View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Revistes Catalanes amb Accés Obert ACTA GEOLOGICA HISPANICA, v. 35 (2000), nº 1,p. 149-164 AWinged Zorotypusin Miocene Amber from the Dominican Republic (Zoraptera: 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 Neoptera 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 insect 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- C AUDELL (1918) (Crampton, 1920; Gurn ey, 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- insects 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 termite 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 Paraneoptera (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); + Dictyoptera 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.