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Research Article
The genus Ismarus Haliday (Hymenoptera, Ismaridae) from China
expand article infoXiao Zhang, Hua-yan Chen, Jing-xian Liu§, Shi-xiao Luo|
‡ Sun Yat-sen University, Guangzhou, China
§ South China Agricultural University, Guangzhou, China
| South China Botanical Garden, Chinese Academy of Sciences, Guangzhou, China
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Abstract

The genus Ismarus Haliday from China is revised. Six species are recognized, of which two species are described as new: Ismarus areolatus Chen, sp. nov., and Ismarus paradorsiger Chen, sp. nov. By examining the pits along the anterolateral margin of mesoscutum and comparing them with the typical notauli in Diapriidae, we conclude that the so-called reduced notauli in Ismarus are actually foveae of the mesoscutual suprahumeral sulcus. An updated key to the Chinese species of the genus is provided.

Keywords

Diaprioidea, hyperparasitoid, key, new species

Introduction

Ismarus Haliday is the only genus in Ismaridae, with 57 described species worldwide (Masner 1976; Johnson 1992; Liu et al. 2011; Comério et al. 2016; Kolyada and Chemyreva 2016; Kim et al. 2018a, b). Available biological data suggest that species of Ismarus are hyperparasitoids of Dryinidae, which are parasitoids of Auchenorrhyncha (Hemiptera), such as leafhoppers, planthoppers and treehoppers (Chambers 1955, 1981; Nixon 1957; Wall 1967; Kozlov 1971; Masner 1976; Jervis 1979; Tussac and Tussac 1991; Olmi 2000). In addition, studies suggested species of Ismarus are likely to be found from wooded areas at higher elevations in warmer climatic zones and at low elevations in cooler climatic zones (Masner 1976; Kim et al. 2018a, b). However, only six species of Ismarus have been recorded from the Oriental regions (Liu et al. 2011; Izadizadeh et al. 2020). The Oriental fauna of Ismarus is still clearly undersampled, considering the diversity of Dryinidae (Xu et al. 2013) and preferred habitats in these regions. In the present study, we examine the species of Ismarus from China, including descriptions of two new species.

Materials and methods

This work is based upon the specimens in the following collections, with abbreviations used in the text: SCAU, Hymenoptera Collection, South China Agricultural University, Guangzhou, China; SYSBM, Sun Yat-sen University, The Museum of Biology, Guangzhou, China.

Abbreviations and morphological terms used in text:

A1, A2, ... A12 antennomere 1, 2, … 12;

POL postocellar line (shortest distance between both posterior ocelli);

OOL oculoocellar line (shortest distance between posterior ocellus and compound eye);

MT Malaise trap;

YPT yellow pan trap;

T1, T2, ... T8 metasomal tergite 1, 2, ... 8.

Morphological terminology otherwise follows Masner (1976). Species of Ismarus from the Oriental and Palearctic regions recently have been quite thoroughly reviewed and keyed (Liu et al. 2011; Kolyada and Chemyreva 2016; Kim et al. 2018a; Izadizadeh et al. 2020), which allow us easily identify the Chinese species in this study.

Specimens were examined using a Nikon SMZ800N microscope. Images and measurements were made using a Nikon SMZ25 microscope with a Nikon DS-Ri 2 digital camera system. Scanning electron micrographs were produced using a Phenom Pro Desktop SEM and a single montage images were generated from image stacks in the program Helicon Images were post-processed with Abobe Photoshop CS6 Extended.

Taxonomy

Ismarus Haliday, 1835

Ismarus Haliday, 1835: 467. Type species Cinetus dorsiger Haliday, 1831, by monotypy.

Entomia Herrich-Schäffer, 1840: 127. Type species Entomia campanulata Herrich-Schäffer, 1840, by monotypy.

Agonophorus Dahlbom, 1858: 289. Type species Ismarus rugulosus Förster, 1850, designated by Muesebeck (1972).

Remarks

In the revision of the New World Ismarinae (then was treated as a subfamily of Diapriidae and contained the only genus, Ismarus), Masner (1976) pointed out that Ismarus “is usually characterized by the low insertion of antennae, transverse head, and the absence (better reduction) of notauli”. This suggests that he was not certain about the status of notauli within the genus. However, in the key to subfamily of Diapriidae and the descriptions of the New World species, he continued to state that “notauli were always present but reduced to anterior pits”. Subsequent researchers followed Masner’s diagnosis of the genus and described these pits as notauli (Liu et al. 2011; Kolyada and Chemyreva 2016; Kim et al. 2018a, b), except Kolyada and Chemyreva (2016) described the anterior pits as the reduced notauli and the rest of the pits along the anterolateral margin of mesoscutum as part of the mesoscutual suprahumeral sulcus. We here examine these pits (Fig. 1A, B) using scanning electron microscopy and compare them with the notauli in a species of Coptera Say (Diapriidae), which has typical notauli (Fig. 1C). The pits in Ismarus are along the anterolateral margin of mesoscutum where the mesoscutual suprahumeral sulcus is located in many groups of Hymenoptera and are different from the pits of the typical notauli such as found in Coptera. The anterior pits in Coptera is located at the anterior margin of the longitudinal grooves of the notauli and apparently are a part of the notauli, while the pits in Ismarus (if they are present) are parallel to the anterolateral margin of the mesoscutum and never present as a part of any longitudinal grooves of the tmesoscutum. Therefore, we here conclude that the pits present along the anterolateral margin of the mesoscutum in Ismarus constitute part of the mesoscutual suprahumeral sulcus and are not notauli. The mesoscutual suprahumeral sulcus can be totally absent (as in Ismarus dorsiger (Haliday)), present as a single pit on each side (as in Ismarus halidayi Foerster, Fig. 1B), or present as a few pits of varying size (as in Ismarus nigritrochanter Liu, Chen & Xu, Fig. 1A).

Table 1.

An updated list of the Chinese species of Ismarus with distribution in China.

Species Distribution in China
Province Ecozone
Ismarus areolatus Chen, sp. nov. Guangdong Oriental
Ismarus halidayi Foerster Sichuan, Guizhou, Yunnan, Tibet Oriental
Ningxia Palaearctic
Ismarus longus Liu, Chen & Xu Yunnan Oriental
Ismarus nigritrochanter Liu, Chen & Xu Yunnan Oriental
Ismarus paradorsiger Chen, sp. nov. Guangdong Oriental
Ismarus parvicellus Liu, Chen & Xu Hainan Oriental
Figure 1. 

Mesosoma, dorsal view, mshs: mesoscutual suprahumeral sulcus; nt: notaulus A Ismarus nigritrochanter Liu, Chen & Xu B Ismarus halidayi Foerster C Coptera sp.

Key to species of Ismarus from China

Females

1 Body mostly pale yellowish (Figs 12, 14); suture between T2 and T3 incomplete (Fig. 12F) I. paradorsiger Chen, sp. nov.
Body entirely black (Figs 2A, 4A, 6, 7A, 9A, 11A, 15A); suture between T2 and T3 complete (Figs 2F, 4F, 7F, 9F, 15F) 2
2 Mesoscutellum areolate (Fig. 2C); lateral pronotal area with a submedian oblique carina (Fig. 2D) Ismarus areolatus Chen, sp. nov.
Mesoscutellum smooth (Figs 4C, 7C, 9C, 15C); lateral pronotal area without carina (Figs 4D, 7D, 9D, 15D) 3
3 Radial cell of fore wing as long as marginal vein (Fig. 5A); median furrow of T2 long, reaching half length of T2 (Fig. 4E) Ismarus halidayi Foerster
Radial cell of fore wing distinctly shorter than marginal vein (Figs 8A, 10A, 16A); median furrow of T2 very short, distinctly less than half length of T2 (Figs 7E, 9E, 15E) 4
4 Mesoscutal suprahumeral sulcus present as a single anterior pit (Fig. 15C); anterior mesoscutellar pit large, longer than length of the mesoscutellar disc (Fig. 15C); radial cell of fore wing small, 0.3 × length of marginal vein (Fig. 16A) Ismarus parvicellus Liu, Chen & Xu
Mesoscutal suprahumeral sulcus present as four or five foveae of varying size (Figs 7C, 9C); anterior mesoscutellar pit distinctly shorter than length of the mesoscutellar disc (Figs 7C, 9C); radial cell of fore wing larger, 0.6 × length of marginal vein (Figs 8A, 10A) 5
5 Second flagellomere 5.0 × as long as wide (Fig. 8B); radial cell of fore wing 3.0 × as long as high (Fig. 8A); antenna black with scape brown, pedicel and first flagellomere dark brown (Fig. 8B); all trochanters brown (Fig. 7A) Ismarus longus Liu, Chen & Xu
Second flagellomere 3.5 × as long as wide (Fig. 10B); radial cell of fore wing 2.0 × as long as high (Fig. 10A); antenna uniformly black (Fig. 10B); all trochanters black (Fig. 9A) Ismarus nigritrochanter Liu, Chen & Xu

Ismarus areolatus Chen, sp. nov.

Figures 2, 3

Diagnosis

This species can be easily distinguished from other Oriental species of Ismarus by the following characters: lateral pronotal area with a submedian oblique carina; mesoscutellum areolate and posterior rim excavate with slightly prominent posterolateral corners; hind tibiae abruptly incrassate.

Description

Female. Body length 2.24–3.15 mm.

Colour. Body black; A1A2 brown, remainder of antenna black; coxae of fore and mid legs black, remainder brown to yellow, with tibia and tarsi becoming paler, hind leg mostly black with trochanter dark brown; wings hyaline, veins blackish brown.

Figure 2. 

Ismarus areolatus Chen, sp. nov., holotype, female (SCAU 3049372) A lateral habitus B head, anterior view C head and mesosoma, dorsal view D head and mesosoma, lateral view E propodeum and basal metasoma, dorsal view F apical metasoma, dorsal view.

Head. Head in dorsal view 2.0 × as wide as long; vertex abruptly sloping behind ocelli; POL as long as OOL; frons with ventrolateral areas densely setose, area below anterior ocellus with scattered setae; facial transverse carina slightly convex ventrally; A3 slightly longer than A4; A4 1.2 × length of A5; A6–A14 with each segment less than 1.5 × as long as wide; A15 approximately 2.0 × longer than wide.

Mesosoma. Dorsal pronotal area rugose-punctate and setose; lateral pronotal area with a submedian oblique carina, rugose ventrally, smooth dorsally; mesoscutum smooth and convex, posterior margin with relatively dense long setae; mesoscutal suprahumeral sulcus absent; humeral sulcus deep and crenulate, 1.7 × length of tegula; mesoscutellum areolate and slightly convex, posterior rim excavate with slightly prominent posterolateral corners; anterior mesoscutellar pit large and deep, as long as length of the mesoscutellar disc, distinctly crenulate posteriorly, median keel strong; mesopleuron smooth and bare, with area below tegula rugulose; metapleuron rugose and covered with dense whitish long setae.

Figure 3. 

Ismarus areolatus Chen, sp. nov., holotype, female (SCAU 3049372) A wings B antenna C hind tibia.

Wings. Radial cell completely closed, very small, 4.4 × as long as wide and 0.2 × as long as marginal vein.

Legs. Fore and mid legs slender; hind tibia abruptly incrassate, its maximum width slightly wider than hind femur.

Metasoma. Petiole distinctly transverse, with strong costae dorsally; base of T2 with six short costae, median furrow short, 0.15× length of T2, remainder of T2 smooth with scattered fine punctures; T3–T7 smooth with scattered fine punctures; T8 coriaceous; sutures between tergites complete and deeply impressed.

Male. Unknown.

Etymology

The name refers to the areolate mesoscutellum.

Material examined

Holotype, female: China: Guangdong, Guangzhou, South China Botanical Garden, forest, 23°10'49.87"N, 113°21'21.33"E, MT1, 7.iv–7.v.2020, Jian-wei Li, SCAU 3049372 (deposited in SYSBM). Paratype: (1 female) China: Guangdong, Maoming, Yunkaishan National Nature Reserve, LSX591, MT-GD32, 22°16'38.37"N, 111°11'45.23"E, 30.iv–29.v.2020, Long-long Chen, SCAU 3049371 (SYSBM).

Ismarus halidayi Förster

Figure 1B, 4, 5, 6

? Entomia campanulata Herrich-Schäffer, 1840: 127 (nomen dubium).

Ismarus halidayi Förster, 1850: 285.

Ismarus longicornis Thomson, 1858: 378. Synonymized by Kolyada and Chemyreva (2016). Ismarus mongolicus Szabó, 1974: 23. Synonymized by Kolyada and Chemyreva (2016).

Material examined

Other material : (11 females, 6 males) China: 2 females and 1 male, Ningxia, Mt. Liupanshan, 35.40°N, 106.38°E, 11–12.vii.2008, Jie-min Yao, Nos. 200808622, 200808859, 200808017 (SCAU); 1 male, Ningxia, Mt. Liupanshan, 35.40°N, 106.38°E, 13–14.vii.2009, Hua-yan Chen, No. 200903337 (SCAU); 2 females, Ningxia, Mt. Liupanshan, 35°29'12"N, 106°20'29"E, 3–14.vii.2009, Huayan Chen, SCAU 3042197, 3042198 (SYSBM); 1 female, Sichuan, Mt. Ermei, 29.61°N, 103.36°E, 7.vii.2009, Jiang-li Tan, No. 200903977 (SCAU); 1 female, Sichuan, Luhuo, 31.38°N, 100.66°E, 30.vi.2009, Jiang-li Tan, No. 200903953 (SCAU); 1 female, Guizhou, Mt. Fanjingshan, 27.92°N, 108.70°E, 2100 m, 30.vii.2001, Yun Ma, No. 200109552 (SCAU); 1 female, Guizhou, Mt. Fanjingshan, 27.92°N, 108.70°E, 12.vii.1993, Song-lin Yao, No. 936734 (SCAU); 1 female, Yunnan, Yongshan County, Huanghua Town, 28.00°N, 103.51°E, 1500 m, 8.x.2010, Wei Dong (SCAU); 1 male, Yunnan, Dali, Mt. Cangshan, 25.63°N, 100.16°E, 5.vi.2009, Jiang-li Tan, No. 200901192 (SCAU); 1 male, Yunnan, Dali, Yunlong County, Tianchi, 25.848602°N, 99.234147°E, 2680.1 m, MT-N1-1, 12–28.vi.2020, You-jing Gong, SCAU 3049366 (SYSBM); 1 female, Yunnan, Dali, Yunlong County, Tianchi, 25.854158°N, 99.239927°E, 2933.3 m, MT-N3-3 14–28.vi.2020, You-jing Gong, SCAU 3049335 (SYSBM); 1 female, 1 male, Yunnan, Dali, Yunlong County, Tianchi, 25.848602°N, 99.234147°E, 2680.1 m, MT-N1-1, 12-28.vi.2020, You-jing Gong, SCAU 3049339 3049340 (SYSBM); 1 male, Tibet, Milin, 29.18°N, 94.20°E, 14.vi.2009, Jiang-li Tan, No.200902367 (SCAU).

Figure 4. 

Ismarus halidayi Foerster, female (SCAU 3044335) A lateral habitus B head, anterior view C head and mesosoma, dorsal view D head and mesosoma, lateral view E propodeum and basal metasoma, dorsal view, arrow indicates median furrow F apical metasoma, dorsal view.

Figure 5. 

Ismarus halidayi Foerster, female (SCAU 3044335) A wings B antenna C hind tibia.

Distribution. China (Ningxia, Sichuan, Guizhou, Yunnan, Tibet).

Figure 6. 

Ismarus halidayi Foerster, male (SCAU 3042197) Lateral habitus.

Ismarus longus Liu, Chen & Xu

Figures 7, 8

Ismarus longus Liu, Chen & Xu, 2011: 51, 53, 59 (original description, keyed, placed in halidayi species group, keyed).

Material examined

Holotype, female: China: Yunnan, Yingjiang, Tongbiguan, 24.60°N, 97.65°E, 2009.v.20, Man-man Wang, No. 200900933 (deposited in SCAU). Paratypes: 1 female, same data as type, No. 200900492 (SCAU); 1 female, Yunnan, Tengchong, Jietou Town, 25.40°N, 98.70°E, 2009.v.13, Man-man Wang, No. 200902486 (SCAU); 1 female, Yunnan, Tengchong, Jietou Town, 25.40°N, 98.70°E, 2009.v.12, Jie Zeng, No. 200902519 (SCAU).

Figure 7. 

Ismarus longus Liu, Chen & Xu, holotype, female (200900933) A lateral habitus B head, anterior view C head and mesosoma, dorsal view D head and mesosoma, lateral view E propodeum and basal metasoma, dorsal view F apical metasoma, dorsal view.

Distribution

China (Yunnan).

Figure 8. 

Ismarus longus Liu, Chen & Xu, holotype, female (200900933) A wings B antenna C hind tibia.

Ismarus nigritrochanter Liu, Chen & Xu

Figures 1A, 9, 10, 11

Ismarus nigritrochanter Liu, Chen & Xu, 2011: 53, 54, 59 (original description, keyed, placed in halidayi species group, keyed).

Coments

This species was originally described based on a singleton female (Liu et al. 2011). Here we record the male: Body length 1.9–2.0 mm (n = 2). Similar to female, trochanters brown; A4 without blade-like carina; A6–A13 with each segment 1.4 × as long as wide.

Figure 9. 

Ismarus nigritrochanter Liu, Chen & Xu, holotype, female (200700989) A lateral habitus B head, anterior view C head and mesosoma, dorsal view D head and mesosoma, lateral view E propodeum and basal metasoma, dorsal view F apical metasoma, dorsal view.

Material examined

Holotype, female. China: Yunnan, Mt. Gaoligongshan, 25.98°N, 98.80°E, 20–21.vii.2006, Zhong-shi Zhou, No. 200700989 (deposited in SCAU). Other material: (2 males) China: 1 male, Yunnan, Diqing, Shangri-la, Jiantang Town, 27°31'45.45"N, 100°5'29.06"E, 2637 m, pine forest, CHY2, MT1, 12.vii–14.viii.2020, Nima, SCAU 3049368 (SYSBM); 1 male, Yunnan, Diqing, Shangri-la, Jiantang Town, 27°31'53.43"N, 100°4'48.75"E, 2643 m, pine forest, CHY6, MT2, 14.viii–18.ix.2020, Nima, SCAU 3044336 (SYSBM).

Figure 10. 

Ismarus nigritrochanter Liu, Chen & Xu, holotype, female (200700989) A wings B antenna C hind tibia.

Distribution

China (Yunnan).

Figure 11. 

Ismarus nigritrochanter Liu, Chen & Xu, male (SCAU 3044336) A lateral habitus B antenna.

Ismarus paradorsiger Chen, sp. nov.

Figures 12, 13, 14

Diagnosis

This species is most similar to the widely distributed I. dorsiger and I. kakamegensis in size and color patterns. It can be easily separated from I. dorsiger by the presence of a mesoscutal suprahumeral sulcus, and from I. kakamegensis by A1, which is distinctly longer than the length of A3 and A4.

Figure 12. 

Ismarus paradorsiger Chen, sp. nov., holotype, female (SCAU 3042199) A lateral habitus B head, anterior view, arrow indicates facial transverse carina convex ventrally C head and mesosoma, dorsal view D head and mesosoma, lateral view E propodeum and basal metasoma, dorsal view F apical metasoma, dorsal view, arrow indicates suture between T2 and T3 obsolete.

Description

Female. Body length 1.95–2.3 mm.

Colour. Body mostly pale yellowish, except mesoscutum, scutellum, metanotum, dorsal T1 and basal T2 variably dark brown; legs pale yellow, except hind tibia and basal tarsus somewhat brown; A1A2 brown, remainder of antenna dark brown; wings hyaline or slightly infuscate medially, veins blackish brown.

Figure 13. 

Ismarus paradorsiger Chen, sp. nov., holotype, female (SCAU 3042199) A wings B antenna C hind tibia.

Head. Head in dorsal view 2.0 × as wide as long; vertex abruptly sloping behind ocelli; POL as long as OOL; frons with ventrolateral areas densely setose, area below anterior ocellus with scattered setae; facial transverse carina slightly convex ventrally; A3 slightly shorter than A4; A4 1.2× length of A5; A6–A14 with each segment approximately 2.0× longer than wide; A15 approximately 3.0× longer than wide.

Mesosoma. Dorsal pronotal area rugose-punctate and setose; lateral pronotal area rugose-punctate ventrally, smooth dorsally; mesoscutum smooth and bare, convex; mesoscutal suprahumeral sulcus present as six small pits; humeral sulcus deep and finely crenulate, slightly shorter than length of tegula; mesoscutellum smooth and slightly convex, posterior rim rounded; anterior mesoscutellar pit small, 0.25 × length of the mesoscutellar disc, distinctly crenulate at bottom, median keel strong; mesopleuron smooth and bare; metapleuron rugose and covered with dense whitish long setae.

Figure 14. 

Ismarus paradorsiger Chen, sp. nov., paratype, female (201000305) Lateral habitus.

Wings. Radial cell completely closed, moderately large, 3.6× as long as wide and as long as marginal vein.

Legs. Fore and mid legs slender; hind tibia abruptly incrassate, its maximum width slightly wider than hind femur.

Metasoma. Petiole distinctly transverse, with irregular longitudinal costae dorsally; base of T2 with five short costae, median furrow short, 0.1× length of T2, remainder of T2 smooth with scattered fine punctures; T3–T8 smooth with scattered fine punctures; suture between T2 and T3 obsolete but the following sutures between tergites distinctly impressed.

Male. Unknown.

Etymology

The name refers to the high degree of similarity with I. dorsiger and is intended to be used as a noun in apposition.

Material examined

Holotype, female: China: Guangdong, Maoming, Yunkaishan National Nature Reserve, LSX704, MT-GD33, 22°17'40.72"N, 111°12'37.97"E, 1480 m, 4–30.vii.2020, Long-long Chen, SCAU 3042199 (deposited in SYSBM). Paratypes: (8 females) China: 1 female, Guangdong, Maoming, Yunkaishan National Nature Reserve, LSX591, MT-GD32, 22°16'38.37"N, 111°11'45.23"E, 30.iv–29.v.2020, Longlong Chen, SCAU 3049370 (SYSBM); 7 females, Yunnan, Yongshan County, Huanghua Town, 28.00°N, 103.51°E, 1500 m, 8.x.2010, Wei Dong, [misidentified as Ismarus dorsiger (Haliday)] Nos. 201000305, 201000308, 201000311, 201000312, 201000314, 201000316, 201000317 (SCAU).

Distribution

China (Guangdong, Yunnan).

Comments

Liu et al. (2011) reported additional distribution records of Ismarus dorsiger (Haliday, 1831) from Yunnan Province of China. However, when we examined the 9 females identified as Ismarus dorsiger (Haliday, 1831) (deposited in SCAU), it turns out that these specimens have mesoscutal suprahumeral sulci, which are absent in I. dorsiger as was documented by Kolyada and Chemyreva (2016) and Kim et al. (2018a, b). Based on additional material collected from Guangdong, we describe them here as a new species. Therefore, I. dorsiger is still unknown from China.

Ismarus parvicellus Liu, Chen & Xu

Figures 15, 16

Ismarus parvicellus Liu, Chen & Xu, 2011: 55, 56, 58 (original description, keyed, placed in halidayi species group, keyed).

Material examined

Holotype, female: China: Hainan, Baisha County, Mt. Jiujialing, 19.21°N, 109.45°E, 2010.VII.18, YPT, Hua-yan Chen, No. 20100013 (deposited in SCAU).

Figure 15. 

Ismarus parvicellus Liu, Chen & Xu, holotype, female (201010013) A lateral habitus B head, anterior view C head and mesosoma, dorsal view, amp: anterior mesoscutellar pit D head and mesosoma, lateral view E propodeum and basal metasoma, dorsal view F apical metasoma, dorsal view.

Distribution

China (Hainan).

Figure 16. 

Ismarus parvicellus Liu, Chen & Xu, holotype, female (201010013) A wings B antenna C hind tibia.

Acknowledgements

Thanks to Zachary Lahey (The Ohio State University) and Elijah J. Talamas (Florida Department of Agriculture and Consumer Services) for providing valuable discussion on morphological characters. This material is based upon work supported in part by the Program of Ministry of Science and Technology of China (2018FY100406) and Guangdong Basic and Applied Basic Research Foundation (2019A1515110610).

References

  • Chambers VH (1955) Some hosts of Anteon spp. (Hym. Dryinidae) and a hyperparasite Ismarus (Hymenoptera. Berytidae). Entomologist’s Monthly Magazine 91: 114–115.
  • Chambers VH (1981) A host for Ismarus halidayi Foerst. (Hym., Diapriidae). Entomologist’s Monthly Magazine 117: 29.
  • Comério EF, Perioto NW, Lara RIR (2016) A New Species of Ismarus Haliday (Hymenoptera: Ismaridae) from Brazil and a New Occurrence Record for Ismarus gracilis Masner. EntomoBrasilis 9(3): 197–201. https://doi.org/10.12741/ebrasilis.v9i3.589
  • Foerster A (1850) Eine Centurie neuer Hymenopteren. Verhandlungen des Naturhistorischen Vereins de Preussischen Rheinlande Westfalens 7: 277–288.
  • Haliday AH (1835) Essay on parasitic Hymenoptera of the Ichneumones Adsciti. Entomological Magazine 2: 458–468.
  • Izadizadeh M, Talebi AA, Kolyada V, Farahani S, Kazerani F, Ameri A (2020) Review of the family Ismaridae Thomson, 1858 (Hymenoptera: Diaprioidea) from Iran. Oriental Insects 1–11. https://doi.org/10.1080/00305316.2020.1758821
  • Jervis MA (1979) Parasitism of Aphelopus species (Hymenoptera: Dryinidae) by Ismarus dorsiger (Curtis) (Hymenoptera: Diapriidae). Entomologist’s Gazette 30: 127–129.
  • Johnson NF (1992) Catalog of Worlds Species of Proctotrupoidea, Exclusive of Platygastridae (Hymenoptera). Memoirs of the American Entomological Institute 51: 1–825.
  • Kim C-J, Copeland RS, Notton DG (2018a) The family Ismaridae Thomson (Hymenoptera, Diaprioidea): first record for the Afrotropical region with description of fourteen new species. African Invertebrates 59(2): 127–163. https://doi.org/10.3897/afrinvertebr.59.24403
  • Kim CJ, Notton DG, Ødegaard F, Lee JW (2018b) Review of the Palaearctic species of Ismaridae Thomson, 1858 (Hymenoptera: Diaprioidea). European Journal of Taxonomy 417: 1–38. https://doi.org/10.5852/ejt.2018.417
  • Kolyada VA, Chemyreva VG (2016) Revision of species of the genus Ismarus Haliday, 1835 (Hymenoptera: Diaprioidea: Ismaridae) of the Russian fauna. Far Eastern Entomologist 318: 1–19.
  • Kozlov MA (1971) Proctotrupoids (Hymenoptera, Proctotrupoidea) of the USSR. Trudy Vsesoyuznogo Entomologicheskogo Obshchestva 54: 3–67.
  • Muesebeck CFW (1972) On the identity of Agonophorus Dahlbom (Hymenoptera: Diapriidae). Proceedings of Entomological Society of Washington 74: 131.
  • Nixon GEL (1957) Hymenoptera, Proctotrupoidea, Diapriidae subfamily Belytinae. Handbooks for the Identification of British Insects 8(3dii): 1–107.
  • Olmi M (2000) Bio-ecologia degli Imenotteri Driinidi e loro impiego in programmi di lotta biologica. In: Lucchi A (Ed.) La Metcalfa negli ecosistemi italiani. ARSIA, Firenze, 93–117.
  • Szabó JB (1974) Neue Arten und Gattungen der Diapriiden aus der Mongolei (Hymenoptera. Diapriidae). Annales Historico-Naturalesmusei Nationalis Hungarici 66: 353–358.
  • Thomson CG (1858) Skandinaviens proctotruper. Belytinae, Ismarinae. Öfversigt af Kongliga Ventenskaps-Akadamiens Förhandlingar 15: 359–380.
  • Tussac H, Tussac M (1991) Récapitulatif d’une collecte de Dryinidae et Diapriidae (Hym. Chrysidoidea et Proctotrypoidea). Entomologiste 47(4): 189–194.
  • Wall I (1967) Die Ismarinae und Belytinae der Schweiz. Entomologische Abhandlungen 35: 123–265.