Contribution to the knowledge of Afrotropical Dryinidae, Embolemidae and Sclerogibbidae (Hymenoptera), with description of new species from Central African Republic and Uganda

Abstract An updated checklist of Dryinidae, Embolemidae and Sclerogibbidae from Central African Republic and Uganda is presented. The following new species of Dryinidae are described: from Central African Republic: Anteon dzanganum sp. n. (Anteoninae); from Uganda: Anteon granulatum sp. n., Anteon kibalense sp. n., Anteon makererense sp. n., Anteon mubfs sp. n. (Anteoninae); Bocchus kibalensis sp. n. (Bocchinae); Dryinus kibalus sp. n. (Dryininae); Gonatopus kanyawarus sp. n. (Gonatopodinae). The following species have been recorded for the first time from Central African Republic: Embolemidae: Ampulicomorpha madecassa Olmi, 1999a; Embolemus capensis Olmi, 1997; Dryinidae: Aphelopus mediocarinatus (Benoit, 1951d), Aphelopus testaceus Olmi, 1991, Aphelopus wittei Benoit, 1951c (Aphelopinae); Anteon cautum Olmi, 1994a, Anteon evertsi Olmi, 1989, Anteon gutturnium (Benoit, 1951b), Anteon inflatrix Benoit, 1951b, Anteon kivuanum (Benoit, 1951c), Anteon semajanna Olmi, Copeland & Guglielmino, 2015, Anteon zairense Benoit, 1951d (Anteoninae); Pseudodryinus townesi (Olmi, 1984) (Dryininae); Echthrodelphax tauricus Ponomarenko, 1970, Gonatopus camerounensis Olmi, 2011, Gonatopus kolyadai Olmi, 2007b, Neodryinus antiquus Benoit, 1954, Neodryinus tussaci Olmi, 2004b (Gonatopodinae); Sclerogibbidae: Probethylus callani Richards, 1939b; Sclerogibba algerica Benoit, 1963, Sclerogibba rapax Olmi, 2005a. The following species have been recorded for the first time from Uganda: Embolemidae: Ampulicomorpha magna Olmi, 1996; Dryinidae: Anteon cautum Olmi, 1994a, Anteon fisheri Olmi, 2003, Anteon hoyoi Olmi, 1984, Anteon kivuanum (Benoit, 1951c), Anteon townesi Olmi, 1984, Anteon zairense Benoit, 1951d (Anteoninae); Bocchus bini Olmi, 1984 (Bocchinae); Dryinus saussurei (Ceballos, 1936) (Dryininae); Echthrodelphax migratorius Benoit, 1954, Neodryinus tussaci Olmi, 2004b (Gonatopodinae). The following further species has been recorded for the first time from Mali: Sclerogibba algerica Benoit, 1963 (Sclerogibbidae); from Ivory Coast: Adryinus oweni Olmi, 1984 (Gonatopodinae); from Cameroon and South Africa: Gonatopus operosus Olmi, 1993 (Gonatopodinae); from Democratic Republic of the Congo and Zambia: Neodryinus antiquus Benoit, 1954 (Gonatopodinae); from South Africa: Anteon striatum Olmi, 2005b (Anteoninae). Including the above new records, 23 species of Dryinidae (previously six), two species of Embolemidae (previously none) and three species of Sclerogibbidae (previously two) are now known from Central African Republic. For Uganda, 39 species of Dryinidae (previously 23), one species of Embolemidae (previously none) and four species of Sclerogibbidae (previously four) are now known. Additional new faunistic records are provided for Cameroon, Democratic Republic of the Congo, Ivory Coast, Mali, South Africa and Zambia.

Afrotropical species of the above three families are poorly known. In recent years research investigation has been intensified in Burundi, Kenya, Madagascar, Mozambique and South Africa (mainly thanks to the efforts of Robert S. Copeland in Burundi and Kenya, Brian L. Fisher in Madagascar, Massimo Olmi in Mozambique and Simon van Noort in South Africa). The above inventory surveys resulted in the descriptions of many new species, and contributed to the publication of updated checklists and Olmi 2007aCopeland 2011 andOlmi et al. 2015 on Burundi andKenya;Olmi et al. 2012 on Mozambique;Olmi 2006Olmi , 2007bOlmi , 2009 on South Africa). However, in spite of the above efforts, the fauna of dryinids, embolemids and sclerogibbids of many countries remains almost unknown.
In addition one of the authors (Simon van Noort) extended his research to two of the lesser known Afrotropical countries, Central African Republic and Uganda, where the number of recorded species was particularly low (in the Central Africa Republic, 6 species of Dryinidae, two species of Sclerogibbidae, no species of Embolemidae; in Uganda, 23 species of Dryinidae, one species of Embolemidae and four species of Sclerogibbidae). The subsequent study of the collected material has resulted in the discovery of eight new species described herein and, along with new distributional data for previously described species of these three families, has provided the opportunity to update the checklist of Dryinids, Embolemids and Sclerogibbids known from Central African Republic and Uganda.

Material and methods
Species descriptions follow the terminology used by Olmi (1984Olmi ( , 1994cOlmi ( , 1999b, Xu et al. (2013), Olmi and Virla (2014) and Olmi and Xu (2015). The measurements reported are relative, except for the total length (head to abdominal tip, without the antennae), which is expressed in millimetres. In the figures of male genitalia the right half is not included. The following abbreviations are used in the descriptions: POL distance between the inner edges of the two lateral ocelli; OL distance between the inner edges of a lateral ocellus and the median ocellus; OOL distance from the outer edge of a lateral ocellus to the compound eye; OPL distance from the posterior edge of a lateral ocellus to the occipital carina; TL distance from the posterior edge of the eye to the occipital carina.

Surveyed areas
The Dzanga-Sangha protected area was surveyed in Central African Republic. This area lies north of the equator and is located in the southwest triangle of the country (Sangha-Mbaéré Prefecture) bordered by Cameroon and the Republic of Congo. The Dzanga-Sangha protected area includes the Dzanga-Ndoki National Park (1220 km 2 ), and the Dzanga-Sangha Dense Forest Special Reserve (3359 km 2 ). The latter is a multiple use zone where logging, traditional hunting, safari hunting and extraction of plants are still allowed under controlled conditions. Annual rainfall is about 1500 mm, with average temperatures ranging between 25°and 29°Celsius. There are two peaks to the rainy season with highest precipitation occurring during the "long rains" from September to November and a second peak during the "short rains" in May and June (Carroll 1997). The eco-region is a part of the Guineo-Congolian lowland rain forest within the Guineo-Congolian regional centre of endemism (White 1983), characterized by the following species: Entandrophragma congoense (Meliaceae); Pentaclethera eetveldeana (Mimoseae); Pericopsis elata (Fabaceae); and Gilbertiodendron dewevrei (Fabaceae). The canopy can reach a height of 60 m. The understory is composed of shrubs, lianas and herbs. Harris (2002) recorded 1090 species of vascular plants in the reserve.
Three separate sites within the forest were sampled (named Camps 1-3). Camp 1 was situated at a marsh clearing, Mabéa Bai, in lowland rainforest 21.4km 53°NE Bayanga, 3°02.01'N, 16°24.57'E, 510m, in the Dzanga-Ndoki National. The vegetation in the Bai marsh clearing is dominated by herbaceous plants including abundant sedges (Cyperaceae) and grasses (Graminae). Characteristic trees of the forest margin include Lophira alata and Berlinia grandiflora. Camp 2 was situated in lowland rainforest on the banks of the Sangha River in the Dzanga-Sangha Dense Forest Special Reserve,12.7km 326°NW Bayanga,3°00.27'N,16°11.55'E,420m. The river is about 500 meters wide at this point and ranges from a depth of 20 cm at the end of the dry season (around March) to as much as 5 meters during the height of the rainy season in September and October when the forests adjacent to the banks are flooded. This seasonally flooded forest has a complex architecture with 15-25 m trees forming a canopy with occasional emergent trees to 40 m with gaps less common; the understorey consists of small trees (5-10 m) with herbs and lianas common, whereas shrubs are almost absent (Harris 2002). Camp 3 was situated in lowland rainforest about 1 km from the banks of the Sangha River in the 38.6km 173°S Lidjombo,2°21.60'N,16°09.20'E,350m. Sampling was conducted in the seasonally inundated riparian forest (see above for species composition) and in mixed species terra firma forest above the flood plain. Details of the sampled habitats (including photographs of the sampling sites) and sampling effort conducted during the WWF expedition to Central African Republic are presented in Azevedo et al. 2015. In Uganda surveys were undertaken within the vicinity of the Makerere University Biological Field Station (MUBFS) (0°33.798'N, 30°21.365'E, 1500 m) in the Kanyawara area of Kibale National Park where 12 main survey sites were sampled. The park is located in western Uganda, approximately 24 km from the eastern edge of the Ruwenzori Mountains and encompasses an area of 766 km 2 and an altitude varying between 1590 m in the north and 1100 m in the south (Chapman et al. 1997). Kibale forest is transitional between lowland rain forest and montane rain forest and is classified as a mid-altitude, moist, evergreen forest with a canopy height typically between 20 and 30 m, but with some trees exceeding 55 m (Chapman et al. 1997, Skorupa 1988, Struhsaker 1997, Wrangham et al. 1994). The park is dominated by moist evergreen forest (57%), with secondary regenerating forest (19%), woodland (4%), grassland (15%), lakes and wetlands (2%), and exotic tree plantations (1%) comprising the remainder (Chapman et al. 1997). Rainfall is bimodal with highest precipitation concentrated during March to May and September to November. Mean annual rainfall totals 1734 mm; mean maximum temperature is 23.7°C; mean minimum temperature is 15.5°C (Rhode et al. 2006). Chapman et al. (1997), Skorupa (1988), andStruhsaker (1997) provide further details of the Kanyawara area.

Sampling methods
The Malaise traps were constructed to the specifications of the Townes design (Townes 1972), and made with a fine-meshed netting (grid size of 0.2 mm), with black walls and a white roof. Yellow pan traps comprised plastic bowls of 165mm diameter x 40 mm depth, which were placed on the forest floor along a linear transect. Sweeping: The sweep net used for sampling was based on the design of Noyes (1982), with an opening area of c. 1300 cm 2 , and a collecting bag constructed from fine-meshed netting with a grid size of 0.2 mm. The yellow pan trap collection method is indicated by the following abbreviation: YPT.

Aphelopus wittei
Distribution. Afrotropical, recorded from almost all sub-saharian countries, from Senegal to Somalia, in addition to Madagascar and Yemen (Olmi and Copeland 2011;Olmi et al. 2015;Olmi and van Harten 2006). Newly recorded from Central African Republic here.

Subfamily Anteoninae Perkins, 1912
Genus  (Olmi 1994a(Olmi , 2009. Newly recorded from Central African Republic and Uganda here. Diagnosis. Female fully winged; head and mesosoma black, except mandible testaceous; head and scutum granulated and reticulate rugose; posterior surface of propodeum reticulate rugose, without longitudinal keels; forewing hyaline, without dark transverse bands or spots, with distal part of stigmal vein less than 0.5 as long as proximal part; segment 4 of protarsus slightly shorter than basal part of segment 5; segment 5 of protarsus (Figs 1A, 4D) with inner side curvilinear, with distinct apical region, basal part shorter than distal part, distal region very short and distal lamellae located near medial lamellae.

Male. Unknown.
Differential diagnosis. Because of the above diagnosis, the new species is similar to Anteon canabense (Benoit, 1951b). The main differences regard the sculpture of the posterior surface of the propodeum and the colour (posterior surface of propodeum granulated, except few irregular keels near margins; head and mesosoma testaceous-reddish, occasionally partly darkened, in A. canabense; posterior surface of propodeum reticulate rugose; head and mesosoma black, except mandible testaceous, in A. dzanganum).
Etymology. The species is named after the type locality, Dzanga-Ndoki National Park.

Hosts. Unknown.
Distribution. Central African Republic.  (Olmi 1989;Olmi et al. 2015). Newly recorded from Central African Republic here. Diagnosis. Male with clypeus not sculptured by longitudinal and subparallel keels; scutum completely granulated and slightly reticulate rugose; posterior surface of propodeum without longitudinal keels; paramere (Figs 1D, 5F) with small distal inner rounded process and proximal membranous process not provided with mosaic sculpture and sensorial setae; distal inner process of paramere with distal apex situated quite close to distal apex of paramere (Figs 1D, 5F).
Female. Unknown. Differential diagnosis. Because of the above diagnosis, the new species is similar to Anteon cautum Olmi, 1994a, A. emeritum Olmi, 1984, and A. whartoni Olmi, in Olmi & Copeland, 2011. The main difference regards the scutum sculpture: completely reticulate rugose and granulated in A. granulatum; not completely reticulate rugose, at most with small surface near anterior margin rugose in the other three species.
Etymology. The species is named after the head completely granulated.
Hosts. Unknown. Distribution. Afrotropical, recorded from almost all sub-saharian countries, from Gambia to Somalia (Olmi 1984(Olmi , 2006, in addition to Madagascar  Diagnosis. Female fully winged; head and mesosoma black, except mandible brown; head completely reticulate rugose; posterior surface of propodeum reticulate rugose, without longitudinal keels, with areolae about as large as those of dorsal surface; forewing with two dark transverse bands, with distal part of stigmal vein less than 0.5 as long as proximal part; segment 4 of protarsus slightly shorter than basal part of segment 5; segment 5 of protarsus (Figs 1B, 6F) with inner side curvilinear, with distinct apical region, with basal part slightly shorter than distal part.
Male. Unknown. Differential diagnosis. Because of the above diagnosis, the new species is similar to Anteon zimbabwense Olmi, 2005b. The main difference regards the segment 5 of the protarsus: with smaller lamellae and distal part slender in A. zimbabwense (Fig. 1F); with lamellae longer and distal part less slender in A. kibalense (Fig. 1B, 6F).
Etymology. The species is named after the type locality, Kibale National Park.  Olmi 1984Olmi , 2006Olmi and van Harten 2006;Olmi et al. 2015). Newly recorded from Central African Republic and Uganda here. Diagnosis. Male with head reticulate rugose and granulated; scutum with anterior third reticulate rugose and remaining surface sculptured by many longitudinal subparallel irregular keels; posterior surface of propodeum not provided with longitudinal keels; propodeum with strong transverse keel between dorsal and posterior surface; paramere (Figs 1E, 7F) without distal inner pointed or rounded process, slightly shorter than penis.
Female. Unknown. Differential diagnosis. Because of the above diagnosis, the new species is similar to Anteon reunionense Olmi, 1987. The main difference regards the sculpture of the scutum: with anterior half reticulate rugose and remaining surface unsculptured, or slightly granulated in A. reunionense; with anterior third reticulate rugose and remaining surface sculptured by many longitudinal subparallel irregular keels in A. makererense.
Male. Unknown. Differential diagnosis. Because of the above diagnosis, the new species is similar to Anteon abditum Olmi, 1994a. The main difference regards the shape of the pronotum: without raised carinae in A. abditum; with raised carina on anterior and lateral margins in A. mubfs.
Etymology. Named after the acronym of Makerere University Biological Field Station, where the holotype was collected. The field station is affectionately called "Mubfs" by those privileged to have experienced a stay there. Noun in apposition.

Hosts. Unknown.
Distribution. Democratic Republic of the Congo and South Africa (Olmi 1984(Olmi , 2006. Newly recorded from Uganda here.

Bocchus bini
Hosts. Unknown. Distribution. Afrotropical, recorded from Ghana to Somalia and Yemen (Olmi and van Harten 2006;Olmi et al. 2015), in addition to Madagascar . Newly recorded from Uganda here. Diagnosis. Male with OPL slightly longer than POL; antennal segment 6 less than twice as long as broad; scutum and scutellum completely reticulate rugose; notauli absent; posterior surface of propodeum with median area crossed by numerous transverse keels.
Female. Unknown. Differential diagnosis. Because of the above diagnosis, the new species is similar to Bocchus seyrigi (Benoit, 1954). The main differences regard OPL/POL ratio (OPL slightly longer than POL in B. kibalensis; OPL more than twice as long as POL in B. seyrigi) and the sculpture of the median area of posterior surface of propodeum (crossed by many transverse keels in B. kibalensis; partly unsculptured in B. seyrigi).
Etymology. The species is named after the type locality, in Kibale National Park. Hosts. Unknown. Distribution. Uganda.
Distribution. Afrotropical, recorded from Angola, Democratic Republic of the Congo, Eritrea, Namibia, Somalia, South Africa, Swaziland, Tanzania, Uganda, Zimbabwe (Olmi 1984(Olmi , 2006(Olmi , 2009. Diagnosis. Female with head black, except mandible and clypeus testaceous; head flat, with posterior margin of vertex convex and TL more than twice as long as POL; head (dorsally viewed) provided with posterior ocelli placed behind imaginary straight line joining posterior edges of eyes; posterior ocelli almost touching occipital carina; head and scutum granulated and reticulate rugose, not sculptured by longitudinal keels or striae; pronotum black, except posterior collar ferruginous; posterior collar of pronotum present; propodeum black; segment 1 of protarsus slightly longer than segment 4; enlarged claw (Figs 2B, 11F) approximately as long as segment 5 of protarsus.

Hosts. Unknown.
Distribution. Recorded from many Afrotropical countries, from Nigeria to Somalia (Olmi and Copeland 2011).
Distribution. Afrotropical, recorded from Burundi, Central African Republic, Democratic Republic of the Congo, Kenya, Mozambique and Uganda .

Subfamily Gonatopodinae Kieffer, in
Distribution. Recorded from many Afrotropical countries, from the Cape Verde Islands, Niger and Senegal to Madagascar and Somalia, in addition to Oman and Egypt (Olmi 1984(Olmi , 1994b(Olmi , 2006Olmi et al. 2015). Newly recorded from Uganda here.
Distribution. Recorded from many Afrotropical countries (from the Cape Verde Islands to Somalia, including Madagascar and South Africa), in addition to many European countries, Oman, Yemen and the United Arab Emirates Olmi 1999bOlmi , 2004cOlmi , 2005cOlmi , 2008van Harten 2000, 2006). Newly recorded from Central African Republic here.
Distribution. Recorded from many countries of the Palaearctic, Afrotropical and Nearctic Regions ; in Africa recorded from many sub-saharian countries, from Benin to South Africa and Ethiopia (Olmi 1984(Olmi , 2006. Hosts. Unknown. Distribution. Cameroon . Newly recorded from Central African Republic here.
Male. Unknown. Differential diagnosis. Because of the above diagnosis, the new species is similar to Gonatopus trochantericus (Benoit, 1954) and G. tulearensis . The main difference regards the metanotum (with sides protruding (rounded protrusions) in G. kanyawarus (Fig. 3A); with sides rounded and not protruding in the other two species (Figs 3D, E)).

Neodryinus tussaci
Distribution. Recorded from many countries of the world, excluding the Neotropical Region . In Africa known from Burundi, Kenya, Madagascar, Somalia, Tanzania, Togo, Uganda and Northern Africa Olmi and Copeland 2011). Known also in Yemen .

Discussion
The checklist of the Dryinidae, Embolemidae and Sclerogibbidae of Central African Republic and Uganda presented in this paper includes 23 species of Dryinidae, two species of Embolemidae and three species of Sclerogibbidae known from Central African Republic; 39 species of Dryinidae, one species of Embolemidae and four species of Sclerogibbidae known from Uganda. Seventeen species of Dryinidae, two species of Embolemidae and one species of Sclerogibbidae were newly recorded from Central African Republic; sixteen species of Dryinidae and one species of Embolemidae were reported for the first time in Uganda.
With 1827 species worldwide (Olmi and Xu 2015), Dryinidae is one of the three largest families in the Chrysidoidea, the other two being Chrysididae and Bethylidae. With the possible exception of Kenya, Madagascar, Mozambique and South Africa, the dryinid fauna of the Afrotropical Region, like that of many families of micro-Hymenoptera, is poorly known. A comparison with the better known Afrotropical countries shows the following situation: in Madagascar 123 species of Dryinidae, six species of Embolemidae and seven species of Sclerogibbidae are recorded ; in South Africa 174 species of Dryinidae (unpublished datum; 119 reported by Olmi (2006)), seven species of Embolemidae (unpublished datum; six reported by Olmi (2006)) and six species of Sclerogibbidae are recorded ; in Mozambique 45 species of Dryinidae (Olmi et al. 2012), no species of Embolemidae (unpublished datum) and three species of Sclerogibbidae are known ; in Kenya 76 species of Dryinidae, six species of Embolemidae and 12 species of Sclerogibbidae are reported by Olmi et al. (2015).
Hosts of Dryinidae are known for 12 of the 52 species reported in the presented checklists in this paper, but no new host associations were established during the recent inventory surveys conducted in Uganda and Central African Republic. No hosts of Embolemidae are known in the entire Afrotropical region. Host associations are better known for the Sclerogibbidae, mainly due to the rearing out of parasitized hosts carried out by Edward S. Ross: the hosts are known for seven of the eight species reported in the presented checklists, in one case as a result of rearing of adult wasps in Uganda.
Further baseline species inventory and establishment of host associations are required across the majority of the countries in the region to enable assessment of overall species richness and biology of the Afrotropical fauna for these families.