Gonatopus Ljungh, 1810

Xu, Zaifu, Olmi, Massimo & He, Junhua, 2013, Dryinidae of the Oriental region (Hymenoptera: Chrysidoidea), Zootaxa 3614 (1), pp. 1-460 : 354-360

18. Genus Gonatopus Ljungh, 1810

Gonatopus Ljungh 1810: 161 ; Richards 1937: 118; Olmi 1993b: 77; Olmi 1994: 66; Olmi 1995c: 787; Olmi 1995g: 5; Olmi 1996b: 182; Olmi 1998c: 72; Olmi 1998d: 177.; Olmi 1999a: 231; He & Xu 2002: 318.; Virla & Olmi 2008: 371.; MoyaRaygoza & Olmi 2010: 100.

Dicondylus Haliday in Curtis 1829 –1930: 110 (synonymized by Olmi, 1993b); type species: Dicondylus bicolor (Haliday in Curtis 1828), designated by Kieffer (1914b); Haliday 1833: 273; Walker 1837: 411; Kieffer 1914b: 72; Richards 1935: 147; Richards 1937: 118; Olmi 1984: 1374; Olmi 1993b: 78.

Labeo Haliday 1833: 273 (nom. praeocc., nec Cuvier 1817) (synonymized by Richards 1937); type species: Labeo vitripennis Haliday 1833 , by monotypy; Kieffer in Kieffer & Marshall 1905: 220; Walker 1837: 429; Dalla Torre 1898: 542.

nec Labea Blanchard 1840: 430 .

Labeola Haldeman 1842: 191 , new name for Labeo Haliday, 1833 (synonymized by Krombein 1979 with Dicondylus Haliday, 1829 –1930).

Pseudogonatopus R. Perkins 1905: 34 (synonymized by Olmi 1993b); type species: Pseudogonatopus kurandae R. Perkins, 1905 , by original designation; Kieffer in Kieffer & Marshall 1906: 488; Kieffer 1914b: 79; Richards 1939: 195; J. Perkins 1976: 32; Chandra 1980c: 192; Olmi 1984: 1191; Olmi 1993b: 78.

Paragonatopus R. Perkins 1905: 40 (synonymized by Olmi 1984); type species: Paragonatopus nigricans R. Perkins 1905 , by monotypy; Kieffer in Kieffer & Marshall 1906: 488; Kieffer 1914b: 87; Richards 1939: 195; Olmi 1979: 218.

Neogonatopus R. Perkins 1905: 42 (synonymized by Olmi 1984); type species: Neogonatopus ombrodes R. Perkins 1905 , by original designation.

Pachygonatopus R. Perkins 1905: 45 (synonymized by Olmi 1984); type species: Pachygonatopus melanias R. Perkins 1905 , by original designation.

Chalcogonatopus R. Perkins 1905: 46 (synonymized by Olmi 1984); type species: Chalcogonatopus gigas R. Perkins 1905 , by original designation.

Eugonatopus R. Perkins 1905: 46 (synonymized by Olmi 1984); type species: Chalcogonatopus (Eugonatopus) pseudochromus R. Perkins 1905 , by original designation.

Mercetia Kieffer in Kieffer & Marshall 1906: 495 (synonymized by Olmi 1996b); type species: Mercetia longicollis ( Kieffer 1905) , by original designation; Olmi 1996b: 182; Kieffer 1914b: 19.

Platygonatopus Kieffer in Kieffer & Marshall 1906: 500 (synonymized by Olmi, 1984); type species: Platygonatopus planiceps ( Kieffer 1904) , designated by Richards (1939).

Apterodryinus R. Perkins 1907: 14 (synonymized by Olmi 1993b); type species: Apterodryinus torvus R. Perkins 1907 , by monotypy; Olmi 1984: 1342; Olmi 1993b: 78.

Agonatopus R. Perkins 1907: 29 (synonymized by Olmi 1984); type species: Agonatopus pallidicornis R. Perkins 1907 , designated by Richards (1939).

Agonatopoides R. Perkins 1907: 33 (synonymized by Olmi 1993b); type species: Agonatopoides synchromus R. Perkins 1907 , by monotypy; Olmi 1984: 1310; Olmi 1993b: 78.

Cyrtogonatopus Kieffer 1907: 19 (synonymized by Olmi 1984); type species: Cyrtogonatopus breviforceps ( Kieffer 1904) , monotypic.

nec Trichogonatopus Kieffer 1909: 336 .

Pristogonatopus Kieffer 1913c: 325 (synonymized by Olmi & Currado 1974); type species: Pristogonatopus dentatiforceps ( Kieffer 1904) , by original designation; Kieffer 1914b: 71; Richards 1939: 195.

Digonatopus Kieffer 1913c: 325 (synonymized by Richards 1939); type species: Digonatopus javanus (R. Perkins 1912) , by monotypy.

Trigonatopus Kieffer 1913c: 325 (synonymized by Olmi 1984); type species: Trigonatopus bifarius (Kieffer in Kieffer & Marshall 1906), by original designation; Kieffer 1914b: 72.

Laberius Kieffer 1914b: 59 , new name for Labeo Haliday 1833 (synonymized by Richards 1937); Richards 1936: 169.

Trichogonatopus Hellén 1930: 2 (synonymized by Olmi 1984); type species: Trichogonatopus lindbergi Hellén 1930 , original designation.

Laberinus Ogloblin 1932: 266 (mispelling).

Metagonatopus Ogloblin 1932: 264 (synonymized by Olmi 1984); type species: Metagonatopus brasiliensis Ogloblin 1932 , by original designation.

Allogonatopus Haupt 1938: 27 (synonymized by Olmi 1984); type species: Allogonatopus procerus Haupt 1938 , by original designation.

Donisthorpina Richards 1939: 201 (synonymized by Olmi 1993b); type species: Donisthorpina formicicola Richards 1939 , by original designation; Olmi 1984: 1304; Olmi 1993b: 78.

Plectrogonatopus Richards 1939: 207 (synonymized by Olmi 1984); type species: Plectrogonatopus striatus ( Kieffer 1905) , by original designation.

Tetrodontochelys Richards 1939: 217 (synonymized by Olmi 1993b); type species: Tetrodontochelys ljunghii ( Westwood 1833) , by original designation; Olmi 1984: 1434; Olmi 1993b: 78.

Epigonatopoides Richards 1939: 221 (synonymized by Olmi 1984); type species: Epigonatopoides pulicarius ( Klug 1810) , by original designation.

Idologonatopus Ogloblin 1953: 111 (synonymized by Olmi 1984); type species: Idologonatopus nigrithorax Ogloblin 1953 , by original designation.

Rhynchogonatopus Benoit 1954: 390 (synonymized by Olmi 1984); type species: Rhynchogonatopus meridionalis Benoit 1954 , by original designation.

Madecagonatopus Benoit 1954: 392 (synonymized by Olmi 1984); type species: Madecagonatopus ankaratrus ( Ceballos 1936) , by original designation.

Pseudogonatopoides J. Williams 1956: 90 (synonymized by Olmi 1984); type species: Pseudogonatopoides mauritianus J. Williams 1956 , by original designation.

Cyrtogonatopoides Ponomarenko 1966: 1579 (synonymized by Olmi 1984); type species: Cyrtogonatopoides borealis Ponomarenko 1966 , by original designation.

Plectrogonatopoides Ponomarenko 1975a: 318 (synonymized by Olmi 1993b); type species: Plectrogonatopoides chortinis Ponomarenko 1975 , by original designation; Olmi 1984: 1431; Olmi 1993b: 78.

Tetradontochelys J. Perkins 1976: 35 (injustified emendation).

Neogonatopoides Abdul-Nour 1976: 274 (synonymized by Olmi 1984); type species: Neogonatopoides claudiae Abdul-Nour 1976 , by monotypy.

Dolichochelys Olmi & Currado 1976: 177 (synonymized by Olmi 1984); type species: Dolichochelys longicornis ( Kieffer 1904) , by original designation.

Megagonatopus Olmi & Currado 1976: 178 (synonymized by Olmi 1984); type species: Megagonatopus birmanus Olmi & Currado 1976 , by original designation.

Acrodontochelys Currado 1976: 13 (synonymized by Olmi 1993b); type species: Acrodontochelys bouceki Currado 1976 , by original designation; Olmi 1984: 1172; Olmi 1993b: 78.

Nogatopus Ponomarenko 1979: 357 (synonymized by Olmi 1984); type species: Nogatopus gobiensis Ponomarenko 1979 , by original designation.

Paradicondylus Olmi 1986: 102 (synonymized by Olmi 1998c); type species: Paradicondylus paulyi Olmi 1986 , by monotypy and original designation; Olmi 1993b: 78; Olmi 1998c: 72.

Type species. Gonatopus formicarius Ljungh, 1810 , by monotypy.

Diagnosis. Female: apterous ( Plates 106B , 126A ) or fully winged ( Plate 106A ); palpal formula 3/2, 4/2, 4/3, 5/ 2, 5/3, or 6/3; antenna with or without rhinaria; pronotum crossed or not by transverse furrow ( Plate 108C, D ); enlarged claw with distal apex pointed and with one large or small subapical tooth ( Plate 107B, C ) (occasionally subapical tooth absent, then enlarged claw with distal group of lamellae ( Plate 107A )); in fully winged forms, segment 5 of protarsus with more than 20 lamellae; tibial spurs 1/0/1. Male: fully winged ( Plate 110A, B ); occipital carina absent or incomplete (in this last case, only present behind and shortly on sides of posterior ocelli); occiput concave; temple present; palpal formula 3/2, 4/2, 4/3, 5/2, 5/3, or 6/3; tibial spurs 1/1/2.

Distribution. Worldwide.

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

World species. Four hundred and thirty nine species are known, fifty eight species in the Oriental region and one species incertae sedis.

Remarks. Larvae of Gonatopus were described by Ponomarenko (1975b) , Carcupino et al. (1996, 1998), Guglielmino & Virla (1998), Virla & Mangione (2000), Guglielmino (2002), Mangione & Virla (2004) and Guglielmino et al. (2006, 2007, 2008). Gonatopus females were divided for convenience by Olmi (1993b) into eleven groups on the basis of the following characters:

Key to the groups of Gonatopus

1. Fully winged ( Plate 106A )....................................................................... Group 11

- Apterous ( Plate 106B )................................................................................. 2

2. Enlarged claw without subapical tooth ( Plate 107A ), or with one very small subapical tooth ( Plate 107B )............... 3

- Enlarged claw with one large subapical tooth ( Plate 107C )..................................................... 5

3. Distal apex of enlarged claw with group of lamellae ( Plate 107A )......................................... Group 8

- Distal apex of enlarged claw without lamellae ( Plate 107C, D )................................................. 4

4. Pronotum not crossed by transverse impression or slightly impressed ( Plate 108A )........................... Group 6

- Pronotum crossed by strong transverse impression ( Plate 108B ).......................................... Group 7

5. Pronotum not crossed by transverse impression or slightly impressed ( Plate 108C )................................. 6

- Pronotum crossed by strong transverse impression ( Plate 108D )................................................ 7

6. Labial palpi 3-segmented......................................................................... Group 9

- Labial palpi 2-segmented......................................................................... Group 5

7. Enlarged claw without lamellae, with or without bristles or peg-like hairs ( Plate 109B )........................ Group 1

- Enlarged claw with lamellae ( Plate 107C ).................................................................. 8

8. Labial palpi 2-segmented............................................................................... 9

- Labial palpi 3-segmented.............................................................................. 10

9. Maxillary palpi composed of two–four segments...................................................... Group 2

- Maxillary palpi 5-segmented..................................................................... Group 10

10. Maxillary palpi 6-segmented...................................................................... Group 4

- Maxillary palpi 5-segmented...................................................................... Group 3

The above groups are distributed in the zoogeographical regions as follows:

Palaearctic: groups 1, 2, 3, 4, 5, 6, 7, 10, 11.

Afrotropical: groups 1, 2, 3, 4, 5, 6, 7, 8, 11.

Oriental: groups 1, 2, 3, 4, 5, 6, 7, 10.

Nearctic: groups 1, 2, 3, 4, 5, 6, 7, 9.

Neotropical: groups 1, 2, 3, 4, 5, 6, 7, 10.

Australian: groups 1, 2, 4, 5, 6, 7, 9.

As far as, it is not possible divide also males into the above groups.

All Oriental species of Gonatopus can be attributed to one of the above eight female groups, except Gonatopus leurosipropodeus He & Xu, 2002 . The attribution of this species to a distinct group is impossible, because it was described on the basis of male specimens and its female is unknown. For this reason, it is attributed to an uncertain group (see “Uncertain group”).

Key to the males

(unknown for the species not present in the key) (males not divided into groups)

1. Notauli incomplete.................................................................................... 2

- Notauli complete..................................................................................... 4

2. Propodeum with dorsal surface dull, granulated, sculptured by irregular striae, and posterior surface shiny, smooth and unsculptured; dorsal surface of propodeum much shorter than posterior surface...................... G. maurus Kieffer

- Propodeum completely smooth and shiny, unsculptured; dorsal surface of propodeum much longer than posterior surface... 3

3. Dorsal process of paramere transverse ( Plate 122C ); clypeus testaceous.............................. G.lucens (Olmi)

- Dorsal process of paramere about parallel to penis, not transverse ( Plate 111A ); clypeus black....... G. nearcticus (Fenton)

4. Notauli posteriorly broadly separated..................................................................... 5

- Notauli posteriorly joint (or very close and almost joint)..................................................... 14

5. Dorsal process of paramere reduced to inner expansion of paramere ( Plate 122B )....... G. flavifemur (Esaki & Hashimoto)

- Dorsal process of paramere not reduced ( Plates 111B , 118D , 119D , 128A , 131A, B )................................ 6

6. Dorsal process of paramere much shorter than paramere ( Plates 128A , 131A, B )................................... 7

- Dorsal process of paramere as long as, or longer, or slightly shorter than paramere ( Plates 111B , 118D , 119D )............ 9

7. Head with POL more than four times as long as OOL; ocelli very large............................ G. rufoniger Olmi

- Head with POL less than three times as long as OOL; ocelli small............................................... 8

8. Dorsal process of paramere very slender ( Plate 131B ); clypeus completely black...................... G. superbus Olmi

- Dorsal process of paramere less slender ( Plate 128A ); clypeus medially black, laterally testaceous... G.javanus (R. Perkins)

9. Face completely or almost completely testaceous or yellow................................................... 10

- Face completely black or brown........................................................................ 11

10. Dorsal process of paramere longer than penis and paramere ( Plate 118D )......................... G. borneanus (Olmi)

- Dorsal process of paramere shorter than penis and paramere ( Plate 111B )...................... G. nigricans (R. Perkins)

11. Antennal segment 3 less than three times as long as broad; dorsal process of parameres longer than penis ( Plate 119D )............................................................................................ G. pyrillae (Mani)

- Antennal segment 3 more than three times as long as broad; dorsal process of parameres shorter than penis ( Plates 111B , 117C , 122D )............................................................................................. 12

12. Dorsal process of paramere almost completely joint to paramere ( Plate 111B ).................. G. nigricans (R. Perkins)

- Dorsal process of paramere completely separated from paramere ( Plates 117C , 122D ).............................. 13

13. Dorsal process of paramere slender ( Plate 122D )............................................. G. pedestris Dalman

- Dorsal process of paramere broader ( Plate 117C )............ G.malesiae (Olmi) type B or G.sarawakensis (Olmi) type B

14. Dorsal process of paramere very reduced ( Plates 117A , 122B ), reduced to inner expansion of paramere................ 15

- Dorsal process of paramere not reduced ( Plates 111B , 117B ).................................................. 16

15. Inner expansion of paramere with many irregular striae ( Plate 122B )................. G. flavifemur (Esaki & Hashimoto)

- Inner expansion of paramere without irregular striae ( Plate 117A )............................... G. nudus (R. Perkins)

16. Dorsal process of paramere much shorter than paramere ( Plates 111B , 131C )..................................... 17

- Dorsal process of paramere approximately as long as paramere ( Plates 117B , 131D )............................... 18

17. Dorsal process of paramere not transverse, with distal apex broad ( Plate 111B ); head and mesosoma mostly black or brown................................................................................... G. nigricans (R. Perkins)

- Dorsal process of paramere transverse, with distal apex pointed ( Plate 131C ); head and mesosoma mostly yellow-testaceous....................................................................................... G. asiaticus (Olmi)

18. Propodeum completely reticulate rugose; dorsal process of paramere broader ( Plate 117B )................................................................................ G.malesiae (Olmi) type A or G. sarawakensis (Olmi) type A

- Propodeum with posterior inclined region reticulate rugose and anterior flat region smooth and unsculptured; dorsal process of paramere slender ( Plate 131D ).................................................... G. leurosipropodeus He & Xu

Remarks. In the above key, there are two species, whose males are not distinguishable, G. malesiae (Olmi) and G. sarawakensis (Olmi) ; the situation is explained in remarks related to G. sarawakensis (see Group 2).

Group 1

One species is known in the Oriental region.

Figures

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 109. A. Gonatopus nearcticus (=Acrodontochelys bouceki Currado), female from Spain, Nerja (from Currado, 1976). Length 3.5 mm. B. Chela of G. nearcticus (=Acrodontochelys sinensis Olmi), female from China, Macau (from Olmi 1984). Scale bar 0.17 mm for B.

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

PLATE 111. A, B. Male genitalia. C. Scutum + metathorax + propodeum. D. Chela. A. Gonatopus nearcticus, from Greece. B. G. nigricans, from Malaya. Different shapes of dorsal process of paramere. C, D. G. bicuspis, holotype (from Olmi 1993d). Scale bar 0.14 mm for A and B, 0.40 mm for C and 0.12 mm for D.

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 118. A. Scutum and metathorax + propodeum, lateral view (from Olmi 1989b). B. Furrow of metathorax + propodeum, frontal view (from Olmi 1989b). C, E. Chelae (from Olmi 1984). D. Male genitalia (from Olmi 1984). A–C. Gonatopus borneanus, holotype. D. G. borneanus, paratype. E. G. insularis, holotype. Scale bar 0.75 mm for A, 0.15 mm for C, 0.18 mm for D and 0.14 mm for E.

PLATE 119. A. Scutum and metathorax + propodeum, lateral view. B. Furrow of metathorax + propodeum, frontal view. C, E. Chelae, holotypes. D. Male genitalia. A–C. Gonatopus pajanensis (from Olmi 1989b). D, E. G. pyrillae (from Olmi 1984). Scale bar 0.29 mm for C, 0.19 mm for D and 0.16 mm for E.

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).

PLATE 126. A. Dorsal view. Length 7.5 mm. B, D, E. Heads. C. Mesosoma. F. Chela. A–C, F. Gonatopus cristatus (=Megagonatopus birmanus Olmi & Currado, 1976), female from Myanmar (from Olmi & Currado, 1976). D, E. Gonatopus daunus, holotype, frontal view (D) and dorsal view (E) (from Olmi 1984). Scale bar 2.27 mm for B, 1.55 mm for C, 0.94 mm for D and E, 0.36 mm for F.

PLATE 128. A, B. Male genitalia. C. Head, holotype. D, F, G. Chelae, holotypes. E. Scutum and metathorax + propodeum, holotype. A. Gonatopus javanus from Japan. B. G. maurus from China (from Xu et al. 2006c). C, D. G. lankanus (from Olmi 2003). E, F. G.lucidus (from Olmi 1984). G. G. maurus, holotype (from Olmi 1984). Scale bar 0.18 mm for A and G, 0.13 mm for B, 0.31 mm for D, 0.56 mm for E, 0.24 mm for F.

PLATE 131. A–D. Male genitalia. E,G, I. Mesosoma. F, H, J.Chelae, holotypes. A. Gonatopus rufoniger from China. B, I, J. G. superbus (from Olmi 1987c). C. G. asiaticus from Japan. D. G. leurosipropodeus, holotype (from He & Xu, 2002). E, F. G. schenklingi, holotype (from Olmi 1984). G, H. G. sitae (from Olmi 1984). Scale bar 0.19 mm for A, 0.18 mm for B, 0.15 mm for C, 0.17 mm for D, 0.21 mm for F, 0.34 mm for H, and 0.32 mm for J.