Deinodryinus R. Perkins, 1907

Olmi, Massimo, Copeland, Robert S. & Noort, Simon Van, 2019, Dryinidae of the Afrotropical region (Hymenoptera, Chrysidoidea), Zootaxa 4630 (1), pp. 1-619 : 217-220

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scientific name	Deinodryinus R. Perkins, 1907
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6. Genus Deinodryinus R. Perkins, 1907

Deinodryinus R. Perkins 1907: 45 ; Kieffer 1914b: 138; Olmi 1984: 118; 1999: 96; Olmi et al. 2000: 146; He & Xu 2002: 98; Virla & Olmi 2008: 369; Moya-Raygoza & Olmi 2010: 91; Azevedo et al. 2010: 880; Xu et al. 2013: 201: Olmi & Virla 2014: 108; Guglielmino & Olmi 2015: 230; Olmi & Xu 2015: 104.

Trisanteon Kieffer 1913a: 300 (synonymized by Olmi 1984); type species: Trisanteon hirticornis ( Kieffer 1911b) , by monotypy and original designation; Kieffer 1914b: 196.

Electrodryinus Ponomarenko 1975c: 126 (synonymized by Olmi 1984); type species: Electrodryinus areolatus Ponomarenko 1975c , by monotypy.

Prioranteon Olmi 1984: 589 (synonymized by Olmi 2007b); type species: Prioranteon casalei Olmi 1984 , by original designation; Olmi 1999: 148.

Type species. Deinodryinus paradoxus R. Perkins 1907 , by subsequent designation of Muesebeck & Walkley 1951.

Diagnosis. ♀: macropterous ( Figs 80A, B View FIGURE 80 ), or micropterous ( Fig. 84A View FIGURE 84 ); palpal formula 6/3; occipital carina complete; vertex of head frequently with two strong oblique keels connecting lateral ocelli to occipital carina; pronotum with distinct anterior collar and posterior disc ( Figs 80A, B View FIGURE 80 ); in macropterous ♀♀, forewing usually with distal part of 2r-rs&Rs vein longer than proximal part ( Figs 80A, B View FIGURE 80 ), less frequently as long as, or shorter than proximal part; enlarged claw without inner proximal prominence bearing bristles ( Fig. 83A View FIGURE 83 ), with one–two bristles or peg-like setae located further distally than proximal prominence ( Fig. 83A View FIGURE 83 ); tibial spurs 1/1/2. ♂ ( Figs 80C, D View FIGURE 80 ): always macropterous (even with micropterous ♀) ( Figs 80C, D View FIGURE 80 ); palpal formula 6/ 3; vertex of head frequently with two strong oblique keels connecting lateral ocelli to occipital carina; antennal setae usually much longer than breadth of antennomeres ( Fig. 80C View FIGURE 80 ), less frequently shorter; forewing usually with distal part of 2r-rs&Rs vein longer than proximal part ( Figs 80C, D View FIGURE 80 ), less frequently as long as, or shorter than proximal part; forewing usually with pterostigma than 4 × as long as broad or more; paramere without dorsal process, usually with one more or less large inner branch wrapping penis ( Fig. 88E View FIGURE 88 ), less frequently with one reduced inner branch ( Figs 86C, D View FIGURE 86 ); tibial spurs 1/1/2.

Distribution. Worlwide.

Hosts. Cicadellidae ( Guglielmino et al. 2013) .

World species. 165 species are known, 28 in the Afrotropical region.

Remarks. Fossil species of Deinodryinus were described by Olmi et al. (2010) and Guglielmino & Olmi (2011). Larvae of Deinodryinus were described by Guglielmino et al. (2017a). Leafhoppers parasitized by Deinodryinus species usually show dryinid sacs dorsally situated between the head and the prothorax ( Fig. 81A View FIGURE 81 ).

Key to species of Deinodryinus

♀♀

1. Micropterous ( Fig. 84A View FIGURE 84 )............................................................................... 2

- Macropterous ( Fig. 80A View FIGURE 80 )............................................................................... 8

2. Propodeal declivity granulate, not rugose, not transversely striate; mesoscutellum humped...................................................................................... D. granulatus Olmi, Copeland & van Noort , sp. nov.

- Propodeal declivity reticulate rugose or completely or partly transversely striate; mesoscutellum humped or flat.......... 3

3. Propodeal declivity completely transversely striate........................................... D. richardsi (Olmi)

- Propodeal declivity reticulate rugose, not or partly transversely striate........................................... 4

4. Metapectal-propodeal disc shiny, unsculptured................................................ D. casalei (Olmi)

- Metapectal-propodeal disc reticulate rugose or granulate...................................................... 5

5. Metapectal-propodeal disc granulate...................................................................... 6

- Metapectal-propodeal disc reticulate rugose................................................................ 7

6. Head testaceous, except ocellar area darkened or black; mesosoma partly testaceous and partly black or darkened; pronotum unsculptured........................................................................... D. capensis Olmi

- Head almost completely black; mesosoma black; pronotum granulate. D. nigropictus Olmi, Copeland & van Noort , sp. nov.

7. Mesosoma completely yellow-testaceous; enlarged claw with one peg-like lamella ( Fig. 92C View FIGURE 92 )............ D. paulyi (Olmi)

- Mesosoma mostly black almost completely black; enlarged claw with two bristles or peg-like setae ( Fig. 92E View FIGURE 92 )................................................................................................ D. prinslooi (Olmi)

8. Posterior surface of pronotum with sharp lateral margins ( Figs 81B View FIGURE 81 , 91A, B View FIGURE 91 )...................................... 9

- Posterior suface of pronotum with rounded lateral margins................................................... 14

9. Body with strong constriction between prothorax and mesothorax ( Figs 91A, B View FIGURE 91 ); frons with three median longitudinal keels (frontal line + two lateral keels joining lateral ocelli to antennal toruli)............................. D. guineensis Olmi

- Body without strong constriction between prothorax and mesothorax ( Figs 81B View FIGURE 81 , 94A View FIGURE 94 ); frons without frontal line, or with complete or incomplete frontal line, without lateral keels joining lateral ocelli to antennal toruli......................... 10

10. Head strongly punctate, unsculptured among punctures ( Fig. 82C View FIGURE 82 ); occasionally anterior third of frons so strongly punctate that it seems reticulate rugose.............................................................................. 11

- Head completely reticulate rugose, or at most with vertex not reticulate rugose................................... 13

11. Protarsomere 1 much shorter than 4; distal region of protarsomere 5 very slender ( Fig. 83A View FIGURE 83 ).................................................................................... D. ambrensis Olmi, Copeland & van Noort , sp. nov.

- Protarsomere 1 as long as, or slightly shorter than 4; distal region of protarsomere 5 robust ( Figs 92B, D View FIGURE 92 ).............. 12

12. Frontal line present in anterior half of frons................................................. D. orangeanus Olmi

- Frontal line absent.............................................. D. piceus Olmi, Copeland & van Noort , sp. nov.

13. Notauli reaching approximately 0.65 × length of mesoscutum.................................. D. insulanus (Benoit)

- Notauli reaching approximately 0.8–0.9 × length of mesoscutum................................... D. pulcher Olmi

14. Antennomere 3 approximately twice as long as 2; protarsomere 5 bearing small lamellae, some of which longer than others ( Figs 83E View FIGURE 83 , 91H View FIGURE 91 )..................................................................................... 15

- Antennomere 3 about 3 ×, or more than 3 × as long as 2..................................................... 16

15. Mesosoma reddish-brown or dark red; protarsomere 5 bearing mostly small lamellae, some of which are much longer than others, and with apex forming obtuse angle with rest of protarsomere ( Fig. 91H View FIGURE 91 )................... D. madecassus (Benoit)

- Mesosoma black; protarsomere 5 bearing small lamellae, all subequal in length, and with apex forming right angle with rest of protarsomere ( Fig. 83E View FIGURE 83 )................................................................. D. danielssoni Olmi

16. Protarsomere 5 bearing lamellae of approximately same length ( Fig. 83B View FIGURE 83 ); species mostly black.......... D. benoiti Olmi

- Protarsomere 5 bearing small lamellae, some of which longer than others ( Fig. 91G View FIGURE 91 ); species mostly testaceous-reddish................................................................................ D. madagascariensis (Benoit)

♂♂

1. Metapectal-propodeal disc with strong transverse posterior keel ( Fig. 96B View FIGURE 96 )....................................... 2

- Metapectal-propodeal disc without transverse posterior keel................................................... 6

2. Paramere very short, much shorter than penis ( Fig. 87C View FIGURE 87 )..................................... D. namorokensis Olmi

- Paramere very long, about as long as penis ( Fig. 87F View FIGURE 87 )........................................................ 3

3. Metapectal-propodeal disc very reduced; transverse posterior keel extending to anterior margin of metapectal-propodeal disc ( Fig. 89F View FIGURE 89 ); distal part of 2r-rs&Rs vein longer than proximal part................................... D. pulcher Olmi

- Metapectal-propodeal disc distinct and large; transverse posterior keel far from anterior margin of metapectal-propodeal disc ( Fig. 96B View FIGURE 96 ); distal part of 2r-rs&Rs vein much shorter than proximal part......................................... 4

4. Propodeal declivity without longitudinal keels...................................... D. sabaeus Olmi & van Harten

- Propodeal declivity with two complete longitudinal keels..................................................... 5

5. Head granulate and reticulate rugose...................................................... D. monticolus Olmi

- Head punctate, unsculptured among punctures............................................ D. umtamvunensis Olmi

6. Paramere very short, much shorter than penis ( Fig. 87C View FIGURE 87 )...................................................... 7

- Paramere very long, about as long as penis ( Fig. 86A View FIGURE 86 )........................................................ 8

7. Head reticulate rugose................................................................ D. namorokensis Olmi

- Head punctate, unsculptured among punctures, occasionally slightly granulate..................... D. danielssoni Olmi

8. Distal part of 2r-rs&Rs vein much or slightly shorter than proximal part......................................... 9

- Distal part of 2r-rs&Rs vein longer than, or as long as proximal part........................................... 11

9. Clypeus yellow-whitish............................................ D. musingilai Olmi, Copeland & Guglielmino

- Clypeus black or brown (rarely with anterior margin brown-testaceous)......................................... 10

10. Proximal inner branch wrapping penis narrow ( Fig. 87E View FIGURE 87 )......................................... D. paulyi (Olmi)

- Proximal inner branch wrapping penis broad ( Fig. 86A View FIGURE 86 )....................................... D. danielssoni Olmi

11. Paramere with one small apical branch not wrapping penis ( Figs 86C, D View FIGURE 86 )....................................... 12

- Paramere with one or two large apical branches wrapping penis ( Figs 86B, F View FIGURE 86 ).................................... 13

12. Paramere with short and slender distal inner branch ( Fig. 88D View FIGURE 88 )..................................... D. rusticus Olmi

- Paramere with large broad inner branch located along distal half of paramere ( Figs 86 View FIGURE 86 C-E).............. D. irreptus Olmi

13. Paramere with two apical branches wrapping penis ( Fig. 86B View FIGURE 86 ); volsellae situated between above two branches.......... 14

- Paramere with one apical branch wrapping penis ( Figs 86F View FIGURE 86 , 88A View FIGURE 88 , 89B, D View FIGURE 89 )....................................... 15

14. Inner apical branch of paramere with apex small ( Fig. 86B View FIGURE 86 )..................................... D. harinhalai Olmi

- Inner apical branch of paramere with apex very large ( Fig. 88F View FIGURE 88 )...................... D. sofiensis Guglielmino & Olmi

15. Apical branch wrapping penis hatchet blade shaped ( Fig. 86F View FIGURE 86 )............................... D. madecassus (Benoit)

- Apical branch wrapping penis not hatchet blade shaped ( Fig. 89D View FIGURE 89 )............................................. 16

16. Head very finely punctate; inner side of paramere not sculptured by papillae ( Fig. 89D View FIGURE 89 ).................. D. suavis Olmi

- Head reticulate rugose, or strongly punctate, with punctures large, deep and similar to areolae; inner side of paramere broadly sculptured or not by papillae ( Figs 88A View FIGURE 88 , 89B View FIGURE 89 )............................................................. 17 17. Apical branch wrapping penis curved, very long and slender ( Fig. 88A View FIGURE 88 ).............................. D. rusticus Olmi

- Apical branch wrapping penis not curved, short and broad ( Figs 89B, C View FIGURE 89 )............................. D. steineri Olmi

References

Azevedo, C. O., Madl, M. & Olmi, M. (2010) A Catalogue of the Bethylidae, Chrysididae, Dryinidae, Embolemidae, Sclerogibbidae and Scolebythidae (Hymenoptera: Chrysidoidea) of the Malagasy Subregion. Linzer biologische Beitrage, 42, 845 - 918.

Guglielmino, A. & Olmi, M. (2011) Revision of fossil species of Deinodryinus, with description of a new species (Hymenoptera, Dryinidae). ZooKeys, 130, 495 - 504. https: // doi. org / 10.3897 / zookeys. 130.1326

Guglielmino, A., Olmi, M. & Buckle, C. (2013) An updated host-parasite catalogue of world Dryinidae (Hymenoptera: Chrysidoidea). Zootaxa, 3740 (1), 1 - 113. https: // doi. org / 10.11646 / zootaxa. 3740.1.1

Guglielmino, A. & Olmi, M. (2015) Revision of the Afrotropical species of Deinodryinus Perkins, 1907 (Hymenoptera: Dryinidae), with description of a new species from Madagascar. African Invertebrates, 56, 229 - 259. https: // doi. org / 10.5733 / afin. 056.0116

Guglielmino, A., Olmi, M., Marletta, A. & Buckle, C. (2017 a) Larval morphology of three species of Anteoninae (Hymenoptera, Dryinidae). Zootaxa, 4320 (3), 470 - 486. https: // doi. org / 10.11646 / zootaxa. 4320.3.4

He, J. & Xu, Z. (2002) Hymenoptera Dryinidae. Fauna Sinica. Vol. 29. Science Press, Beijing, 464 pp.

Kieffer, J. J. (1911 b) Nouveaux Bethylides et Dryinides exotiques du British Museum de Londres. Annales de la Societe scientifique de Bruxelles, 35, 200 - 233.

Kieffer, J. J. (1913 a) Serphides de l'Ile de Lucon. Bollettino del Laboratorio di Zoologia generale e agraria della Regia Scuola superiore d'Agricoltura di Portici, 7, 189 - 192.

Kieffer, J. J. (1914 b) Bethylidae. Das Tierreich. Vol. 41. R. Friedlander und Sohn, Berlin, 595 pp.

Moya-Raygoza, G. & Olmi, M. (2010 [ 2008 ]) A catalogue of Dryinidae of Mexico, with descriptions of the opposite sexes of some species (Hymenoptera Chrysidoidea). Frustula entomologica, New Series, 31, 77 - 112.

Muesebeck, C. F. W. & Walkley, L. M. (1951) Family Dryinidae. In: Muesebeck, C. F. W., Krombein, K. V. & Townes, H. K. (Eds.), Hymenoptera of America North of Mexico. Synoptic catalogue. Agriculture Monograph 2. U. S. Department of Agriculture, Washington, D. C., pp. 1034 - 1043.

Olmi, M. (1984) A revision of the Dryinidae (Hymenoptera). Memoirs of the American Entomological Institute, 37, I-XII + 1 - 1913.

Olmi, M. (1999) Hymenoptera Dryinidae-Embolemidae. Fauna d'Italia. Vol. 37. Edizioni Calderini, Bologna, 425 pp.

Olmi, M. (2007 b) New species of Afrotropical Dryinidae (Hymenoptera: Chrysidoidea), with description of a new genus and a new subfamily. African Invertebrates, 48, 199 - 232.

Olmi, M. & Virla, E. G. (2014) Dryinidae of the Neotropical region (Hymenoptera: Chrysidoidea). Zootaxa, 3792, 1 - 534. https: // doi. org / 10.11646 / zootaxa. 3792.1.1

Olmi, M. & Xu, Z. (2015) Dryinidae of the Eastern Palaearctic region (Hymenoptera: Chrysidoidea). Zootaxa, 3996 (1), 1 - 253. https: // doi. org / 10.11646 / zootaxa. 3996.1.1

Perkins, R. (1907) Parasites of leaf-hoppers. Report of Work of the Experiment Station of the Hawaiian Sugar Planters' Association, Division of Entomology, Bulletin, No. 4, 5 - 59.

Ponomarenko, N. G. (1975 c) [A new hymenopteran species from Baltic amber]. Paleontologicheskiy Zhurnal, 1, 126 - 128. [in Russian; English translation: Paleontological Journal, 9 (1), 124 - 126]

Virla, E. G. & Olmi, M. (2008) Dryinidae. In: Claps, L. E., Debandi, G. & Roig-Junent, S. (Eds.), Biodiversidad de Artropodos Argentinos. Vol. 2. Sociedad Entomologica Argentina, Mendoza, pp. 357 - 372.

Xu, Z., Olmi, M. & He, J. (2013) Dryinidae of the Oriental region (Hymenoptera: Chrysidoidea). Zootaxa, 3614 (1), 1 - 460. https: // doi. org / 10.11646 / zootaxa. 3614.1.1

Figures

FIGURE 80. Habitus of Deinodryinus sp. A, B: ♀ in dorsal (A) and lateral (B) view. C, D: ♂ in lateral (C) and dorsal (D) view. Scale bar: A: 4.15 mm; B: 3.36 mm; C: 1.69 mm; D: 5.3 mm.

FIGURE 81. A: Leafhopper parasitized by Deinodryinus paulyi (Olmi). B: Deinodryinus ambrensis Olmi, Copeland & van Noort sp. nov.: ♀ holotype in dorsal view; mesoscutum sculpture. C, D: Deinodryinus casalei (Olmi), ♀ holotype in dorsal (C) and lateral (D) view. Scale bar: A: 2.13 mm.

FIGURE 82. Deinodryinus ambrensis Olmi, Copeland & van Noort sp. nov., ♀ holotype: habitus in dorsal (A) and lateral (B) view; head in frontal view (C); forewing (D).

FIGURE 83. Chelae of Deinodryinus species. A: D. ambrensis Olmi, Copeland & van Noort sp. nov., holotype; B: D. benoiti Olmi, holotype; C: D. capensis Olmi, holotype; D: D. casalei (Olmi), holotype; E: D. danielssoni Olmi, ♀ from South Africa, 32°57.732’S 18°05.985’E; F: D. granulatus Olmi, Copeland & van Noort sp. nov., holotype. Scale bar: A: 0.08 mm; B: 0.19 mm; C, D: 0.09 mm; E, F: 0.07 mm.

FIGURE 84. ♀♀ of Deinodryinus species. A: D. casalei (Olmi), holotype. B, C: D. granulatus Olmi, Copeland & van Noort sp. nov., holotype in dorsal (B) and lateral (C) view. Scale bar: A: 1.80 mm; C: 1.69 mm.

FIGURE 86. ♂ genitalia of Deinodryinus species. A: D. danielssoni Olmi, ♀ from South Africa, Skoorsteenkop. B: D. harin- halai Olmi, holotype. C: D. irreptus Olmi, holotype. D, E: D. irreptus Olmi, parameres of ♂♂ from Madagascar, 19°42.34’S 44°43.5’E (D) and 20°2.42’S 44°39.44’E (E). F: D. madecassus (Benoit, allotype. Scale bar: A: 0.12 mm; B, C, F: 0.15 mm; D, E: 0.08 mm.

FIGURE 87. ♂ genitalia of Deinodryinus species. A: D. monticolus Olmi, holotype; B: D. musingilai Olmi, Copeland & Guglielmino, holotype; C, D: D. namorokensis Olmi, holotype (C) and ♂ from Madagascar, 16°03.06’S 45°54.30’E (D); E: D. paulyi (Olmi) (holotype of D. nigerensis Olmi); F: D. pulcher Olmi, paratype. Scale bar: A, D: 0.12 mm; B: 0.09 mm; C, F: 0.10 mm; E: 0.08 mm.

FIGURE 88. ♂ genitalia of Deinodryinus species. A–D: D. rusticus Olmi: holotype (A); paramere of ♂♂ from Madagascar, 18°28.24’S 47°57.36’E (B), 24°39.13’S 43°59.48’E (C) and 24°14.26’S 46°17.16’E (D); E: D. sabaeus Olmi & van Harten, holotype; F: D. sofiensis Guglielmino & Olmi, paratype. Scale bar. A–C: 0.12 mm; D: 0.09 mm; E: 0.08 mm; F: 0.10 mm.

FIGURE 89. A–E. ♂ genitalia of Deinodryinus species. A–C: D. steineri Olmi: holotype (A); parameres of ♂♂ from Mada- gascar, 12°30.52’S 49°10.53’E (B) and 21°15.99’S 47°25.21’E (C); D: D. suavis Olmi, holotype; E: D. umtamvunensis Olmi, holotype. F: metapectal-propodeal disc + propodeal declivity of D. pulcher Olmi ♂ in dorsal view. Scale bar:A: 0.11 mm; B, C: 0.10 mm; D: 0.12 mm; E: 0.09 mm; F: 0.27 mm.

FIGURE 91. A–C: Deinodryinus guineensis Olmi, ♀ holotype: prothorax + mesoscutum in dorsal (A) and lateral (B) view; chela (C). Deinodryinus insulanus (Benoit), ♀ from Madagascar, Perinet: prothorax in dorsal (D) and lateral (E) view; chela (F). Chelae of Deinodryinus species holotypes: G: D. madagascariensis (Benoit); H: D. madecassus (Benoit). Scale bar:A: 0.57 mm; B: 0.96 mm; C: 0.08 mm; D: 1.42 mm; E: 2.55 mm; F, G: 0.12 mm; H: 0.09 mm.

FIGURE 92. Chelae of Deinodryinus species holotypes. A: D. nigropictus Olmi, Copeland & van Noort sp. nov.; B: D. orangeanus Olmi; C: D. paulyi (Olmi); D: D. piceus Olmi, Copeland & van Noort sp. nov.; E: D. prinslooi (Olmi); F: D. pulcher Olmi. Scale bar: A: 0.10 mm; B: 0.12 mm; C: 0.06 mm; D: 0.08 mm; E, F: 0.14 mm.

FIGURE 94. Habitus of Deinodryinus species. A: D. piceus Olmi, Copeland & van Noort sp. nov., ♀ holotype in dorsal view; head vertex sculpture. B–E: D. prinslooi (Olmi), ♀ from South Africa, 32°21.260’S 26°23.001’E: habitus in dorsal (B) and lateral (C) view; head and mesosoma in dorsal view (D); head in frontal view (E). D. pulcher Olmi, ♀ holotype in dorsal view. Scale bar: A: 1.48 mm; F: 2.62 mm.

FIGURE 96. A: Deinodryinus richardsi (Olmi), ♀ holotype chela; B: metapectal-propodeal disc + propodeal declivity of Deinodryinus sabaeus Olmi & van Harten, ♂ paratype. C–D: ♂ genitalia of Lonchodryinus hovanus Olmi: holotype (C) and ♂ from Madagascar, 12°31.13’S 49°10.46’E (D). E–F: chelae of Lonchodryinus species holotypes: L. madagascolus Olmi, Copeland & van Noort sp. nov. (E) and L. pauliani (Benoit) (F). Scale bar: A: 0.11 mm; B: 0.20 mm; C, D, F: 0.13 mm; E: 0.11 mm.