Systematic Entomology (2020), 45, 349–364 DOI: 10.1111/.12400 The evolution of Zoraptera YOKOMATSUMURA 1,2,ROLFG.BEUTEL 1*,JOSÉA.RAFAEL 3*, IZUMIYAO 4*,JOSENIRT.CÂMARA 5*,SHEILAP.LIMA 3 KAZUNORIYOSHIZAWA 4 1E G,I ü Z Evh,Fh Sh Uv J,J,G, 2D F Mh Bh,Z I,K Uv,K,G, 3I N P Az,M,Bz, 4S E,Sh A,Hkk Uv,S,J 5Uv F Pí, B J,Pí, Bz Abstract. Z h .B w x wh h kw, w h h DNA -v h. Th h hw h Z v h , h w . Th h h h v h . Th v h P. P h h h Z w h v, h h . Th h h P , v h k Gw L . Th x v f v h , hv, k hh v h.W v v h h v.O v / h , hk . Introduction B et al., 2014; Mh et al., 2014; Ch, 2018). I h h v (vw Z h h I Mh- Mh et al., 2014;K et al., 2016; B et al., 2017), G. Th wh h wh Z (G &E, 2005; h P - v (Yhzw, 2011; M et al., 2014; C:Yk M,E G,I ü Z Evh,Fh Sh Uv W &P, 2014; Mh et al., 2014, 2015; M J,E. 1, 07743 J,G &D F- et al., 2015; W et al., 2019). R,W et al. Mh Bh,Z I,K (2019) h h h - Uv, Bh G 1–9, 24118, K,G. P v v, E-: k..h@.;R G. B, h hh wh M E G,I ü Z Evh, et al. (2014). Th v Z h Fh Sh Uv J,E. 1, 07743 J,G. D, h w h E-: .@-.;Jé A. R,I N P Az,Cx P 2223, 69067-375 M, AM, h . Th h h - Bz. E-: @.v.;Kz Yhzw,S P w ‘ -w E,Sh A,Hkk Uv, 060-8589 wh f v h S,J. E-: @..hk.. w ’ (W et al., 2019). C h w h Z (..Fh & B, 2008; ∗Th h h wk. Mh et al., 2014;M et al., 2015), © 2019 Th Ah. Systematic Entomology h Jh W &S L h R E S. 349 Th h h Cv C A L, whh , , v h wk . 350 Y. Matsumura . h x v h - h h x v h h ‘’ . Th v h h h h v .Hwv, wh w h x h f v h h k h h x (Mh et al., 2014). h h v ( E, 2003). I h v - Th h h- (.. Ph,M,B, wh Z . Th x Oh), 44 x 16 x v . Th v- Z (Mh et al., 2018, 2019; h v v - Ch &S, 2019). Kkvá-Pk & Pk (1993) x h h ( S wh Z h w v.Hwv, et al., 2012). Th h v wh h h x kw v v wh v w h (Ch, 1989). C, h w z h . wh Zorotypus E (2000) wh ‘h h- h ’. Lkw, Ch & Ch (2000) w z wh Materials and methods Zorotypus (E, 2000). S h,E’ f h w (R &E, M h w h fx 2006; T & Wh, 2012; Mh et al., 2013; Y wh 80–99.5% h (F. 1). T Zoroty- et al., 2015; W et al., 2016; Ch, 2018). Th pus novobritannicus w w h Ah C- Zorotypus v , < 3 , ,Bh Y Uv,Pv, UT, U.S.A. I , v , . Th k 31 v 21–22 w wh h x v (F. 2). W h YK16-10 h ( h) x v E Z. 6. W (Mh et al., 2014; Ch, 2018). C Z. huxleyi Z. hamiltoni h h P h , Z. 6 h (W et al., 2019), Z hv .O w hz h, hv P,H,Eh, z.Hwv, h v O,Z Ahh. Th w h v wh Ahh. Th h x w . GBk ( h Th k v - T S1, S2). h x h . Th hh vf hv v v, wh (Hü, 2007; M- DNA extractions, amplifcations, sequences and alignment et al., 2014), h wk x (..G, 1938). I , -h P h w w : w (..G, 18S RNA H 3, h 16S RNA 12S 1938; Nw, 1978, 2000; Mh et al., 2013). T RNA. Th k w 18S kw, h h P. (18S-Z-: 3′-ATT AAG CCA TGC AAG TGT CAG-5′; D h 18S-Z-: 3′-TTA RYA TAT GCT ATT GGW GCT GG-5′), kw wh Z, , , Z. zimmermani H 3 (H3-E-: 3′-AAR GCY CCW MGM AAR (G, 1939) h Z. barberi (G, 1938). CAR CT-5′;H3-E-: 3′-TGR ATR GCR CAV AGR TTR S (G, 1938; GTR TCY TC-5′), 16S (16S: 3′-CCG GTC TGA ACT CAG P, 1949, 1951; Hw, 1974, 1976; R &E, ATC ACG T-5′; 16S-L: 3′-CGC CTG TTT ATC AAA 2006; Hü, 2007; R et al., 2008; Mh et al., AAC AT-5′) 12S (12S-Z-: 3′-TGG CGG YRW DWW 2013, 2018; W et al., 2016; Kˇák et al., 2017; Y &L, RWT YTW TYR GRG G-5′; 12S-Z-: 3′-TTA CTM TYA 2017), k h f AAT CCA CCT TC-5′). Mh DNA x, PCR .Ah vf h f w Yhzw &Jh h h , h (2008). S f w z hk (.. h G-T E V G, 1938). Th v h - (P,M, WI, U.S.A.) w - x.M .A - w hw h , -w w f h .R RNA h h whh h w mat 6.5 (Kh & S, 2013) wh kw (Sh, 1978; Ch, 1994, 1995; Mh et al., h Q-INS- , whh 2011; D et al., 2013). Th w -h RNA .A w ©2019 Th Ah. Systematic Entomology h Jh W &S L h R E S. 45, 349–364 The evolution o Zoraptera 351 Fig. 1. Z . (A) Z. C R; (B) Z. asymmetristernum K; (C–F) Z. caudelli M. Phh T. K (A) T. K (B–F). [C f vw w.]. , w x h w h x x . (F S1). Pv hw h h h h Model selection and phylogenetic estimation (..K, 2004; Yhzw & Jh, 2005; Ihw et al., 2011; M et al., 2014; W Th h w et al., 2019). Th, w h h - partitioninder 2.3.3 (L et al., 2017), h M et al., (2014) wh h h. Th H 3 W et al., (2019) h w h . (h ) RNA (h ) w f I , w h partitioninder . Th -f h h h Z ( F S1). ©2019 Th Ah. Systematic Entomology h Jh W &S L h R E S. 45, 349–364 352 Y. Matsumura . Fig. 2. C h h.D Bz hw h . W x kh iq-tree 1.6.3 v E (T 1). F , (N et al., 2015), wh 10 000 k- h Rh h (412 M) w h h h wh - (H et al., 2018) x Ev et al. (2019). I h v.T h , , h h x 450 M w iq-tree w .A h - Z-P v M et al. , w h (2014) T et al. (2015). h F. 3. A B w F h mcmctree , w f h - mrbayes (R &Hk, 2003). W h v 419 M h w h wh h 1 000 000 - P–P h T et al. , w v 1000 . Th (2015). B h , h f 10% w x -, 50% w baseml h paml w k. - w Th GTR + G w wh = 0.5, whh w . tracer h beast w k (Bk- x h et al., 2014) w hk h h Mkv h M jmodeltest (P, 2008) h .W C (MCMC) h v. 1 000 000 , h v w v 50 . Th f 10% h v w Divergence time estimation x -.W w hk h h MCMC h v ( R F v , B h w et al., 2017). h w mcmctree h paml 4.8 w F h beast , w h clade ages k k (Y, 2007) beast 2.6 (Bk et al., 2014). (Mh et al., 2017). Th w w : Th w w BEAST M T h , x k M et al. (2014) T et al. (2015): 160 h k , h-h h .W M h v P; 130 M h 20 , h f 10% v I; 99 M h h v w x -. tracer h ©2019 Th Ah. Systematic Entomology h Jh W &S L h R E S. 45, 349–364 The evolution o Zoraptera 353 Fig. 3. C x kh iq-tree. N- w .Ak h h w .M Z hhh ( x ). [C f vw w.]. Table 1. A , -x- (DEC) ( L;R &Sh, 2008) h - F x C A (M) w rasp 3.2 (Y et al., 2015). D-v Pronemoura angustithorax P 161.0–160.5 (DIVA; R, 1997) w v h Sorellembia estherae E 99.9 DEC .Hwv, v hw h DIVA Valditermes brennenae I 136.4–130.0 w f wh x wh- v (Kh, 2010). Fv h w f:A beast w k (Bk et al., 2014) w (AF), I (IM), N (NA), N (NT) hk h h MCMC h v. A. Th x w h w w, Biogeographical analysis w w wh. F x , h w A w h h . Th h h mcmctree .O h w h kw w x h .W h . ©2019 Th Ah. Systematic Entomology h Jh W &S L h R E S. 45, 349–364 354 Y. Matsumura . Character evolution wh Z. asymmetristernum (AF). Th - h Z. .1 (AF) w h W h w h :() A + N (77.15%) A (22.85%) ;() (F.