Gonatopodinae Kieffer, 1906

Xu, Zaifu, Olmi, Massimo & He, Junhua, 2013, Dryinidae of the Oriental region (Hymenoptera: Chrysidoidea), Zootaxa 3614 (1), pp. 1-460 : 315-316

VI. Subfamily Gonatopodinae Kieffer in Kieffer & Marshall, 1906

Gonatopodinae partim Kieffer in Kieffer & Marshall 1906: 499.

Gonatopodinae partim Kieffer 1907: 12.

Gonatopodini Kieffer 1914b: 67; Richards 1939: 188.

Dryinini partim Muesebeck & Walkley 1951: 1034.

Gonatopodinae Richards 1953: 51 ; Krombein 1979: 1244; Olmi 1984: 1050; Olmi 1993a: 195; Olmi 1993b: 77; Olmi 1994: 64; Olmi 1995b: 503; Olmi 1995d: 134; Olmi 1999a: 213; Olmi & Bechly 2001: 42; He & Xu 2002: 287; Engel 2005: 490; Olmi & Virla 2006: 408; Virla & Olmi 2008: 371

Type genus. Gonatopus Ljungh, 1810 , designated by Olmi (1984).

Diagnosis. Female: usually apterous ( Plates 106B , 108C ); occasionally fully winged ( Plate 96A , 100A , 106A ); rarely micropterous (not present in the Oriental region) or brachypterous (present in Oriental region); ocelli present; occipital carina usually absent, rarely incomplete and only present behind and on sides of posterior ocelli; mandible with four teeth progressing larger from anterior one to posterior; palpal formula 2/1, 2/2, 3/2, 4/2, 4/3, 5/2, 5/3, 6/2, or 6/3; antennal segments 5–10 without tufts of long hairs; antenna with or without rhinaria; pronotal tubercle absent ( Plates 96B , 100B ); in winged females, metanotum very reduced; in macropterous species fore wing with stigmal vein and pterostigma ( Plates 96A, B , 100A , 106A ), pterostigma very narrow; in macropterous species fore wing with three cells enclosed by pigmented veins (costal, median and submedian) ( Plates 96A, B , 100A , 106A ); protarsus chelate ( Plates 96A, B , 106B , 108C ); chela with rudimentary claw ( Plates 93B , 107B ); protrochanter long and slender, more than twice as long as broad ( Plate 96B ); usually tibial spurs 1/0/1; rarely (in Neodryinus R. Perkins ) 1/0/2. Male: fully winged ( Plate 94 , 110A, B ); occipital carina usually absent, rarely incomplete and only present behind and on sides of posterior ocelli, rarely complete (in Echthrodelphax R. Perkins ); occiput usually concave ( Plate 121C ), occasionally straight; mandible with three teeth progressing larger from anterior one to posterior; palpal formula 2/1, 2/2, 3/2, 4/2, 4/3, 5/2, 5/3, 6/2, or 6/3; epicnemium visible, because lateral regions of prothorax not continuous with mesopleura (as in plate 2H); mesosternum fused with mesopleura and not distinct; fore wing with three cells enclosed by pigmented veins (costal, median and submedian) ( Plates 94A , 110A, B ); fore wing with stigmal vein and pterostigma (pterostigma always very narrow) ( Plates 94A , 110A, B ); dorsal process of paramere always present ( Plate 95B ), occasionally reduced ( Plates 117A , 122B ); tibial spurs 1/1/2.

Distribution. Worldwide.

Hosts. Cicadellidae (except Idiocerinae , Macropsinae , Typhlocybinae ), Tropiduchidae , Lophopidae , Ricaniidae , Acanaloniidae , Flatidae , Delphacidae , Issidae , Nogodinidae , Meenoplidae ( Guglielmino & Olmi, 1997, 2006, 2007).

World species. Five hundred and fifty three species are known, eighty one in the Oriental region, including one species incertae sedis.

World genera. Twelve genera are known, six in the Oriental region.

Remarks. The following six genera are not quoted from the Oriental region: Epigonatopus R. Perkins, 1905 (Australian) , Eucamptonyx R. Perkins, 1907 (Australian, Nearctic, Neotropical), Gynochelys Brues, 1906 (Afrotropical) , Pareucamptonyx Olmi, 1991 (Neotropical) , Pentagonatopus Olmi, 1984 (Australian, Nearctic), Trichogonatopus Kieffer, 1909 (Nearctic, Neotropical).

Key to the genera

Females

1. Females fully winged; rarely brachypterous ( Plates 96A, B , 100A , 106A )......................................... 2

- Females apterous ( Plates 106B , 108C ).................................................................... 5

2. Tibial spur formula 1/0/2............................................................ 1. Neodryinus R. Perkins

- Tibial spur formula 1/0/1............................................................................... 3

3. Notauli absent or hardly visible (then incomplete or complete).................................... 2. Adryinus Olmi

- Notauli clearly distinct and complete...................................................................... 4

4. Species small, with palpal formula variable from 3/2 to 4/2, or 5/2, rarely 5/3 or 6/3; segment 5 of protarsus with less than 15 medial lamellae ( Plates 100C , 102B, C )............................................ 3. Echthrodelphax R. Perkins

- Species large, with palpal formula 6/3; segment 5 of protarsus with more than 20 medial lamellae.................................................................................................... 5. Gonatopus Ljungh partim

5. Palpal formula 2/1.............................................................4. Haplogonatopus R. Perkins

- Palpal formula different................................................................................ 6

6. Palpal formula 6/2................................................................... 6. Esagonatopus Olmi

- Palpal formula different.......................................................... 5. Gonatopus Ljungh partim

Males (unknown in Adryinus Olmi )

1. Head with a distinct complete occipital carina ( Plate 101A )............................. 3. Echthrodelphax R. Perkins

- Head without occipital carina, or with an incomplete occipital carina (then carina only present behind ocellar triangle and shortly on sides of posterior ocelli)....................................................................... 2

2. Temple absent..................................................................... 1. Neodryinus R. Perkins

- Temple distinct ( Plate 110A )............................................................................ 3

3. Palpal formula 2/1.............................................................4. Haplogonatopus R. Perkins

- Palpal formula different................................................................................ 4

4. Palpal formula 6/2................................................................... 6. Esagonatopus Olmi

- Palpal formula different.......................................................... 5. Gonatopus Ljungh partim

Figures

PLATE 93. Chelae. A. Neodryinus dolosus, holotype (from Olmi 1984). B. N. grandis, holotype (Xu et al. 2011a). C. N. isoneurus, holotype (from Xu & He, 1997e). D. N. javanus, female from Malaysia (from Olmi 1984). Scale bar 0.37 mm for A and C, 0.31 mm for B and D.

PLATE 94. Neodryinus javanus, Male. Length 3.4 mm.

PLATE 95. A, B. Male genitalia. C. Pronotum in lateral view, holotype. D, E. Chelae, holotypes. A. Neodryinus javanus from Malaya (from Olmi 1984). B. N. karaentensis, holotype (from Olmi 2011a). C, D. N. leptopus (from Olmi 1984). E. N. lineatus (from Xu et al. 2003). Scale bar 0.26 mm for A, 0.21 mm for B, 1.19 mm for C, 0.29 mm for D and E.

PLATE 96. Neodryinus malayanus, female holotype (from Olmi 1991). Length 5.0 mm. A. Dorsal view. B. Lateral view.

PLATE 100. Echthrodelphax fairchildii. A, B. Female in dorsal view (A) and lateral view (B). Length 2.5 mm. C. Chela of female from Japan. Scale bar 0.16 mm for C. (From Olmi 1984)

PLATE 101. Echthrodelphax fairchildii, males from Hawaii, Oahu, Ewa (from Olmi 1984). Length 1.8 mm. A. Dorsal view. B. Lateral view.

PLATE 102. A, B, E, F. Male genitalia. C, D. Chelae. A. Echthrodelphax fairchildii from Hawaii (from Olmi 1984). B, C. E. laotianus from Thailand (from Olmi 1998e). C. E. laotianus, holotype (from Olmi 1984). D. E. rufus, paratype (arolium removed) (from Olmi 1984). E, F. E. rufus from China, paramere in ventral side (E) and dorsal view (F) (from Olmi 1995a). Scale bar 0.13 mm for A, 0.08 mm for B, 0.14 mm for C, 0.11 mm for D, E and F.

PLATE 106. A. Gonatopus longicollis, female from Spain. Length 4.8 mm (from Olmi 1999a). B. G. asiaticus, holotype. Length 3.0 mm (from Olmi 1984).

PLATE 107. Chelae, holotypes. A. Gonatopus paulyi (from Olmi 1986). B. G.abjectus (from Olmi 1984). C. G. bengalensis (from Olmi 1984). D. G. lucens (from Olmi 1984). Scale bar 0.15 mm for A, 0.14 mm for B, 0.21 mm for C and 0.20 mm for D.

PLATE 108. A. Gonatopus pedestris, female from Italy, lateral view. Length 3.0 mm. B. Mesosoma, lateral view of G. rufoniger (=G. indianus Olmi), from India. Length 2.0 mm. C. Female of G. bicolor, lateral view. Lenght 2.9 mm. D. G. acuminatus, holotype, lateral view. Length 3.12 mm. (From Olmi 1984)

PLATE 110. Gonatopus nearcticus (=Acrodontochelys ugandanus (Benoit)), male from Uganda. Length 3.0 mm (from Olmi 1984). A. Dorsal view. B. Lateral view.

PLATE 117. A–C. Male genitalia. D, H. Chelae, holotypse. E. Head. F, G. Mesosoma. A. Gonatopus nudus from Malaya. B. G. sarawakensis or malesiae, var. a. C. G. sarawakensis or malesiae, var. b. D. G. sarawakensis. E, F, H. G. validus. G. G. validus, female from China. Scale bar 0.24 mm for A, 0.27 mm for B, 0.29 mm for C, 0.16 mm for D, 0.12 mm for H. (From Olmi 1984).

PLATE 121. Gonatopus flavifemur. A. Female from Viet Nam. Length 3.0 mm. B. Chela of female from Japan (from Olmi 1984). C. Male (=Dicondylus indianus Olmi) from India (from Olmi 1984). Length 2.6 mm. Scale bar 0.20 mm for B.

PLATE 122. A. Male, lateral view. B–D. Male genitalia. E. Chela, holotype. A. Gonatopus flavifemur (=Dicondylus indianus Olmi) from India. Length 2.6 mm. B. G. flavifemur from Australia. C. G. lucens from Japan. D. G. pedestris from Finland. E. G. lucens. Scale bar 0.17 mm for B, 0.10 mm for C, 0.15 mm for D and 0.19 mm for E. (From Olmi 1984).