Research Article
Research Article
First records of Trichina Meigen, Euthyneura Macquart and Oedalea Meigen (Diptera, Hybotidae) from North Africa, with descriptions of two new species
expand article infoLaila Zouhair, Patrick Grootaert§, Kawtar Kettani
‡ Abdelmalek Essaadi University, Tetouan, Morocco
§ Royal Belgian Institute of Natural Sciences, Brussels, Belgium
Open Access


Trichina Meigen, 1830, Euthyneura Macquart, 1836 and Oedalea Meigen, 1820 are reported for the first time in North Africa from northern Morocco, with seven species including two ones new to science, based on material collected at nine sites located in the Moroccan sector of the Mediterranean Intercontinental Biosphere reserve (MIBR). These new records represent also the first evidence of the occurrence of Trichininae and Oedaleinae subfamilies throughout North Africa and bring the total of Moroccan hybotid fauna to 51 species. The new species are described and illustrated. A key to Moroccan Trichina species is provided.


Hybotidae, Morocco, new records, new species, North Africa, Oedaleinae, Trichininae


Trichina Meigen, Euthyneura Macquart and Oedalea Meigen genera were previously included within the Trichiniini and Oedaleini tribes, which were included for a long time beside the Ocydromiini tribe in Ocydromiinae subfamily in Chvála’ s (1983) classification. More recent classifications, such as Sinclair and Cumming (2006) and Wahlberg and Johanson (2018), now treat these three tribes as separate subfamilies within the Hybotidae.

The genus Trichina attributed to the subfamily Trichininae consists of small black species (2.5–3 mm), with small mouthparts, large wing stigma extending to tip of R2+3, and hind tibiae dilated towards tip, and whose adults are known as predaceous in both sexes and occur for a long period during the summer in somewhat shaded humid biotopes (Chvála 1983). They are often found in the low grasses in deciduous forests, and are also observed at the tips of branches or along forest margins from approximately sunset until dusk (Chvála 1983). Larvae of Trichina occur probably in the soil (Smith 1989). The genera Euthyneura and Oedalea are both assigned to the Oedaleinae subfamily historically considered as Oedaleini tribe, which includes the most abundant of all flower-visiting empidoids in the Holarctic region (Marshall 2012). Species of Euthyneura are small black flies (2–3 mm) with a relatively long proboscis directed forward and simple legs devoid of long bristles; they are usually found on flowers and low herbage, but also on conifers (Chvála 1983). Among the representatives of the family Hybotidae, this genus represents one of the few groups whose adults feed only on nectar and pollen and are entirely flower visitors (Chvála 1983; Chandler 1992; Shamshev and Kustov 2012), while their larvae develop in rotting wood (Chvála 1983). The Oedalea species are more robust than those of Euthyneura. These are predaceous flies with conspicuously long antennae, shiny black thorax and raptorial hind legs (Chvála 1983; Shamshev 2020) and whose adults can usually be swept in low numbers from leaves of trees and shrubs, while the larvae can be bred from dead wood (Beuk 1992).

Various studies have been recently carried out on the aforementioned genera, mainly in Europe in the west Palaearctic region (Kanavalová et al. 2021). Thus, Euthyneura is known from the Palaearctic region with seven species (Shamshev and Kustov 2012) whereas data for Trichina and Oedalea in this region remain very poor and do not exceed seven species for Trichina and fourteen for Oedalea known to date from Europe (Barták and Kubík 2009; De Jong et al. 2014; Evenhuis and Pape 2021). As for North Africa, studies on hybotids are even scarcer and rather fragmentary, as is the case for Morocco where only 44 species of Hybotidae are recorded so far (Kettani et al. 2022), reflecting the lack of studies on such an important family of Diptera in the region.

In the present paper, we report the first record of Trichina, Euthyneura and Oedalea genera from the whole of North Africa, thus representing the first evidence of the occurrence of Trichininae and Oedaleinae subfamilies in North Africa, knowing that all species found so far in the west Palaearctic region of the genera of these subfamilies have only been recorded in Europe (De Jong et al. 2014), and no records have been reported until now in North Africa. Importantly, the genus Euthyneura is recorded here for the first time in the entire Mediterranean region.

In addition to these first records, we provide evenly here diagnosis and descriptions of two species described as new to science, belonging to the genus Trichina: Trichina azizi Zouhair & Grootaert sp. nov. and Trichina rifensis Zouhair & Grootaert sp. nov.

Materials and methods

The studied specimens originate from the entomological field surveys undertaken by the first author (LZ) over the years 2020 and 2021, and by the third author (KK) between 2017 and 2019. Specimens are preserved in 70° alcohol and housed in the private collection of the first author (PCLZ) at the University Abdelmalek Essaadi (Tetouan, Morocco) and in the private collection of the second author (PCPG). Most of the collected specimens were sampled using sweep net, and some with Malaise trap.

The specimens examined were mainly collected in mixed forests and riparian areas at nine sites located in the northern part of Morocco (Table 1, Fig. 1). With the exception of two forest sites situated within unprotected areas (S8, S9), the remaining sampling sites belong to the National Park of Talassemtane (NPTL) and the Project of Natural Park of Bouhachem (PNPB), which are considered the most important protected areas in terms of conservation in Morocco. The nine sites are located in the Rif, which consists of a mountainous chain located in the northernmost part of Morocco. This chain represents, along with the Atlas mountain region, the most important endemic areas of the country. It is important to highlight that all the studied sites belong also to the Mediterranean Intercontinental Biosphere Reserve (MIBR) which encompasses southern Spain and northern Morocco, and is known as a reservoir of biodiversity with regard to the Moroccan part (Rosas et al. 2017; Bachar et al. 2021).

Table 1.

Coordinates, altitudes and localities of the studied sites.

Code Site Protected area, locality Province Altitude Geographical coordinates Habitat Collecting tool
S1 Tissgris PNPB, Hmmadesh Tetouan 505 m 35.397°N, 5.522973°W Mixed forest Sweep net
S2 Oued Stah PNPB, Bni Aarouss Larache 766 m 35.2764°N, 5.531389°W River bank Sweep net
S3 Oued Asellam PNPB, Moulay Abdessalam Tetouan 1267 m 35.2657°N, 5.483595°W River bank Sweep net
S4 Lemtahane PNPB, Dar Abdessalam Tetouan 964 m 35.2708°N, 5.434864°W Mixed forest Malaise trap
S5 Sefihat telj NPTL, Talembote Chefchaouen 1745 m 35.1852°N, 5.211767°W Fir forest Sweep net
S6 Bouslimane NPTL, Jbel Bouslimane Chefchaouen 1350 m 35.0971°N, 5.14505°W Fir forest Sweep net
S7 Oued El Ferda NPTL, Akoumi Chefchaouen 447 m 35.237°N, 5.176283°W River bank Sweep net
S8 Aïn Lahcen Unprotected area, Aïn Lahcen Tetouan 316 m 35.5601°N, 5.578017°W Pine forest Sweep net
S9 Fifi Unprotected area, Bab Taza Chefchaouen 1332 m 34.9803°N, 5.2266°W Mixed forest Sweep net
Figure 1. 

Location of the nine studied sites in the north of Morocco.

In the Rif, the climate is of the Mediterranean type and is composed of two distinct periods, a hot and dry summer and the other a relatively cold and rainy winter (Al Karkouri 2017). The bioclimate is primarily wet with high humidity which allows the growth of a rich flora, including nearly all of the Moroccan forest species such as deciduous groves, the endemic fir forests of Morocco (Abies marocana), the Atlas Cedar tree (Cedrus atlantica), Pine forests (Pinus pinaster), Cork Oak (Quercus suber), Olives, Thuja articulata (Tetraclinis articulata), Kermes Oak (Quercus coccifera), Holm Oak (Quercus ilex), and the coccifera which have formed extensive forests in the western part of the Rif (Taiqui 1997).

Most species were recognized using a stereomicroscope, but for the new ones it was necessary to make a preparation of the male genitalia for their accurate identification. The male terminalia were removed from the insect body, macerated in the 10% KOH for 24h in order to dissolving the tissues. When all darkly sclerotized structures were transparent enough, the maceration process was stopped by removing the terminalia from the KOH and bringing them to a vial with tap water. After at least 5 minutes a drop of vinegar was added to the vial to stop the maceration process completely. After about a minute the terminalia were rinsed in vial with tap water again and after a few minutes transferred to 70% ethanol for 10 minutes and transferred to pure glycerin. They were preserved in the glycerol to be later observed under the microscope and drawn using a camera obscura.

Morphological terminology and abbreviations largely follow Chvála (1983), except for the male terminology, which mostly follows Sinclair and Cumming (2006). The arrangement of taxa follows also Sinclair and Cumming (2006).


Family HYBOTIDAE Meigen, 1820

Subfamily TRICHININAE Chvála, 1983

Genus Trichina Meigen, 1830

Trichina elongata Haliday, 1833

Fig. 7A

Material examined

1♂. Morocco, Rif, Sefihat telj, NPTL, 1745 m, 04.iii.2020, sweep net, leg. L. Zouhair, PCLZ; 1♂. Rif, Akoumi, Oued El Ferda, NPTL, 447 m, 13.iii.2021, sweep net, leg. L. Zouhair, PCLZ.


Widespread throughout Europe, including the European part of Russia (Shamshev and Kustov 2006). First record for Morocco.

Trichina opaca Loew, 1864

Fig. 7B

Material examined

1♂. Morocco, Rif, Fifi, 1332 m, 6.v.2021, sweep net, leg. L. Zouhair, PCLZ.


Known from Central and Northern Europe (Shamshev and Kustov 2006). First record for Morocco.

Trichina unilobata Chvála, 1981

Fig. 7C

Material examined

1♂. Morocco, Rif, Aïn Lahcen, 316 m, 10.i.2020, sweep net, leg. L. Zouhair, PCLZ.


Spain, Turkey. First record for Morocco.

Descriptions of new species

Trichina azizi Zouhair & Grootaert, sp. nov.

Figs 3A–D, 7D

Material examined

Holotype. 1♂. Morocco, Rif, Bouslimane, NPTL, 1350 m, 28.iv.2019, sweep net, leg. K. Kettani, PCLZ.


(S6: Bouslimane) (Fig. 2): The holotype was swept from the foothills of Bouslimane mountain located at the south of the national park of Talassemtane. The habitat consists of a fir forest (Abies marocana) dominated by a humid bioclimate as part of the mesomediterranean zone. This fir formation grows on brown fersiallitic soil.

Figure 2. 

Type habitat of Trichina azizi Zouhair & Grootaert, sp. nov. in the fir forest at Bouslimane locality (Photo: K. Kettani, 28.04.2019).

Differential diagnosis

Trichina azizi sp. nov. has 3rd antennal segment considerably long and slender like in T. elongata according to the specimen we have and to the description in Chvála (1983), but it differs in length since it is less than 3.4 times as long as broad in T. elongata (Barták and Kubík 2009) whereas it is more than 3.6 times as long as broad in the new species. Both species differ also in the length of the stylus, which in the new species is little longer than half-length of 3rd segment; while in T. elongata, the stylus is less than half the length of the 3rd segment. The bare terminal part of antennae is shorter than the pubescent thicker basal part which is the same in both. The new species has only two pairs of scutellar setae, like in T. elongata and also in T. clavipes and T. bilobata. In addition, it can be recognized by yellowish hind trochanter, femora, tibiae and metatarsi, and by the lack of a ventral spine on hind trochanters, like in T. opaca. Male terminalia with the hypandrial projection is rather long, like in T. elongata, but in the latter it is more slender than in the new species. The left surstylus is similar in structure to that of T. elongata, but it is not sharpened apically like in T. elongata. The right surstylus is spiny apically and sub-apically, which is the most important diagnostic character in this new species. Cerci are long and well developed but the left cercus is longer than the right one.


This species is dedicated to the father (Aziz Zouhair) of the first author.


Male. Small brown species (2.8 mm) (Fig. 7D).

Head. Brown in ground colour. Ommatidia bicoloured. Antennae brown, with postpedicel considerably long and slender with 0.22 mm long over 0.06 mm wide, stylus is 0.134 mm long (0.6 × the length of postpedicel hence a little longer than half the length), its bare terminal part shorter than the pubescent thicker basal part. Palpus small, brownish, with a brownish bristle. Proboscis paler. Face narrow, less than width of scape.

Thorax. Entirely brown, (unfortunately, all thoracic setae are broken, but according to setae follicles, 2 pairs of scutellar setae). Mesonotum shining black except for the extreme anterior border that bears some sparse microtrichia, the notopleural depression with some denser microtrichia, but is still subshining black. Wings very faintly brown yellowish infuscated, veins pale brown, pterostigma yellowish brown and indistinct, costa running to tip of M1, squamae including fringes brown, halters brown yellowish. Legs almost uniformly brownish leaving hind trochanter, femora, tibia, and hind metatarsi yellowish. Fore femur slender with paler hairs; mid femur with 2 ventral rows of dark and thick bristles; hind femur longer than mid and fore femora, with a row of thick and long spine-like bristles. No spine on the hind trochanter. Fore tibia more slender at base, covered with small paler hairs; mid tibia very slender than fore tibia and covered mostly with dark hairs; hind tibia much longer than fore and mid tibiae, laterally compressed and dilated towards tip.

Abdomen. Brown, tergites and sternites covered with scattered black setulae. Terminalia somewhat large and blackish brown, with spiny right surstylus in apical and sub-apical parts (Fig. 3B, D), left surstylus longer and slender in comparison with right surstylus, hypandrial projection moderately long, slender (Fig. 3D). Cerci longer than wider, the left cercus longer than the right cercus (Fig. 3B).

Figure 3. 

Trichina azizi Zouhair & Grootaert sp. nov., Holotype male, terminalia A right surstylus, lateral view B epandrium, dorsal view C left surstylus, lateral view D epandrium, ventral view. Scale: 0.1 mm. (Drawn by Patrick Grootaert).

Female. (Unknown).

Abbreviations: ae: aedeagus (phallus); cer: cercus; hyp: hypandrium; hypp: hypandrial projection; lel: left epandrial lamella; ls: left surstylus; rel: right epandrial lamella; rs: right surstylus.

Trichina rifensis Zouhair & Grootaert, sp. nov.

Figs 5A–D, 7E

Material examined

Holotype. 1♂. Morocco, Rif, PNPB, Lemtahane, 964 m, 7.v.2017–30.v.2017, Malaise trap, leg. K. Kettani, PCLZ.


(S4: Lemtahane) (Fig. 4): The Malaise trap was set up in a scrubland composed of Pinus pinaster and some fruit trees growing on siliceous soil. The bioclimate is subhumid and favours thermo-Mediterranean vegetation.

Figure 4. 

Type habitat of Trichina rifensis Zouhair & Grootaert sp. nov., in Lemtahane (Photo: K. Kettani, 07.05.2017).

Differential diagnosis

The newly described species Trichina rifensis sp. nov., is very similar to T. opaca as it is described in Chvála (1983) and according to the key of Trichina in Barták and Kubík (2009) and to our T. opaca specimen, in: the absence of the ventral spine in hind trochanters, and in the colour of legs which are extensively blackish to blackish brown, with the knees and extreme base of tibiae often paler. It is similar to T. opaca also in having the same number of scutellar pairs setae (3 pairs). The wings are conspicuously darkened brown, veins blackish, squamae including fringes and halters deep black in both species. However, both species differ in the pruinosity of the mesonotum: in the new species, the mesonotum is shining black but anteriorly there is a very narrow stripe with weak microtrichia and the notopleural depression is more distinctly set with microtrichia (not grey dusted), while in T. opaca the mesonotum is entirely thinly brownish pollinose and consequently rather dulled, and differ also in the length of the 3d antennal segment that is less than 3.2 times as long as broad in T. opaca, while it is more than 3.2 times (4 times) as long as broad in T. rifensis sp. nov. The male terminalia in the new species is with the hypandrial projection much longer than in T. opaca, the structure of the left surstylus is similar in the two species so that is C-shaped in both. The right surstylus of the new species is spiny in the apical and sub-apical parts contrary to T. opaca, which forms an important differential diagnosis character, cerci are equal in length and shape in T. opaca, while in the new species, the left cercus is narrower and longer than the right cercus which is shorter and conical in shape.


The new species is named rifensis, after the Moroccan Rif region where it was found.


Male. Black species with body small (3 mm) Fig. 7E.

Head. Black in ground colour. Eyes meeting on frons for a long distance. A distinct prominent ocellar tubercle with 2 pairs of ocellar bristles, anterior pair as long as posterior one, occiput and vertex finely greyish pollinose, covered with black and distinct hairs. Face linear, narrow less than width of scape. Antennae entirely black, inserted at middle of head in profile with postpedicel is 0.24 mm long over 0.06 mm wide, stylus (apical naked part missing) has the basal part and the thickened second segment 0.079 mm long. Palpus short with a very long black apical bristle, 1.5 times as long as palpus and a subapical bristle half as long as apical one. Proboscis yellowish, pointing obliquely forward and covered with several brownish hairs.

Thorax. Polished black with all hairs black and long. Acr biserial, dc uniserial, ending in 1 pair of very long prescutellars, 1 humeral, 1 notopleural, 1 postalar bristles, 3 scutellar pairs. Mesonotum covered with microtrichia. Mesonotum shining black but anterior part with a very narrow stripe with weak microtrichia, notopleural depression more distinctly set with microtrichia (not grey dusted). Wings conspicuously brownish, stigma blackish brown extending to tip of vein R2+3, veins blackish brown, costa reaching vein M1, squamae, including fringes, and halters black. Legs mostly covered with hairs, extensively blackish leaving base of hind tibia and knees paler. No spine on hind trochanter. Fore femur rather slender, covered with paler hairs and bearing 6–8 black bristles apically; mid femur with 2 rows of paler bristles and 2–4 black bristles apically; hind femur much longer than fore and mid femora with ventral and dorsal paler bristles. Fore tibia rather slender, with black hairs; mid tibia with distinct a pair of black and strong bristles; hind tibia much longer than fore and mid tibiae, laterally compressed and dilated towards tip, with 2 rows of black bristles, with one dorsal black bristle.

Abdomen. Tergites and sternites blackish, covered with scattered black setulae. Terminalia black, with right surstylus somewhat robust and spiny in sub-apical and apical parts (Fig. 5B, D), left surstylus slender and hypandrial projection long and slender. Right cercus shorter, conical, left cercus longer and narrower (Fig. 5B).

Figure 5. 

Trichina rifensis Zouhair & Grootaert sp. nov., Holotype male terminalia A right surstylus, lateral view B epandrium, dorsal view C left surstylus, lateral view D epandrium, ventral view. Scale: 0.1 mm. (Drawn by Patrick Grootaert).

Female. (Unknown)

Abbreviations: ae: aedeagus (phallus); cer: cercus; hyp: hypandrium; hypp: hypandrial projection; lel: left epandrial lamella; ls: left surstylus; rel: right epandrial lamella. rs: right surstylus.

General comments

Despite the male terminalia of the two new species described above being very similar in structure, it is easy to distinguish between the both species: Trichina azizi sp. nov. has yellowish hind trochanters, femora tibiae and metatarsi, so it is very distinct from Trichina rifensis sp. nov., which has entirely black hind trochanters, femora, tibiae (except at base) and tarsi, as may be seen in Fig. 7D, E.

Compared to other species of Trichina described so far, Trichina rifensis sp. nov., is very close to Trichina opaca in most morphological characters (colour of legs and wings, shape of antennae, number of scutellar pairs, absence of hind trochanters spine(s)) but as we previously noted in the differential diagnosis to this new species, it differs clearly in the male terminalia. As to Trichina azizi sp. nov., is similar to T. elongata in the shape of the antennae, number of scutellar pairs and absence of the trochanter spine, but it is easy to differentiate these species by the male terminalia as we have described in the differential diagnosis above. The common character between the right surstylus of the both new species is that they are spiny, which is a character not described in Trichina at all.

Key to the Moroccan species of Trichina Meigen (males)

This key is compiled by referring to the key of Trichina in Barták & Kubík (2009).

1 The whole of mesoscutum microtrichose. 3d antennal segment usually less than 3.2 times as long as broad T. opaca Loew, 1864
The central parts of mesoscutum without microtrichiae. 3d antennal segment more than 3.4 times as long as broad 2
2 Face broader T. unilobata Chvála, 1983
Face narrow 3
3 Hind trochanter with posterior to ventral spine(s)-like setae. Legs uniformly brownish. Right surstylus not spiny in apical and sub-apical parts T. elongata Haliday, 1833
Hind trochanter without posterior to ventral spine(s)-like setae. Hind trochanter, femora, tibia and metatarsi yellowish (T. azizi sp. nov.) or at least knees and base of hind tibia paler (T. rifensis sp. nov.). Right surstylus spiny in apical and sub-apical parts 4
4 3 pairs of scutellar setae T. rifensis sp. nov.
2 pairs of scutellar setae T. azizi sp. nov.

Subfamily OEDALEINAE Chvála, 1983

Genus Euthyneura Macquart, 1836

Euthyneura myrtilli Macquart, 1836

Fig. 7F

Material examined

1♀. Morocco, Rif, PNPB, Lemtahane, 964 m, 7.v.2017–30.v.2017, Malaise trap, leg. K. Kettani, PCLZ; 1♀. Rif, PNPB, Oued Asellam, 1267 m, 22.iii.2021, sweep net, leg. L. Zouhair, PCLZ ; 1♀. Rif, PNPB, Oued Stah, 766 m, 2.v.2021, sweep net, leg. L. Zouhair, PCLZ.


Common in northern and central Europe, absent in the south (Chvála and Vonicka 2008); also European part and Western Siberia of Russia (Shamshev 2016). First record for Morocco.

Genus Oedalea Meigen, 1820

Oedalea portugalica Barták & Grootaert, 2021

Figs 6A–E, 7G

Material examined

1♀. Morocco, Rif, PNPB, Tissegris, 505 m, 20.iv.2021, sweep net, leg. L. Zouhair, PCLZ.


Known up to present only from the type locality in Portugal. First record for Morocco.


Oedalea portugalica was described by Barták & Grootaert (2021) in Kanavalová et al. (2021) from Portugal. The male terminalia were not illustrated or described, and are provided herein for the first time.

Terminalia (Fig. 6) small. Cerci (Fig. 6B) digitiform, covered toward tip with long setae, right cercus (Fig. 6B) slightly longer and more narrowed apically than left cercus. Epandrium covered with long setae (Fig. 6A, C), right epandrial lamella somewhat longer, broader and more elongated towards apex (Fig. 6A) than left epandrial lamella (Fig. 6C).

Figure 6. 

Oedalea portugalica Barták & Grootaert, 2021. Male terminalia A right epandrial lamella, lateral view B epandrium, dorsal view C left epandrial lamella, lateral view D hypandrium, ventral view E left postgonite with aedeagus (phallus). Scale: 0.1 mm. (Drawn by Patrick Grootaert).

In Kanavalová et al. (2021), O. portugalica was compared with O. stigmatella. In the terminalia of O. stigmatella (figs 401–404 in Chvála 1983), the shape of the right and left epandrial lamellae is different, so that in O. portugalica there is a deep medial apical incision in the hypandrium (it is forked Fig. 6D), while in O. stigmatella the fork is shallow and broader (fig. 403 in Chvála 1983). The tip of the left postgonite is very characteristic and different. A sharp protrusion is apical in O. stigmatella (fig. 404 in Chvála 1983) while it is lateral (subapical) in O. portugalica (Fig. 6E seen from the right, while on Fig. 6A it is below the right epandrial lamella seen from the right side).

Abbreviations: ae: aedeagus (phallus); cer: cercus; hyp: hypandrium; lep: left epandrial lamella; lpgt: left postgonite; rep: right epandrial lamella; rpgt: right postgonite.

Figure 7. 

A Trichina elongata Haliday, 1833 B Trichina opaca Loew, 1864 C Trichina unilobata Chvála, 1981 D Trichina azizi Zouhair & Grootaert sp. nov. E Trichina rifensis Zouhair & Grootaert sp. nov. (Left antennal style missing) F Euthyneura myrtilli Macquart, 1836 G Oedalea portugalica Barták & Grootaert, 2021.


The current study contributes significantly to the enrichment of the faunistic database of the Hybotidae fauna of Morocco in particular and that of North Africa in general. Seven species belonging to three genera (Trichina, Euthyneura and Oedalea) which were unprecedentedly reported in North Africa as well as their respective subfamilies (Trichininae and Oedaleinae) are newly recorded in the region, increasing the total of the known hybotid species of Morocco from 44 species to 51. The description of two new species belonging to the Trichina genus provides also an important contribution to the hybotid fauna. Our results show that the distribution of the three mentioned genera has expanded beyond Europe where they were originally recorded.

These findings highlight the richness of the Moroccan biodiversity, particularly in the Western Rif, which is considered a biodiversity hotspot, where the new species were found, and which constitutes the only part of Morocco included in the Mediterranean Intercontinental Biosphere Reserve (MIBR) due to the great specific richness and the high rate of endemism recorded there (Bachar et al. 2021). Indeed, of the 51 species of Hybotidae occurring in Morocco, 25 of them are cited exclusively in the Rif.

As noted by Stark (2008) regarding the weak diversity of the subfamilies Trichininae and Oedaleinae compared to the Tachydromiinae, our results exhibit the same pattern following our study of ancient and recent hybotid samples collected from various habitats in Morocco by the first and the third authors, where Tachydromiinae subfamily was remarkably always dominant. However, this can also be explained by the fact that the Moroccan Hybotidae fauna is strongly under-collected and there is still at least as much to discover there. There is no doubt that the number of species of Hybotidae occurring in Morocco as in North Africa will increase when collecting is more intensified.


  • Al Karkouri J (2017) Les milieux montagneux marocains à l’épreuve du changement climatique (cas de la montagne rifaine). Hespéris-Tamuda LII 1: 237–267.
  • Bachar M, ElYacoubi H, Zidane L, Rochdi A (2021) Ethnomedicinal and traditional phytotherapeutic plants used in Bouhachem Natural Regional Park (Rif of Morocco): Case of Bni-Leit and Al-Oued districts. Journal of Pharmacy & Pharmacognosy Research 9(3): 284–312.
  • Beuk PLTh (1992) A remarkable aggregation of flies above a road surface, including ten species new to the dutch fauna (Diptera). Entomologische berichten 52(8): 105–110.
  • Chandler P (1992) Anthalia beatricella sp. nov. and two other additions to the British list of Oedaleini (Diptera, Empididae). Dipterists Digest 12: 16–22.
  • Chvála M (1981) Empididae (Insecta: Diptera) from southern Spain, with descriptions of twenty new species and notes on Spanish fauna. Steenstrupia 7: 113–177.
  • Chvála M (1983) The Empidoidea (Diptera) of Fennoscandia and Denmark. II. General part. The families Hybotidae, Atelestidae and Microphoridae. Fauna entomologica Scandinavica 12: 1–279.
  • Chvála M, Vonicka P (2008) Hybotidae (Diptera) of the Jizerské hory Mts and Frýdlant region (northern Bohemia, Czech Republic). Sborník Severočeského muzea, Přírodní Vědy 26: 237–251. [ISBN 978-80-87266-00-7]
  • De Jong Y, Verbeek M, Michelsen V, Bjørn PDP, Los W, Steeman F, Bailly N, Basire C, Chylarecki P, Stloukal E, Hagedorn G, Wetzel FT, Glöckler F, Kroupa A, Korb G, Hoffmann A, Häuser C, Kohlbecker A, Müller A, Güntsch A, Stoev P, Penev L (2014) Fauna Europaea - all European animal species on the web. Biodiversity Data Journal 2: e4034.
  • Evenhuis NL, Pape T [Eds] (2021) Systema Dipterorum, version 3.5. [updated 20 October 2021]
  • Haliday AH (1833) Catalogue of Diptera occurring about Holywood in Downshire. Entomologist′s monhly Magazine 1: 147–180.
  • Kanavalová L, Grootaert P, Kubík Š, Barták M (2021) Four new West Palaearctic species and new distributional records of Hybotidae (Diptera). ZooKeys 1019: 141–162.
  • Kettani K, Ebejer MJ, Ackland DM, Bächli G, Barraclough D, Barták M, Carles-Tolrá M, Černý M, Cerretti P, Chandler P, Dakki M, Daugeron C, De Jong H, Dils J, Disney H, Droz B, Evenhuis N, Gatt P, Graciolli G, Grichanov IY, Haenni JP, Hauser M, Himmi O, MacGowan I, Mathieu B, Mouna M, Munari L, Nartshuk EP, Negrobov OP, Oosterbroek P, Pape T, Pont AC, Popov GV, Rognes K, Skuhravá M, Skuhravý V, Speight M, Tomasovic G, Trari B, Tschorsnig HP, Vala JC, von Tschirnhaus M, Wagner R, Whitmore D, Woźnica AJ, Zatwarnicki T, Zwick P (2022) Catalogue of the Diptera (Insecta) of Morocco – an annotated checklist, with distribution and bibliography. ZooKeys 1094: 1–466.
  • Loew H (1864) Ueber die schlesischen Arten der Gattungen Tachypeza Meig. (Tachypeza, Tachista, Dysaletria) und Microphorus Macq. (Trichina und Microphorus). Zeitschrift für Entomologie. Breslau 14: 1–50.
  • Macquart PJM (1836) Description d’un nouveau genre d’insectes diptères de la famille des tanystomes. Annales de la Société Entomologique de France 5: 517–520.
  • Marshall SA (2012) Flies. The Natural History & Diversity of Diptera. Firefly Books, 616 pp.
  • Rosas M, Daniëls PP, Infante F (2017) Ethnomycological study of the Moroccan sector of the Intercontinental Biosphere Reserve of the Mediterranean. Boletin de la Sociedad Micologica de Madrid 41: 117–127.
  • Shamshev IV (2016) An annotated checklist of empidoid flies (Diptera: Empidoidea, except Dolichopodidae) of Russia. Proceedings of the Russian Entomological Society 87: 3–183.
  • Shamshev IV (2020) First record of the genus Oedalea Meigen (Diptera: Hybotidae) from Vietnam, with description of a new species and a key to species of the Oriental Region. Russian Entomological Journal 29(2): 218–221.
  • Smith KGV (1989) An introduction to the immature stages of British flies: Diptera larvae, with notes on eggs, puparia and pupae. Handbooks for the Identification of British Insects. Part 14. Royal entomological society of London 10: 163.
  • Stark A (2008) Species of the genus Oedalea Meigen, 1820 (Diptera: Hybotidae): An element of the canopy fauna in European forests? In: Floren A, Schmidl J (Eds) Canopy arthropod research in Europe, bioform entomology, Nuremberg. 105–117.
  • Wahlberg E, Johanson KA (2018) Molecular phylogenetics reveals novel relationships within Empidoidea (Diptera). Systematic Entomology 43(4): 619–636.
login to comment