New records for a catalogue of Chalcididae (Hymenoptera, Chalcidoidea) from Morocco

Abstract

New records for a catalogue of Chalcididae (Hymenoptera, Chalcidoidea) from Morocco
Three species of Chalcididae (Hymenoptera, Chalcidoidea) were newly recorded from Morocco during a study carried out in the Maâmora forest between 2012 and 2014: Hockeria bifasciata (Walker, 1834), H. mengenillarum (Silvestri, 1943) and Proconura decipiens (Masi, 1929). P. decipiens (Masi, 1929) stat. rev. will be removed from synonymy with P. nigripes (Fonscolombe, 1832). This study includes bibliographical research and revision of specimens deposited in the National Museum of Natural History, Scientific Institute of Rabat (Morocco). Twenty-six species and fourteen genera belonging to the family Chalcididae (Hymenoptera, Chalcidoidea) are now catalogued from Morocco.

Data published through GBIF (Doi: 10.15470/nochzr)

Key words: Hymenoptera, Chalcididae, New data, Maâmora, Morocco

Resumen

Nuevos registros para un catálogo de Chalcididae (Hymenoptera, Chalcidoidea) de Marruecos
Se han registrado tres nuevas especies de Chalcididae (Hymenoptera, Chalcidoidea) en Marruecos, a partir de un estudio realizado en el bosque de Maâmora entre 2012 y 2014: Hockeria bifasciata (Walker, 1834), H. mengenillarum (Silvestri, 1943) y Proconura decipiens (Masi, 1929). P. decipiens (Masi, 1929) stat. rev. dejará de considerarse sinónimo de P. nigripes (Fonscolombe, 1832). Este estudio se ha completado con investigación bibliográfica y la revisión de los especímenes depositados en el Museo Nacional de Historia Natural, Instituto Científico de Rabat (Marruecos). El artículo hace referencia a veintiséis especies y catorce géneros pertenecientes a la familia Chalcididae (Hymenoptera, Chalcidoidea) presentes en Marruecos.

Datos publicados en GBIF (Doi: 10.15470/nochzr)

Palabras clave: Hymenoptera, Chalcididae, Nuevos datos, Maâmora, Marruecos

Resum

Nous registres per a un catàleg de Chalcididae (Hymenoptera, Chalcidoidea) del Marroc
S’han registrat tres noves espècies de Chalcididae (Hymenoptera, Chalcidoidea) al Marroc, a partir d’un estudi portat a terme al bosc de Maâmora entre 2012 i 2014: Hockeria bifasciata (Walker, 1834), H. mengenillarum (Silvestri, 1943) i Proconura decipiens (Masi, 1929). P. decipiens (Masi, 1929) stat. rev. deixarà de ser considerat sinònim de P. nigripes (Fonscolombe, 1832). Aquest estudi s’ha completat amb recerca bibliogràfica i la revisió dels espècimens dipositats al Museu Nacional d’Història Natural, Institut Científic de Rabat (Marroc). L’article fa referència a vint-i-sis espècies i catorze gèneres pertanyents a la família Chalcididae (Hymenoptera, Chalcidoidea) presents al Marroc.

Dades publicades a GBIF (Doi: 10.15470/nochzr)

Paraules clau: Hymenoptera, Chalcididae, Noves dades, Maâmora, Marroc

Introduction

Chalcididae Latreille, 1817 (Hymenoptera, Chalcidoidea) is a moderately large family with worldwide distribution. The family currently contains 85 genera and 1,564 species placed in five subfamilies (Noyes, 2019). All Chalcididae increase in abundance on approaching the Mediterranean rim (De Lombaerde et al., 2002). All Chalcididae are parasitoids of larvae or pupae of other insects, mostly Lepidoptera and Diptera (Delvare, 1995) and are described as host group specific endoparasitoids (Dajoz, 2010) of Lepidoptera, Diptera, Coleoptera larvae, and even some species of Tenthredinidae (Hymenoptera, Symphyta) and some Neuroptera (Bouček, 1988; Herting, 1978). One-hundred and twenty-two hosts can be parasitized by 25 species of Chalcididae; of these, 89 are Lepidoptera, 16 Diptera, 13 Hymenoptera, and four Coleoptera (Ferrer, 2010). The literature concerning Chalcididae in Morocco is limited to notes on just one or a few species. The first, Neochalis barbara Benoist, 1921 (= Neochalcis fertoni Kieffer, 1899) was reported in 1921 by Benoist from a Casablanca aviation camp (Casablanca-Settat region), and this was followed by an account by De Lépiney (1927, 1930) of Brachymeria tibialis (Walker, 1834) as a parasitoid of the gypsy moth Lymantria dispar (Linnaeus, 1758) (Lepidoptera, Lymantriidae) as a defoliator causing serious damage to cork oak. Other species, mainly hosts of this family, have been cited by Bléton and Fieuzet (1939), Steffan (1948), Bouček (1952), Chakir and Fraval (1985), Villemant (1989), Villemant and Fraval (1993). Delvare et al. (1999) reported that Belaspidia is represented by two species. The Chalcidoidea Database mentions only nine species for Morocco (Noyes, 2019).

The aim of this study is to present for the first time a faunistic synthesis of Moroccan Chalcididae wasps with new records and information about their distribution.

Material and methods

The material studied here belongs to samples collected by the authors in the field together with revision of the material deposited in the MNHNM. Bibliographical data have also been added in this study.

Study area

The specimens were collected at three locations within the forest of Maâmora, Morocco (fig. 1): (1) Station 1: Taïcha, located near the forestry post Taïcha in the western Maâmora, canton A, commune Sidi Taïbi, 34° 13′ 29.5″ N, 6° 39′ 04.8″W, 36 m; (2) Station 2: Al Maha, located in a private reserve in South Eastern Maâmora, canton D, commune Aïn Johra, 34° 06′ 34.2″ N, 6° 19′ 04.0″ W, 175 m; and (3) Station 3: Aïn Assou, located near the forestry post Aïn Assou in the northern Maâmora, canton D, commune Kceïbia, 34° 12′ 07.9″ N, 6° 15′ 10.8″ W, 84 m. The climate of the Maâmora forest is in general thermo-Mediterranean, sub-humid at the first station and semi-arid at the other two stations (Métro and Sauvage, 1955). Geographical positions are given using coordinates in Merchich’s geographic reference system.

Fig. 1. Study sites and location of sampling trap in Maâmora forest with its different cantons (A, B, C, D and E). Fig. 1. Puntos de estudio y emplazamiento de las trampas de muestreo en el bosque de Maâmora con las distintas divisiones del mismo (A, B, C, D y E).

Sampling method

All samples were collected by K. Kissayi (first author), using one Malaise trap per station once a week over three periods: 26 April to 8 October 2012, 24 May to 23 September 2013 and 27 November 2013 to 14 June 2014. After sorting, specimens were preserved in 70 % alcohol prior to identification.

Identifications

Identifications were carried out by G. Delvare (CIRAD) and K. Kissayi using keys in Habu (1960, 1962, 1966), Bouček (1952, 1956, 1988), Bouček and Sedivy (1954), Ferrière and Kerrich (1958), Bouček and Narendran (1981), Narendran (1984, 1986, 1989), Delvare (1988, 1992, 1993), Bouček and Delvare (1992), Bouček and Halstead (1997), Narendran and van Achterberg (2016) and Delvare (2017).

Abbreviations used in the text

CIRAD, Center for International Cooperation in Agronomic Research for Development, Montpellier, France.
MNHNM, National Museum of Natural History, Scientific Institute, Rabat, Morocco.
MNHN, National Museum of Natural History, Paris, France.
ITSHM, Institute of Specialized Technicians in Horticulture, Meknes, Morocco.
NHM, Natural History Museum Bern, Switzerland.
GDPC, Gérard Delvare personal collection.
KKPC, Khadija Kissayi personal collection.

Results

A total of four subfamilies, 14 genera and 26 species of Chalcididae were recorded from Morocco: Chalcidinae (7 species), Cratocentrinae (1 species), Dirhininae (2 species) and Haltichellinae (16 species) (see dataset published in GBIF, Doi: 10.15470/nochzr).

Annotated list of Chalcididae species

The list of Moroccan Chalcididae shown below is arranged by subfamilies, tribes, genera and species in alphabetical order. Under distribution in Morocco we include localities and dates of capture (Morocco’ indicates that a more precise locality is unavailable). Remarks. The genus and/or species mentioned for the first times from Morocco are indicated by *.

1. New records from Morocco

Subfamily Haltichellinae Ashmead, 1904
Tribe Haltichellini Ashmead, 1904
Hockeria Walker, 1834

*Hockeria bifasciata Walker, 1834 (fig. 2)
Distribution in Morocco: Fir forest of the National Park of Talassemtane.
Examined material: Maâmora Forest, canton A, commune Sidi Taïbi, Taïcha, 1♀, 2 VII 2013, canton D, commune Aïn Johra, Al Maha, 1♀, 13 VIII 2013, K. Kissayi leg. (KKCP).
Remarks: this species has a Palaearctic and Indian distribution and is identified for the first time from Morocco. It was previously cited from North Africa from Algeria by Lucas (1849).

Fig. 2. Hockeria bifasciata Walker, 1834, ♂, habitus side view. (Photo by K. Kissayi.) Fig. 2.Hockeria bifasciata Walker, 1834, ♂, habitus, vista lateral. (Fotografía de K. Kissayi.)

*Hockeria mengenillarum (Silvestri, 1943)
Examined material: Maâmora Forest, canton D, commune Aïn Johra, Al Maha, 1♀, 13 VIII 2013, K. Kissayi leg. (GDCP).
Remarks: Hitherto only known from Italy (Silvestri, 1943) and Turkey (Öncüer, 1991). New from North Africa (Morocco).

Tribe Hybothoracini Bouček, 1952
*Proconura Dodd, 1915

*Proconura decipiens Masi, 1929 (fig. 3)
Examined material: Maâmora forest, canton D, commune Kceïbia, Aïn Assou, 1♂, 26 III 2013; 1♀, 1♂, 02 VIII 2013; 1♀, 14 VIII 2012; 1♀, 29 VIII 2012, K. Kissayi leg. (KKCP).
Remarks: new record from North Africa (Morocco).

Fig. 3. Proconura decipiens (Masi, 1929), ♀, habitus side view. (Photo by C. Locatelli.) Fig. 3. Proconura decipiens (Masi, 1929), ♀, habitus vista lateral. (Fotografía de C. Locatelli.)

2. Species previously known in Morocco

There are species already mentioned in the literature but not found in the present study, and also species deposited in museums in addition to those already cited in the catalogue of Noyes (2019).

Subfamily Chalcidinae Latreille, 1817
Tribe Brachymeriini Mani, 1938
Brachymeria Westwood, 1829

Brachymeria minuta (Linnaeus, 1767)
Examined material: Gardens of ITSHM, 4♀♀, 2♂♂, 16 XII 2015, K. Kissayi leg.
Remarks: species previously mentioned in Morocco by Jourdan and Rungs (1934: 210) in Rabat and in Tangier (MNHNM, Rabat) and this record is not included in Noyes (2019).

Brachymeria rugulosa Förster, 1859
Remarks: species previously mentioned in Morocco by Fabre and Mouna (1983: 4) in Ras-al-Ma, Azrou, Aïn Kahla.

Brachymeria secundaria (Ruschka, 1922)
Remarks: species previously mentioned in Morocco by Villemant and Fraval (1993: 93) in Maâmora forest and this record is not included in Noyes (2019).

Brachymeria tibialis (Walker, 1834)
Examined material: Tangier, 1919, J. De Gaulle leg. (MNHNM, under the name Chalcis minuta with an erroneous identification and the specimens under the name Chalcis intermedia); Maâmora forest, 7 specimens, 1924, A. Théry leg., L. Berland det. (MNHNM, under name Chalcis flavipes).
Remarks: this species was previously mentioned in Morocco from Maâmora forest (De Lépiney 1930: 87; Villemant and Fraval, 1993: 93; Villemant, 1989: 126); Daïet Achleff, Middle Atlas, about 1,800 m, 19 VII 1938 (Bleton and Fieuzet, 1939: 62), Middle Atlas (Fabre and Mouna, 2005: 93), Azrou Forests (De Lépiney and Mimeur, 1932: 72); Larache, Aïn felfel, Jbel Tazekka, Chaoun (Hérard and Fraval, 1980: 44), Fir forest of Talassemtane National Park (Benyahia, 2016: 167); Zaer forest (De Lépiney and Mimeur, 1932: 72), Mechrâ el Kettane, Bled Dendoun, Kenitra
(De Lépiney, 1927: 165).

Brachymeria vitripennis Förster, 1859
Remarks: this species was previously mentioned in Morocco by Anonymous (1938: 4) from Tadla, Doukkala, Loukkos, Moulouya, Gharb, Tetouan, Tangier, Haouz. This record is not included in Noyes (2019).

Tribe Chalcidini Fabricius, 1775
Chalcis Fabricius, 1787

Chalcis myrifex (Sulzer, 1876) (fig. 4)
Examined material: Tangier, ♀, 1914, Ernest André leg. (MNHNM, under the name Smicra myriflex).
Remarks: species mentioned in Europe, North Africa and North America. First record from Morocco; previously mentioned in North Africa from Tunisia (Bouček, 1952, 1977).

Fig. 4. Chalcis myrifex (Sulzer, 1876), ♀, habitus lateral view. (Photo by K. Kissayi.) Fig. 4. Chalcis myrifex (Sulzer, 1876), ♀, habitus vista lateral. (Fotografía de K. Kissayi.)

*Chalcis sispes (Linnaeus, 1760) (fig. 5)
Examined material: Tangier, 4 specimens, 1911, J. Vachal leg.; same locality, 7 specimens, 1915, Ernest André leg. (MNHN).
Remarks: species mentioned in Europe and North America. First record from Morocco; previously mentioned in North Africa from Tunisia (Bouček, 1952).

Fig. 5. Chalcis sispes (Linnaeus, 1760), habitus dorsal view. (Photo by K. Kissayi.) Fig. 5. Chalcis sispes (Linnaeus, 1760), habitus vista dorsal. (Fotografía de K. Kissayi.)

Subfamily Cratocentrinae Steffan, 1951

The subfamily Cratocentrinae (Hymenoptera, Chalcididae) was reviewed and reassessed by Heraty et al. (2013) and recently by Delvare (2017) and Abul-Sood et al. (2018).

Cratocentrus Cameron, 1907

Cratocentrus fastuosus (Masi, 1944)
Examined material: Rabat, 1♀, 22 VII 1938, Ch. Rungs leg., J. R. Steffan det.; Beni Amer, 32° 30′ N, 6° 42′ W, 1♂, 1♀, 17 V 1947, Ch. Rungs leg., J. R. Steffan det. (MNNH).
Remarks: species previously mentioned in Morocco from Taghazoute, 20 km N Agadir, Volubilis, Moulay Idriss (Abul-Sood et al., 2018: 500), Herting (1973: 31, 53) and by
Steffan (1959: 313) without locality; the examined material probably corresponds to Steffan records published in 1959.

Subfamily Dirhininae Ashmead, 1988
Tribe Dirhinini Ashmead, 1904
Dirhinus Dalman, 1818

Dirhinus (Dirhinus) aff. hesperidum (Rossi, 1790)
Remarks: this species was reported by Delvare and Copeland (2018) from Moulay Idriss, the ancient city of Volubilis, and 18 km N Souk-El Arbaa du Gharb, but it is not included in Noyes (2019).

Dirhinus himalayanus Westwood, 1836
Remarks: this species was reported by Blanchot (1992: 7) from Zagora, a large number in swine facility but it is not included in Noyes (2019).

Subfamily Haltichellinae Ashmead, 1904
Tribe Haltichellini Ashmead, 1904
Antrocephalus Kirby, 1883

Antrocephalus cameroni Delvare, 2017
Remarks: this species was reported by Delvare (2017: 257) from Oued Assaka, and Moulay Idriss, ancient city of Volibulis, GDPC (G. Delvare, pers. comm.).

Antrocephalus hypsopygiae Masi, 1928
Remarks: this species collected from Oued Cherrat, NHM (G. Delvare, pers. comm.); previously mentioned from Morocco by Lotfalizadeh et al. (2012: 79).

Belaspidia Masi, 1916

Belaspidia obscura Masi, 1916
Remarks: this species mentioned in Morocco from Essaouira-Agadir road, 1 km N Oued Ibouzoule (Delvare et al., 1999: 338); Tata Province, Tichekji; Province of Chtouka Aït Baha, Aït Baha (Stefanescu et al., 2012: 88, 91).

Belaspidia tussaci Delvare, 1999
Remarks: species described from Morocco material collected from Arbaoua (Delvare et al., 1999: 339).

Chirocera Latreille, 1825

Chirocera pectinicornis (Latreille, 1809)
Remarks: species known from Morocco (Steffan, 1957: 140) not included in Noyes (2019).

Haltichella Spinola, 1811

Haltichella rufipes (Olivier, 1791)
Remarks: species mentioned in Morocco from Rabat (Rungs, 1936: 18), not included in Noyes (2019).

Hockeria Walker, 1834

Hockeria singularis Bouček, 1952
Remarks: species know in Morocco locality not indicated (Steffan, 1953: 11).

Hockeria unicolor Walker, 1834
Examined material: Maâmora Forest, canton D, commune Aïn Johra, Al Maha, 1♀, 06 IX 2012, K. Kissayi leg.
Remarks: species previously mentioned in Morocco by (Bouček, 1952: 69).

Neochalcis Kirby, 1883

Neochalcis fertoni Kieffer, 18
Examined material: Maâmora forest, 1 specimen, 18 VI 1925 (MNHNM under the name Neochalis barbara).
Remarks: species previously mentioned in Morocco from Casablanca (Benoist, 1921: 119).

Tribe Hybothoracini Bouček, 1952
Bucekia Steffan, 1951

Bucekia differens (Bouček, 1949)
Remarks: species mentioned in Morocco (Falahatpisheh et al., 2018: 258) not included in Noyes (2019).

Lasiochalcidia Masi, 1929

Lasiochalcidia pubescens (Klug, 1834)
Remarks: species know in Egypt, Iran and Morocco; in Morocco from Tangier (Bouček, 1952: 104, Steffan, 1951a: 5, 1951b: 77) but these records are not included in Noyes (2019).

Lasiochalcidia rubripes (Kieffer, 1899)
Remarks: species know from Italy (Corsica only), Hungary and Morocco; in Morocco it was reported by Falahatpisheh et al. (2018: 260) but this record is not included in Noyes (2019).

Psilochalcis Kieffer, 1905

Psilochalcis benoisti (Steffan, 1948)
Remarks: species distributed in Europe and North Africa. In Morocco it was reported by Steffan (1948: 121, 1951a: 61; Bouček, 1956: 244).

Discussion

This study presents data, for the first time, on 26 species of Chalcididae (Hymenoptera, Chalcidoidea) belonging to 14 genera and 4 subfamilies in Morocco. Of these, Hockeria bifasciata Walker, 1834, H. mengenillarum (Silvestri, 1943), and the genus Proconura represented by P. decipiens (Masi, 1929), found in cork oak forest of Maâmora, are new records from Morocco. These species are possibly economically relevant because they are parasitoids of various pests. Hockeria bifasciata is reported as a primary parasitoid of some families of Lepidoptera, and H. mengenillarum is also a primary parasitoid of Strepsiptera (Bouček, 1952). Hosts quoted for P. nigripes require confirmation because P. nigripes and P. decipiens are two distinct species that have been erroneously synonymized (G. Delvare, pers. com.). Moreover, according to the literature, four genera (Bucekia Steffan, Chirocera Latreille, Dirhinus Dalman and Haltichella Spinola) and 12 species are here reported for the first time in Morocco. Special mention has been made of the genus Chalcis Fabricius never previously reported in Morocco but represented by two species. Consequently, this study has enriched the Moroccan Chalcididae fauna described given by Noyes (2019) by 17 species. Most new records were made by visiting MNHNM (Rabat) and MNHN (Paris) and those newly identified during this work are summarized in table 1 and the dataset published in GBIF (Doi: 10.15470/nochzr). From among all the Moroccan Chalcididae, one species, Belaspidia obscura Masi, can be considered endemic to Morocco.

Table 1. The newest genera and species of Chalcididae recorded in Morocco during this study: * genera and species not included in Noyes (2019); ** genera and/or species newly identified. Tabla 1. Nuevos géneros y especies de Chalcididae registrados en Marruecos durante este estudio: * géneros y especies no incluidos en Noyes (2019); ** generos y/o especies identificados por primera vez.

Knowledge of Chalcididae, particularly in the North African region, is still very incomplete and has several shortcomings. Comparison of Moroccan Chalcididae with the fauna of other North African countries reveals that Morocco currently has greater species richness of this family (24 species) than its neighbouring countries. In Algeria, Egypt, Tunisia, Libya and Mauritania 21, 20, 15, 13 and 1 species are cited, respectively. Nevertheless, these data do not reflect the reality according to Noyes (2019); because unfortunately no recent revisions are available and no recent studies have been conducted in all countries in the region.

Finally, despite the economic relevance of various species of this family and of other families of Chalcidoidea, they have not been the focus of studies in Moroccan until our research project that began in 2012 in the cork oak forest of Maâmora (Kissayi and Benhalima, 2016, 2017a, 2017b, 2018; Delvare et al., 2017; Kissayi et al., 2017a, 2017b). Located in the western Palaearctic region, between the Mediterranean Sea and the Atlantic Ocean, in addition to being one of the 34 hotspots of biodiversity, Morocco most probably hosts a large entomofauna. Faunistic studies and further investigations on chalcids wasps are thus strongly recommended in other parts of the country and indeed throughout the entire North African region.

Acknowledgements

We would like to thank Dr Gérard Delvare (CIRAD, Montpellier, France) for identification and validation of the species. Our sincere thanks go to Mrs. Claire Villemant, responsible for the collection of Hymenoptera at the Museum of Natural History in Paris (France), and Wouter Dekoninck curator of collection, Museum of the Royal Institute of Natural Sciences in Brussels (Belgium) for his great hospitality and for facilitating Mrs. Khadija Kissayi’s consultations on Hymenoptera in the museums, and to Mrs. Camille Locatelli, responsible for digitization of the collections, Royal Belgian Institute of Natural Sciences, for making beautiful photos.

References

Anonymous, 1938. La cassidae de la Betterave Cassida vittata Wildl et Olonga Illig. Service de la défense des végétaux Ministère de l’Agriculture, Maroc: 1–6.

Abul-Sood, M. I., Gadallah, N. S., Hossni, M. T., Delvare, G., 2018. The subfamily Cratocentrinae (Hymenoptera: Chalcididae): reappraisal of their morphological characters and review of the West Palaearctic species, with the description of two new species. Zootaxa, 4377(4): 490–516.

Benoist, R., 1921. Description d’une nouvelle espèce du genre Euchalcis (Hym. Chalcididae). Bulletin de la Société entomologique de France, 8: 118–120.

Benyahia, Y., 2016. Inventaire de la biodiversité entomologique (Coléoptères et Hyménoptères) pour une gestion patrimoniale de la sapinière du Parc National de Talassemtane (Rif, Maroc). PhD thesis, Université Hassan II.

Blanchot, Ph., 1992. Nouveau répertoire bibliographique et nouvelles données bio-écologiques sur les parasites de Musca domestica Linné (Diptera: Muscidae). EPHE, Biologie et Evolution des Insectes, 5: 1–54.

Bléton, C. A., Fieuzet, L., 1939. Notes sur quelques auxiliaires observés dans la région de Fès. Bulletin de la Société des Sciences Naturelles du Maroc, 19: 57–65.

Bouček, Z., 1977. A faunistic review of the Yugoslavian Chalcidoidea (Parasitic: Hymenoptera). Acta Entomologica Jugoslavica, 13 (Supplement): 1–145.

– 1952. The first revision of the European species of the family Chalcididae (Hymenoptera). Sborník Entomologického Oddeleni Národního Musea v Praze, 27 (Supplement 1): 1–108.

– 1956. A contribution to the knowledge of the Chalcididae, Leucospidae and Eucharitidae (Hymenoptera, Chalcidoidea) of the Near East. Bulletin of the Research Council of Israel, 5B: 227–259.

– 1988. Australasian Chalcidoidea (Hymenoptera). A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International, Wallingford, United Kingdom.

Bouček, Z., Delvare, G., 1992. On the New World Chalcididae (Hymenoptera). Memoirs of the American Entomological Institute, 53: 1–466.

Bouček, Z., Halstead, J. A., 1997. Chapter 6. Chalcididae. In: Annotated Keys to the Genera of Nearctic Chalcidoidea (Hymenoptera): 151–164 (G. A. P. Gibson, J. T. Huber, J. B. Woolley, Eds.). National Research Council Research Press, Ottawa, Ontario, Canada.

Bouček, Z., Narendran, T. C., 1981. Indian chalcid wasps (Hymenoptera) of the genus Dirhinus parasitic on synanthropic and other Diptera. Systematic Entomology, 6: 229–251.

Bouček, Z., Sedivy, J., 1954. Die Hymenopteren-Parasiten von Hyphantria cunea Drury in der Tschechoslowakei. Folia Zoologica Entomologica, 3: 169–189.

Chakir, S., Fraval, A., 1985. Native and introduced enemies of Lymantria dispar (Lep., Lymantriidae) in the Mamora forest, Morocco: study along a transect, 1976-1982. Actes de l’Institut Agronomique et Vétérinaire Hassan II, Maroc, 5(1-2): 27–36.

Dajoz, R., 2010. Dictionnaire d’Entomologie. Lavoisier/tec et doc, France.

De Lépiney, J., 1927. Les insectes nuisibles du Chêne-liège dans la forêt de la Mamora (Maroc). Annales Epiphyties, 13: 145-174.

– 1930. Contribution à l’étude du complexe biologique de Lymantria dispar L. Mémoire de la Société des Sciences naturelles du Maroc, 23: 1–100.

De Lépiney, J., Mimeur, J.-M., 1932. Notes d’entomologie agricole et forestière du Maroc. Mémoires de la Société des sciences naturelles du Maroc, 31: 1–195.

De Lombaerde, J., Pauly, A., Vago, J. L., 2002. Les Chalcididae de Belgique et du Nord de la France (Hymenoptera Chalcidoidea). Notes fauniques de Gembloux, 49: 95–101.

Delvare, G., 1988. Some important morphological features of the Chalcidini (Hymenoptera: Chalcididae) and their implications in the classification of the tribe. In: Advances in Parasitic Hymenoptera Research: 25–64 (V. K. Gupta, Ed.). Leiden, The Netherlands.

– 1992. Les Chalcididae (Hymenoptera) d’importance économique dans les palmeraies d’Amérique tropicale. Bulletin de la Société entomologique de France, 97(4): 349–372.

– 1993. A reclassification of the Chalcidini (Hym., Chalcididae) with a check list of the New World species. Memoirs of the American Entomological Institute, 53: 119–457.

– 1995. The Family Chalcididae. In: The Hymenoptera of Costa Rica: 289–298 (P. Hanson, I. Gauld, Eds.). Oxford Science Publications, Oxford.

– 2017. Order Hymenoptera, family Chalcididae. Arthropod fauna of the UAE, 6: 225–274.

Delvare, G., Copeland, R. S., 2018. Four-horned wasps, description of some remarkable Dirhinus (Hymenoptera, Chalcididae) from Kenya, with a discussion of their taxonomic placement. Zootaxa, 4374(3): 301–349.

Delvare, G., Kissayi, K., Benhalima, S., 2017. Revue des espèces ouest-paléarctiques du genre Tineophoctonus (Hymenoptera: Encyrtidae: Encyrtinae). Bulletin de la Société Entomologique de France, 122 (2): 191–198.

Delvare, G., Pujade-Villar, J., Rasplus, J. Y., 1999. Révision des Blaspidia Ouest-Paléarctique (Hymenoptera: Chalcidoidea). Annales de la Société Entomologique de France (n.s.), 35(3-4): 329–345.

Fabre, J. P., Mouna, M., 1983. Sur la présence au Maroc d’une tordeuse Acleris undulala Walsingham (Lep. Tortricidae) grave ravageur des cèdres: Cedrus atlantica Mannetti, Cedrus libani Barrel. Extrait du procès-verbal de la séance du 11 mai 1983. Académie d’Agriculture de France, 642–647.

– 2005. Pest insects of cedars: Cedrus atlantica Manetti, C. Libani A. Richard and C. Brevifolia Henry in Mediterranean area. In: Entomological Research in Mediterranean Forest Ecosystems: 89–104 (F. Lieutier, D. Ghaioule, Eds.). INRA Editions, Paris.

Falahatpisheh, A., Fallahzageh, M., Dousti, A., Delvare, G., 2018. Review of Iranian Chalcididae (Hymenoptera, Chalcidoidea) with nomenclatural notes. Zootaxa, 4394(2): 251–269.

Ferrer, E. P., 2010. Composición de la familia Chalcididae (Hymenoptera: Chalcidoidea) en Cuba. Cocuyo, 18: 48–54.

Ferrière, Ch., Kerrich, G. J., 1958. Agaonidae, Leucospidae, Chalcididae, Eucharitidae, Perilampidae, Cleonymidae and Thysanidae. Handbook for the identification of British Insects, 8(2)(a): 1–40.

Habu, A., 1960. A revision of the Chalcididae (Hymenoptera) of Japan with descriptions of sixteen new species. Bulletin the National Institute of agricultural sciences, series C (Phytopathology and Entomology), 11: 132–363.

– 1962. Chalcididae, Leucospididae and Podagrionidae (Insecta: Hymenoptera). Chiyode Publishing & Printing Co., Ltd., Tokyo, Japan.

– 1966. Descriptions of some Brachymeria species of Japan (Hymenoptera: Chalcididae). Kontyu, 34(1): 22–28.

Hérard, F., Fraval, A. 1980. La répartition et les ennemis naturels de Lymantria dispar (L.) (Lep. Lymantriidae) au Maroc; 1973-1975. Acta oecologica, Oecologia applicata, 1(1): 35–48.

Heraty, J. M., Burks, R. A., Cruaud, A., Gibson, G. A. P., Liljeblad, J., Munro, J., Rasplus, J. -Y., Delvare, G., Janšta, P., Gumovsky, A., Huber, J., Woolley, J. B., Krogmann, L., Heydon, S., Polaszek, A., Schmidt, S., Darling, D. C., Gates, M. W., Mottern, J., Murray, E., Dal Molin, A., Triapitsyn, S. V., Baur, H., Pinto, J. D., van Noort, S., George J., Yoder M., 2013. A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera). Cladistics, 29(5): 466–542.

Herting, B., 1973. Coleoptera to Strepsiptera. A catalogue of parasites and predators of terrestrial arthropods. Section A. Host or Prey/Enemy.3. Commonwealth Agricultural Bureaux, Institute of Biological Control.

– 1978. Neuroptera, Diptera, Siphonaptera. A catalogue of parasites and predators of terrestrial arthropods. Section A. Host or Prey/Enemy. Commonwealth Agricultural Bureaux, Commonwealth Institute of Biological Control.

Jourdan, M. L., Rungs, Ch., 1934. Observations sur quelques Hyménoptères du Maroc. Bulletin de la Société des Sciences Naturelles du Maroc, 14: 204–213.

Kissayi, K., Benhalima, S., 2016. Contribution à la connaissance des Encyrtidae (Hymenoptera: Chalcidoidea) du Maroc, nouvelles données et comparaison avec la faune d’Afrique du Nord. Annales de la Société Entomologique de France, 52(6): 343–353.

– 2017a. First focused survey of Eupelmidae (Hymenoptera: Chalcidoidea) in Morocco with three new records. Annales de la Société Entomologique de France, 53(3): 211–218.

– 2017b. First report of the family of Signiphoridae Howard, 1894 (Hymenoptera: Chalcidoidea) in Morocco. Revue Marocaine Protection des Plantes, 11: 29–31.

– 2018. Ophelimus maskelli (Ashmead, 1900) et son parasitoïde Closterocerus chamaeleon Girault, 1922 (Hymenoptera: Eulophidae) au Maroc. Bulletin OEPP/EPPO Bulletin, 48(2): 293–299.

Kissayi, K., Benhalima, S., Douaik, A., 2017a. Diversité des Encyrtinae (Hymenoptera: Chalcidoidea) collectés par pièges Malaise au sein de la forêt de la Maâmora. 8ème Meeting IOLB-WPRS Group Integrated Protection in Oak Forest. IOBC-WPRS Bulletin, 127: 45–49.

Kissayi, K., Benhalima, S., Smaili, M. C., 2017b. New records of Aphelinidae (Hymenptera: Chalcidoidea) from Morocco and comparison with North Africa region fauna. Journal of Entomology and Nematology, 9(7): 55– 62.

Lotfalizadeh, H., Ebrahimi, E., Delvare, G., 2012. A contribution to the knowledge of family Chalcididae (Hymenoptera: Chalcidoidea) in Iran. Journal of Entomological Society of Iran, 31(2): 67–100.

Lucas, P. H., 1849. Histoire naturelle des animaux articulés. Insectes (suite). Exploration Scientifique de l’Algérie pendant les années 1840, 1841, 1842. Sciences Physiques, Zoologie, 3, Paris.

Métro, A., Sauvage, C., 1955. Flore des végétaux ligneux de la Mamora. La nature au Maroc. La Société des sciences naturelles et physiques du Maroc.

Narendran, T. C., 1984. Key to Indian genera of the family Chalcididae (Hymenoptera: Chalcidoidea). Entomophaga, 29(4):431–438.

– 1986. Family Chalcididae. In: The Chalcidoidea (Insecta: Hymenoptera) of India and the adjacent countries: 174-178 (B. R. Subba Rao & M. Hayat, Eds). Oriental Insects, Florida.

– 1989. Oriental Chalcididae (Hymenoptera: Chalcidoidea). Department of Zoology, University of Calicut, Kerala, India.

Narendran, T. C., van Achterberg, C., 2016. Revision of the family Chalcididae (Hymenoptera, Chalcidoidea) from Vietnam, with the description of 13 new species. ZooKeys, 576: 1–202.

Noyes, J. S., 2019. Universal Chalcidoidea Database [online]. <http://www.nhm.ac.uk/our-science/data/chalcidoids/database/listChalcids.dsml> [Accessed on 2 January 2019].

Öncüer, C., 1991. A catalogue of the parasites and predators of insect pests of Turkey. no. 550, Ege Universitesi Ziraat Facultesi; Yayinlari.

Rungs, Ch., 1936. Observations sur quelques Hyménoptères du Maroc. Bulletin de la Société des Sciences Naturelles du Maroc, 16: 15–31.

Silvestri, F., 1943. Studi sugli Strepsiptera Insecta. III Descrizione e biologia di specie italiane di Mengenilla. Bollettino del Laboratorio di Zoologia Generale e Agraria della Facoltà Agraria in Portici, 32: 197–282.

Steffan, J. R., 1948. Deux nouveaux genres d’Haltichellinae (Hym. Chalcididae). Bulletin de la Société Entomologique de France, 5: 118–123.

– 1951a. Les espèces françaises d’Haltichellinae (Hyménoptères Chalcididae). Feuille des Naturalistes, 6(1-2): 1–7.

– 1951b. Contribution à l’étude des Haltichellinae « groupe Euchalcidia » (Hymenopt. Chalcididae) conservés au Muséum national d’Histoire naturelle de Paris. Mémoires du Muséum National d’Histoire Naturelle, Série Zoologie, 4(2): 51 86

– 1953 Les espèces françaises de Haltichellinae (Hyménoptères Chalcididae) (Suite). Cahiers des Naturalistes, Bulletin des Naturalistes Parisiens (N.S.), 8: 7–12.

– 1957. Révision du genre Chirocera Latr., Tanyotorthus Stef. et Tanycoryphus Cam. (Hym. Chalcididae). Annales de la Société Entomologique de France, 126: 139-158.

– 1959. Révision de la tribu des Cratocentrini (Hym., Chalcididae). Acta Entolologica Musei Natlalis Pragae, 33(549): 287–325.

Stefanescu, C., Askew, R. R., Corbera, J., Shaw, M., 2012. Parasitism and migration in southern Palaearctic populations of the painted lady butterfly, Vanessa cardui (Lepidoptera: Nymphalidae). European Journal of Entomologie, 109: 85–94.

Villemant, C., 1989. Ennemis des chenilles et chrysalides. In: Lymantria dispar: 125–143 (A. Fraval, Ed.). Actes Editions, Rabat.

Villemant, C., Fraval, A., 1993. La faune entomologique du chêne-liège en forêt de la Mamora (Maroc). Ecologia Méditerranea, 19: 89–98.