Ca rp e n t e r , J.M., Wh e e l e r , W.C., 1999. Cladística numérica, análi- should be expected that pre-Cenozoic fossils of Zoraptera sis simultáneo y fil o genia de hexápodos. Bol. S.E.A., 26, and Embiidina will even t u a l l y be found. These fossils will 333-346. ce rt a i n l y be more phyl og e n e t i c a l l y significant than Z. Caudell, A. N ., 1918. Zo r otypus hubbardi , a new species of the goe l e t i . Unfortu n a t e l y, outside of an undescribed, Perm i - order Zoraptera from the United States. Canad. Entomol., an fossil of a putative embiid (Kuk a l ov á - P eck, 1991), 50(11), 375-381. there are no Mesozoic or Paleozoic fossils of the Zo- Caudell, A. N ., 1920. Zoraptera not an apterous order. Proc. ent. raptera or Embiidina, only embiids in Baltic and Domini- Soc. Wash., 22(5), 84-97. can amber. Kuk a l ov á - P eck (1991: Fig. 6.19B) figured an Chao, R.-F., Chen, C.-S., 2000. For m o s o z o r os newi , a new undescribed fossil from the Per mian of the Urals, whi c h genus and species of Zoraptera (Insecta) from Tai w an. Pan - has virtu a l l y no thoracic differentiation, and a pair of ter- Pac. Entomol., 76(1), 24-27. minal appendages that are either cerci or male genitalia. Choe, J.C., 1989. Zo r otypus gurneyi , new species, from Pan a m a The right appendage is either smaller or else has only the and redescription of Z. barberi G u rn ey (Zoraptera: Zorotyp- base intact (e.g., if genitalic, the gonostylus is lost); ap- idae). Ann. ent. Soc. Am e r ., 82(2), 149-155. parent asymmetry has led Kuk a l ov á - P eck to conclude that Choe, J.C., 1992. Zoraptera of Panama with a review of the the fossil is of embiid affinities. Wing venation is wea k l y morphology, systematics, and biology of the order. In D. pr e s e r ved , and what little venation that is figured shows Quintero, A. Aiello (eds.). Insects of Panama and no diagnostic features for Embiidina. When the Zo- Mesoamerica: Selected Studies, 249-256, Oxford Univ. raptera-Embiidina diver gence took place is, thus, purely Press, Oxford. conjectural at this point. Choe, J.C., 1994a. Sexual selection and mating system in Zo r otypus gurneyi Choe (Insecta: Zoraptera). I. Dominance hi e r a r c h y and mating success. Behav. Ecol. Sociobiol., 34, ACK N OW L E D G E M E N T S 87-93. Choe, J.C., 1994b. Sexual selection and mating system in MSE was supported by the generosity of Mr. Robert G. Zo r otypus gurneyi Choe (Insecta: Zoraptera). II. Determi - Goelet, Trustee and Chairman Emeritus of the American Muse- nants and dynamics of dominance. Behav. Ecol. Sociobiol., um of Natural History board of trustees, who also provi d e d 34, 233-237. funds for the acquisition of this wonderful specimen. His sense Choe, J.C., 1995. Courtship feeding and repeated mating in of museum priorities - collections and collection-based research Zo r otypus barberi (Insecta: Zoraptera). Behav., - is deeply appreciated. We are grateful to Dr. Edward S. Ross 49(6), 1511-1520. for the donation of specimens of Clothodidae and informa t i o n Choe, J.C., 1997. The evolution of mating systems in the Zo- on Embiidina; to Molly G. Rightmyer for commenting on the raptera: Mating variations and sexual conflicts. In J.C. manuscript and assistance in preparing the figures; to Drs. Niels Choe, B. J. Crespi (eds.). The Evolution of Mating Systems P. Kristensen and J. Minet for val u a b le revi e ws of the manu- in Insects and Arachnids, 130-145, Cambridge Univ. Press, script; and, even though we could not agree with their conclu- Cambridge. sions, to Drs. Ward C. Wh e e l e r , Michael Whiting, Quentin D. Crampton, G.C., 1920. Some anatomical details of the remark- Wh e e l e r , and James M. Carpenter for sharing with us their pa- ab le winged zorapteron, Zo r otypus hubbardi s [sic] Caudell, per on hexapod phyl og e n y. with notes on its relationships. Proc. ent. Soc. Wash., 22(5), 98-106. D e l a m a r e - D e b o u t eville, C., 1951. Caracteres sex u e l s RE F E R E N C E S cephaliques de type intermédiaire chex les mâles aptères de Zo r otypus Delamarei [sic] Paulian. Le Naturaliste Mal- Bey - B i e n k o, G.Y., 1936. Insectes dermaptères. In Fauna of the gache, 3, 37-38. U.S.S.R., 1-239, Acad. Sci. Press, Moscow. Engel, M.S., 2000. A new Zo ro t y p u s from Per u, with notes on Boudreaux, H.B., 1979. Ar thropod Phyl og e n y with Special Ref- related neotropical species (Zoraptera: Zorotypidae). J. erence to Insects, Wil e y-Interscience, viii+320 pp., New Yor k . Kansas ent. Soc., 73(1), 10-19.

163 Grimaldi, D.A., 1995. The age of Dominican amber. In K.B. Or thoptera (s.l.), Odonata, Neuroptera. Bull. Acad. Sci. U. Anderson, J.C. Crelling (eds.). Am b e r , Resinite, and Fos s i l R. S. S., 19, 569-598. Resins, 203-217, Am e r . Chem. Soc., Washington. Minet, J., Bourgoin, T., 1986. Phyl o génie et classification des Gu rn e y, A. B ., 1938. A synopsis of the order Zoraptera, with He xapodes (Arthropoda). Cah. Liaison, O. P. I. E., 20(4), notes on the biology of Zo r otypus hubbardi Caudell. Proc. 23-28. ent. Soc. Wash., 40(3), 57-87. Ne w, T.R., 1978. Notes on neotropical Zoraptera, with descrip- Hennig, W., 1953. Kritische Bemerkungen zum phyl og e n e t i s - tions of two new species. Syst. Entomol., 3, 361-370. chen System der Insekten. Beitr. Entomol., 3, 1-85. Poi n a r , G.O., Jr., 1988. Zo r otypus palaeus, new species, a fossil Hennig, W., 1969. Die Stammesgeschichte der Insekten. Fra n k - Zoraptera (Insecta) in Dominican amber. J. New York ent. fu r t am Main, Kramer, 436 pp. Soc., 96(3), 253-259. Hennig, W., 1981. Insect Phyl og e n y. New York, Wil ey - I n t e r - Rasnistyn, A. P ., 1980. Origin and evolution of hym e n o p t e r o u s science, xxii+514 pp. insects. Tr. Paleontol. Inst. Akad. Nauk SSSR, 174, 1-190. Hw ang, F.-S., 1974. Zo r otypus chi n e n s i s [sic], a new species [In Russian]. from China. Acta Entomol. Sinica, 17(4), 423-427. [In Chi- Rasnitsyn, A. P ., 1998. On the taxonomic position of the insect nese, with English summary] . order Zorotypida = Zoraptera. Zool. Anz., 237, 185-194. It u rr a l d e - V inent, M.A., MacPhee, R.D.E., 1996. Age and paleo- Rentz, D.C . F ., Kevan, D.K.McE., 1991. Dermaptera (Earwigs). ge o graphical origin of Dominican amber. Science, 273, In Naumann, I.D. (ed.). The Insects of Australia: A Text b o o k 1850-1852. for Students and Research Wor k ers, 360-368, Ithaca, Cor- It u rr a l d e - V inent, M.A., MacPhee, R.D.E., 1999. Pal e og e o gra - nell Univ. Press. ph y of the Caribbean Region: Implications for Cenozoic Ri e gel, G.T., 1963. The distribution of Zo r otypus hubbardi (Z o - bi og e o gra p h y. Bull. Am e r . Mus. Nat. Hist., 238, 1-95. raptera). Ann. ent. Soc. Am e r ., 56, 744-747. Ka r ny, H.H., 1922. Zorapteren aus Süd-Sumatra. Treubia, 3(1), Sh a r o v, A.G., 1966. The position of the orders Glosselyt r o d e a 14-37. and Caloneurodea in the systematics of insects. Pal e o n t o l . Ka r ny, H.H., 1932. Psocoptera. Insects of Samoa, 7(4), 117-129. Zh., 1966, 84-93. [In Russian]. Kristensen, N.P ., 1975. The phyl og e n y of hexapod “orders”. A Sh e t l a r , D.J ., 1978. Biological observations on Zo r otypus hub- critical revi e w of recent accounts. Z. zool. Syst. Evol - ba rd i Caudell (Zoraptera). Entomol. News, 89(9-10), 217- Forsch., 13, 1-44. 223. Kristensen, N.P ., 1991. Phyl og e n y of extant hexapods. In Nau- Si l v estri, F., 1913. Descrizione di un nuovo ordine di insetti. mann, I.D. (ed.). The Insects of Australia: A Textbook for Boll. Lab. Zool. Gen. Ag r. Por tici, 7, 193-209. Students and Research Wor k ers, 2nd edition, 125-140, Itha- Si l v estri, F., 1946. Descrizione di due specie neotropicali di ca, Cornell Univ. Press. Zo ro t y p u s (Insecta, Zoraptera). Boll. Lab. Ent. Ag r. Port i c i , Kristensen, N.P ., 1995. For ty yea r s ’ insect phyl o genetic system- 7, 1-12. atics: Hennig’s “Kritische Bemerkungen...”, and subsequent Til ly a r d , R.J., 1926. The Insects of Australia and New Zealand. de velopments. Zool. Beitr., N.F ., 36, 83-124. Sy d n e y, Angus and Robertson Ltd., xi+[5]+560 pp. Kuk a l ov á - P eck, J., 1985. Ephemeroid wing venation based up- Vis h n i a k ova, V.H., 1980. Earwigs from the Upper of the on new gigantic and basic morph o l - Karatau Range (Insecta, Fo r ficulida). Paleontol. Zh., ogy , phyl og e n y, and metamorphosis of pterygote insects 1980(1), 78-94. [In Russian]. (Insecta, Ephemerida). Canad. J. Zool., 63, 933-955. Wei d n e r , H., 1969. Die Ordnung Zoraptera oder Bodenläuse. Kuk a l ov á - P eck, J., 1991. Fossil history and the evolution of Entomol. Z., 79, 29-51. he xapod structures. In Naumann, I.D. (ed.). The Insects of Wei d n e r , H., 1970. Zoraptera (Bodenläuse). Handbuch Zool., Australia: A Textbook for Students and Research Wor ke r s , 4(2/15), 1-12. 2nd edition, 141-179, Ithaca, Cornell Univ. Press. Wei d n e r , H., 1976. Eine neue Zo ro t y p u s -A r t von den Gala- Kuk a l ov á - P eck, J., Peck, S.B., 1993. Zoraptera wing struc t u r e s : pagosinseln, Zo r otypus leleupi sp. n. (Zoraptera). Mission Evidence for new genera and relationship with the blattoid or- zo o l o gique belge aux Iles Galapagos et en Ecuador, 3, 161- ders (Insecta: Blattoneoptera). Syst. Entomol., 18, 333-350. 176. Ma rt y n o v, A. V ., 1925. To the knowledge of fossil insects from Wh e e l e r , W.C., Whiting, M., Wh e e l e r , Q.D ., Carpe n t e r , J.M., In Jurassic beds in Tur k estan. 2. Raphidioptera (continued), press. The phyl og e n y of the extant hexapod orders. Cladistics.

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