Corresponding author: Torsten Dikow (
Academic editor: Pierfilippo Cerretti.
The
The southern African
At the start of this review,
As can be seen from the above information,
This review is based on an additional 61 specimens from numerous natural history collections accumulated over the past 35 years representing all previously known species. Such an increase in specimen number, their geographic occurrence expanding the range of the genus considerably, and substantial morphological variation suggested that a few new species might be represented among the material. However, this not the case and to the contrary the number of valid species is reduced to three by synonymy.
Morphological terminology and abbreviations for setae follows
The female genitalia and male terminalia are first excised and macerated in 10% potassium hydroxide (
The following data on species occurrences are given (where available): country, state/province, county, locality, geographic co-ordinates (formatted in both decimal and degrees minutes seconds latitude/longitude), elevation (in meters), date of collection (format: yyyy-mm-dd), habitat information, sampling protocol (if other than hand netting), collector, catalog number (a unique specimen number and any other identifying number), depository (institution and collection code), number of specimens and sex, life stage, name of person who identified the specimen, and any other previous identifications (note that for synonymized species the holotype still retains its status as a primary type specimen and therefore the particular material examined list will include two (or more) holotypes; see the entry under 'previousIdentifications' for the original identification by the author). Each specimen is listed with a unique specimen number (either an institutional catalog number or an AAM-XXXXXX number used by the senior author) that will allow the re-investigation as well as provide a unique
Institutions providing specimens are listed below, together with the abbreviations used in the text when citing depositories (institutionCode), a link to the record in the Global Registry of Biodiversity Repositories (
Lucid Builder: illustrated, multi-entry, matrix-based identification key –
Lucid Phoenix: illustrated, dichotomous identification key –
The genus (Fig.
Males are unique in the arrangement of their medially fused aedeagal prongs and the yellow to light brown abdominal coloration. Females in contrast are more generalized and similar to other female
Head: brown, in general grey pubescent; width distinctly greater than thorax, interocular distance on vertex larger than at ventral eye margin, vertex between compound eyes ± horizontally straight, medially only slightly below dorsal eye margin, parafacial area about as wide as ½ the width of central facial gibbosity; facial gibbosity distinct, well-developed and discernible in lateral view; mystax yellow, covering entire facial gibbosity; frons not elevated, predominantly apubescent; vertex entirely grey pubescent; postgena lightly grey pubescent; setation: vertex yellow, frons white or yellow, ocp setae white, pocl macrosetae absent; ocellar triangle apubescent; proboscis light brown, very short, vestigial, knob-like; labellum small, as wide as prementum, as long as prementum, unsclerotized laterally; maxillary palpus cylindrical, yellow, minute.
Antenna: brown, scape and pedicel white setose dorsally and ventrally; postpedicel cylindrical in proximal ½, symmetrically bulbous in distal ½, ≥ 4.0 times as long as combined length of scape and pedicel, asetose; apical seta-like sensory element situated apically in cavity on postpedicel.
Thorax: brown, lightly grey pubescent; scutum uniformly black or uniformly brown, surface entirely smooth, lightly grey pubescent, broad sublateral stripes apubescent, scutal setation comprised of scattered long white to yellow setae; dc setae pre- and postsuturally white or yellow, acr setae absent, lateral scutal setae white or yellow, npl setae 0, spal setae 0, pal setae 0; antepronotum dorso-medially with V-shaped indentation; postpronotal lobe light brown, grey pubescent; proepisternum, lateral postpronotum, and postpronotal lobe long white setose; scutellum grey pubescent, asetose, apical scutellar setae absent; mesopostnotum, anatergite, and katatergite partly grey pubescent, mesopostnotum apubescent, mesopostnotum asetose, anatergite asetose, katatergite long white setose; katatergite ± flat; anterior anepisternum white setose, supero-posterior anepisternum long white setose; posterior anepimeron asetose, katepimeron asetose; metanepisternum grey pubescent, asetose, metepimeron ± flat, yellow, grey pubescent, long white setose; infra-halter sclerite asetose or white setose.
Leg: light brown, setation yellow; pro, mes, and met coxa lightly white pubescent, short yellow setose; met trochanter setose medially; femur brown or light brown, met femur ± cylindrical only slightly wider than pro and mes femur, in distal ½ macrosetose, 1 antero-ventral and 1 postero-ventral row of macrosetae, postero-ventrally regular setose only; pro, mes, and met tibia straight, met tibia cylindrical, ventral keel absent, latero-posteriorly regular setose only; pro and mes tarsomere 1 longer than tarsomere 2, but less than combined length of tarsomeres 2–3, met tarsomere 1 as long as combined length of tarsomeres 2–4; pulvillus well-developed, as long as well-developed claw, and as wide as base of claw; empodium absent.
Wing: length = 6.6–8.1 mm; hyaline throughout, veins light brown, microtrichia absent; cells r1, r4, r5, m3, + cu
Abdomen: yellow to brown; setation comprised of dense white setae, surface entirely smooth; T1 brown, T2 brown anteriorly and postero-medially, otherwise yellow, T3–7 brown and yellow posteriorly; T1 and anterior ½ of T2 long white setose, remaining T short white setose; T predominantly apubescent; S1–7 light brown; S1 asetose, S2–7 sparsely white setose; S predominantly apubescent; T2–4 parallel-sided and not constricted waist-like; bullae on T2 black, transversely elongate, surface entirely smooth, T2 surface anterior to bullae smooth.
♂ terminalia: Fig.
Head: brown, facial gibbosity light brown; mystax white or yellow, sparsely covering entire facial gibbosity; setation: vertex white or yellow, pocl macrosetae light brown; maxillary palpus light brown.
Thorax: light brown, predominantly grey pubescent; scutum uniformly brown, predominantly brown pubescent, narrow sublateral stripes (wider anteriorly) and lateral and posterior margins grey pubescent, scutal setation comprised of scattered short yellow setae; proepisternum, lateral postpronotum, and postpronotal lobe short white setose; mesopostnotum, anatergite, and katatergite lightly grey pubescent, katatergite short white setose; anterior anepisternum white to yellow setose, supero-posterior anepisternum short white to yellow setose; metepimeron light brown; infra-halter sclerite asetose.
Leg: femur brown; pulvillus reduced, half length of well-developed claw.
Wing: length = 9.6–12.7 mm.
Abdomen: setation comprised of sparsely scattered short yellow setae; T1–7 brown, T2–6 light brown medially; T1–7 sparsely yellow setose; S1–7 brown; S1–7 sparsely short yellow setose; bullae on T2 oval.
♀ genitalia (Fig.
This rather small species (wing length in males 6.6–8.1 mm and in females 9.6–12.7 mm) is distinguished from congeners by the entirely grey pubescent vertex, the short postpedicel (only about 4 times as long as combined length of scape and pedicel), the grey pubescent scutellum, and the few white setae on the infra-halter sclerite, which are absent in some females.
Namibia (Hardap, Karas) and South Africa (Northern Cape) (Fig.
Namibia: Hardab: Excelsior No. 127 (
Not known to occur in any of the southern African biodiversity hotspots (Cape Floristic Region, Maputaland-Pondoland-Albany, or Succulent Karoo) (Fig.
Head: brown, facial gibbosity light brown, in general densely grey pubescent; width distinctly greater than thorax, interocular distance on vertex larger than at ventral eye margin, vertex between compound eyes ± horizontally straight, medially only slightly below dorsal eye margin, parafacial area about as wide as ½ the width of central facial gibbosity; facial gibbosity distinct, well-developed and discernible in lateral view; mystax white, densely covering entire facial gibbosity; frons not elevated, predominantly apubescent; vertex entirely white pubescent; postgena white pubescent; setation: vertex white, frons white, ocp setae white, pocl macrosetae white; ocellar triangle apubescent; proboscis yellow, very short, vestigial, knob-like; labellum small, as wide as prementum, as long as prementum, unsclerotized laterally; maxillary palpus cylindrical, yellow, minute.
Antenna: brown, scape and pedicel white setose dorsally and ventrally; postpedicel cylindrical in proximal ½, symmetrically bulbous in distal ½, ≥ 7.0 times as long as combined length of scape and pedicel, asetose; apical seta-like sensory element situated apically in cavity on postpedicel.
Thorax: brown, lightly grey pubescent; scutum predominantly black, anteriorly and laterally yellow to light brown, surface entirely smooth, lightly grey pubescent, scutal setation comprised of long white setae with distinct rows of long dorsocentral setae and dense lateral scutal setae; dc setae pre- and postsuturally white, acr setae absent, lateral scutal setae white, npl setae 0, spal setae 0, pal setae 0; antepronotum dorso-medially with V-shaped indentation; postpronotal lobe yellow, white pubescent; proepisternum, lateral postpronotum, and postpronotal lobe long white setose; scutellum apubescent, asetose, apical scutellar setae absent; mesopostnotum, anatergite, and katatergite grey pubescent, mesopostnotum asetose, anatergite asetose, katatergite long white setose; katatergite ± flat; anterior anepisternum white setose, supero-posterior anepisternum long white setose; posterior anepimeron asetose, katepimeron asetose; metanepisternum grey pubescent, asetose, metepimeron ± flat, yellow, grey pubescent, long white setose; infra-halter sclerite white setose.
Leg: yellow to light brown, setation predominantly white; pro, mes, and met coxa lightly white pubescent, long white setose; met trochanter setose medially; femur yellow to light brown, met femur ± cylindrical only slightly wider than pro and mes femur, in distal ½ macrosetose, 1 antero-ventral and 1 postero-ventral row of macrosetae, postero-ventrally long white, erect setose with setae arranged in distinct row; pro, mes, and met tibia straight, met tibia cylindrical, ventral keel absent, latero-posteriorly long white, erect setose with setae arranged in distinct row; pro and mes tarsomere 1 longer than tarsomere 2, but less than combined length of tarsomeres 2–3, met tarsomere 1 as long as combined length of tarsomeres 2–4; pulvillus well-developed, as long as well-developed claw, and as wide as base of claw; empodium absent.
Wing: length = 9.6–12.2 mm; hyaline throughout, veins light brown, microtrichia absent; cells r1, r4, r5, m3, + cu
Abdomen: yellow to brown; setation comprised of dense white setae, surface entirely smooth; T1–2 anteriorly yellow otherwise brown, T3 antero-medially brown otherwise yellow, T4–7 yellow to light brown; T1 and anterior ½ of T2 long white setose, remaining T short white setose; T predominantly apubescent; S1–7 yellow; S1–7 short white setose; S predominantly apubescent; T2–4 parallel-sided and not constricted waist-like; bullae on T2 black, transversely elongate, surface entirely smooth, T2 surface anterior to bullae smooth.
♂ terminalia: Fig.
This large species (wing length in males 9.6–12.2 mm, females unknown) is distinguished from congeners by the entirely white pubescent vertex and postgena, by the long postpedicel (about 7 times as long as combined length of scape and pedicel), the long white scutal setation, the yellow postpronotal lobes, the densely grey pubescent mesopostnotum, anatergite, and katatergite, the yellow to light brown coloration of the legs, and the long, erect white setae dorsally on the metathoracic femur.
Namibia (Karas) and South Africa (Northern Cape) (Fig.
South Africa: Northern Cape: Vioolsdrift (
Not known to occur in any of the southern African biodiversity hotspots (Cape Floristic Region, Maputaland-Pondoland-Albany, or Succulent Karoo) (Fig.
Head: brown, in general lightly silver pubescent; width distinctly greater than thorax, interocular distance on vertex larger than at ventral eye margin, vertex between compound eyes ± horizontally straight, medially only slightly below dorsal eye margin, parafacial area about as wide as ½ the width of central facial gibbosity; facial gibbosity distinct, well-developed and discernible in lateral view; mystax white, densely covering entire facial gibbosity; frons not elevated, predominantly apubescent; vertex predominantly apubescent, only lateral margin grey pubescent; postgena lightly grey pubescent; setation: vertex white, frons white, ocp setae white, pocl macrosetae absent; ocellar triangle apubescent; proboscis brown, short, about ½ length of oral cavity; labellum small, as wide as prementum, as long as prementum, unsclerotized laterally; maxillary palpus cylindrical, light brown, minute.
Antenna: brown, scape and pedicel white setose dorsally and ventrally; postpedicel cylindrical in proximal ½, symmetrically bulbous in distal ½, ≥ 7.0 times as long as combined length of scape and pedicel, asetose; apical seta-like sensory element situated apically in cavity on postpedicel.
Thorax: brown, lightly grey pubescent; scutum uniformly brown, surface entirely smooth, apubescent, scutal setation comprised of long white setae with distinct rows of long dorsocentral setae and dense lateral scutal setae; dc setae pre- and postsuturally white, acr setae absent, lateral scutal setae white, npl setae 0, spal setae 0, pal setae 0; antepronotum dorso-medially with V-shaped indentation; postpronotal lobe light brown, grey pubescent; proepisternum, lateral postpronotum, and postpronotal lobe long white setose; scutellum apubescent, asetose, apical scutellar setae absent; mesopostnotum, anatergite, and katatergite lightly grey pubescent, mesopostnotum asetose, anatergite long white setose, katatergite long white setose; katatergite ± flat; anterior anepisternum white setose, supero-posterior anepisternum long white setose; posterior anepimeron long white setose, katepimeron long white setose; metanepisternum grey pubescent, asetose, metepimeron ± flat, yellow, grey pubescent, long white setose; infra-halter sclerite white setose.
Leg: light brown to brown, setation predominantly white; pro, mes, and met coxa lightly white pubescent, long white setose; met trochanter setose medially; femur light brown to brown, met femur evenly clubbed in distal 3/4, in distal ½ macrosetose, 1 antero-ventral and 1 postero-ventral row of macrosetae, postero-ventrally long white, erect setose proximally with setae arranged in distinct row; pro, mes, and met tibia straight, met tibia cylindrical, ventral keel absent, latero-posteriorly long white, erect setose with setae arranged in distinct row; pro and mes tarsomere 1 longer than tarsomere 2, but less than combined length of tarsomeres 2–3, met tarsomere 1 as long as combined length of tarsomeres 2–4; pulvillus well-developed, as long as well-developed claw, and as wide as base of claw; empodium absent.
Wing: length = 7.1–8.9 mm; hyaline throughout, veins brown, microtrichia absent; cells r1, r4, r5, m3, + cu
Abdomen: yellow to brown; setation comprised of dense white setae, surface entirely smooth; T1 brown, T2 predominantly yellow with brown medially and antero-laterally, T3–7 yellow with brown antero-laterally; T1 and anterior ½ of T2 long white setose, remaining T short white setose; T predominantly apubescent; S1–7 light brown; S1–7 short white setose; S predominantly apubescent; T2–4 parallel-sided and not constricted waist-like; bullae on T2 black, transversely elongate, surface entirely smooth, T2 surface anterior to bullae smooth.
♂ terminalia: Fig.
Head: mystax white, covering entire facial gibbosity, sparse; pocl macrosetae white.
Antenna: postpedicel ≥ 5.0–≥ 6.0 times as long as combined length of scape and pedicel.
Thorax: scutum predominantly brown pubescent, narrow sublateral stripes (wider anteriorly) and lateral and posterior margins grey pubescent, scutal setation comprised of scattered short white setae; scutellum grey pubescent proximally, apubescent distally; supero-posterior anepisternum short white setose; posterior anepimeron short white setose, katepimeron short white setose; metepimeron light brown or yellow, grey pubescent, short white setose.
Leg: setation yellow; met femur ± cylindrical only slightly wider than pro and mes femur, postero-ventrally regular setose only; met tibia latero-posteriorly regular setose only; pulvillus reduced, half length of well-developed claw.
Wing: length = 10.9–14.2 mm; hyaline throughout, slightly brown stained along veins.
Abdomen: setation comprised of sparsely scattered short yellow setae; T1–5 brown with yellow posterior margin, T6 brown (sometimes yellow posteriorly), T7 brown; T1–7 sparsely yellow setose; S1–7 brown; S1–7 sparsely short yellow setose.
♀ genitalia: 8–9 acanthophorite spines per plate.
This large species (wing length in males 7.1–8.9 mm and in females 10.9–14.2 mm) is distinguished from congeners by the wing venation in that cell r5 is open and therefore M1 terminates in C (and not in R1), the predominantly apubescent vertex, the short proboscis that is only about half the length of the oral cavity, the long white setose anatergite, and the setose katepimeron.
Namibia (Erongo, Karas, Kunene) (Fig.
Females of this species (as
Females usually oviposited in shallow depressions, such as hoof prints and in particular on the lip of these prints, in the sandy Kuiseb river bed and followed a
This species exhibits substantial intra-specific variation (Figs
Namibia: Karas: Seeheim (
Not known to occur in any of the southern African biodiversity hotspots (Cape Floristic Region, Maputaland-Pondoland-Albany, or Succulent Karoo) (Fig.
1 | Anatergite setose; cell r5 open (M1+2 terminating into C); proboscis about ½ length of oral cavity; katepimeron setose |
|
– | Anatergite asetose; cell r5 closed (M1+2 terminating into R1); proboscis short, less than ½ length of oral cavity; katepimeron asetose |
|
2 | Infra-halter sclerite (ventral to halter base and posterior to metathoracic spiracle) with only very few white setae (Fig. |
|
– | Infra-halter sclerite long white setose (Fig. |
|
There is considerable sexual dimorphism and the setation, for example on the anatergites, is always easier to observe in males. Intraspecific variation in the abdominal coloration, especially in females, is likewise substantial.
The antepronotum is anteriorly not entire as in all
A small sclerite ventral to the halter and posterior to the metathoracic spiracle, here termed the infra-halter sclerite (Fig.
The knowlegde of the biology of
Although not directly observed by Wharton, he suggests that mating with teneral or very young females might occur in
These data provide a glimpse at the temperature tolerance within species as, for example,
The
We would like to thank the museum curators who made specimens available through loans and for their hospitality when visiting the collections. We thank Fritz Geller-Grimm (Frankfurt a.M., Germany) for donating the sole recently collected
TD conceptualized the project and gathered material from numerous collections. SL worked on this review during her NSF-funded summer internship at the Biodiversity Synthesis Center at the Field Museum of Natural History, Chicago, IL, USA where she was trained and supervised by TD in 2012.
lateral
dorsal
ventral
Distribution of
♂ (AAM-003021) dorsal (Morphbank #
♂ (AAM-003021) lateral (#
♀ (AAM-003000) dorsal (#
♀ (AAM-003000) lateral (#
Map of southern Africa with elevational relief and biodiversity hotspots (in grey) showing distribution of
dorsal (Morphbank #
lateral (#
lateral
dorsal
ventral
♂ (AAM-002988, head rotated 180˚) dorsal (Morphbank #
♂ (AAM-002988, head rotated 180˚) lateral (#
♀ (AAM-000456, Holotype) dorsal (#
♀ (AAM-000456, Holotype) lateral (#
lateral
dorsal
ventral
posterior
♀ holotype of
♀ (AAM-002924, #
♂ (AAM-002826, #
♂ (AAM-003046, #
Antepronotum and infra-halter sclerite of
Antepronotum with V-shaped indentation of
Sparsely setose infra-halter sclerite of
Densely setose infra-halter sclerite of
Seasonal incidence of
Month |
|
|
|
January | – | – | – |
February | – | √ | √ |
March | √ | √ | √ |
April | √ | – | √ |
May | √ | – | √ |
June | – | – | √ |
July | – | – | – |
August | – | – | – |
September | – | – | – |
October | – | √ | √ |
November | – | – | – |
December | – | – | – |
Month |
|
|
|
|||||||||||||
Maltahöhe | Upington | Aus | Keetmanshoop | Vioolsdrift | Gobabeb | Keetmanshoop | Khorixas | |||||||||
°C | coll. | °C | coll. | °C | coll. | °C | coll. | °C | coll. | °C | coll. | °C | coll. | °C | coll. | |
January | 34 | – | 37 | – | 24 | – | 36 | – | 20 | – | 30 | – | 36 | – | 31 | – |
February | 33 | – | 37 | – | 24 | ✓ | 35 | – | 20 | – | 30 | ✓ | 35 | – | 27 | – |
March | 31 | – | 34 | ✓ | 24 | – | 34 | – | 19 | ✓ | 32 | ✓ | 34 | – | 24 | – |
April | 28 | – | 30 | ✓ | 23 | – | 30 | – | 19 | – | 31 | ✓ | 30 | ✓ | 24 | ✓ |
May | 25 | ✓ | 26 | – | 22 | – | 26 | – | 20 | – | 29 | – | 26 | ✓ | 28 | ✓ |
June | 21 | – | 23 | – | 22 | – | 23 | – | 19 | – | 27 | ✓ | 23 | – | 26 | – |
July | 22 | – | 23 | – | 21 | – | 23 | – | 18 | – | 26 | – | 23 | – | 26 | – |
August | 24 | – | 25 | – | 19 | – | 25 | – | 17 | – | 24 | – | 25 | – | 25 | – |
September | 27 | – | 29 | – | 20 | – | 29 | – | 17 | – | 26 | – | 29 | – | 29 | – |
October | 30 | – | 32 | – | 21 | – | 32 | ✓ | 18 | – | 28 | – | 32 | – | 30 | – |
November | 32 | – | 34 | – | 22 | – | 34 | – | 18 | – | 30 | – | 34 | – | 32 | – |
December | 33 | – | 37 | – | 23 | – | 36 | – | 19 | – | 29 | – | 36 | – | 30 | – |
Natural-language species descriptions in SDD format
Data type: morphological
Brief description: The XML file includes the natural-language species descriptions in SDD (Structure of Descriptive Data) format.
File: oo_6075.sdd
Average annual temperature at Aus
Data type: image, graph
Brief description: Average temperature Aus
File: oo_4629.png
Average annual rainfall at Aus
Data type: image, graph
Brief description: Average rainfall Aus
File: oo_4630.png
Average annual temperature at Gobabeb
Data type: image, graph
Brief description: Average temperature Gobabeb
File: oo_4616.png
Average annual rainfall at Gobabeb
Data type: image, graph
Brief description: Average rainfall Gobabeb
File: oo_4617.png
Average annual temperature at Keetmanshoop
Data type: image, graph
Brief description: Average temperature Keetmanshoop
File: oo_4618.png
Average annual rainfall at Keetmanshoop
Data type: image, graph
Brief description: Average rainfall Keetmanshoop
File: oo_4619.png
Average annual temperature at Khorixas
Data type: image, graph
Brief description: Average temperature Khorixas
File: oo_4620.png
Average annual rainfall at Khorixas
Data type: image, graph
Brief description: Average rainfall Khorixas
File: oo_4621.png
Average annual temperature at Maltahöhe
Data type: image, graph
Brief description: Average temperature Maltahöhe
File: oo_4626.png
Average annual rainfall at Maltahöhe
Data type: image, graph
Brief description: Average rainfall Maltahöhe
File: oo_4627.png
Average annual temperature at Upington
Data type: image, graph
Brief description: Average temperature Upington
File: oo_4622.png
Average annual rainfall at Upington
Data type: image, graph
Brief description: Average rainfall Upington
File: oo_4623.png
Average annual temperature at Vioolsdrift
Data type: image, graph
Brief description: Average temperature Vioolsdrift
File: oo_4624.png
Average annual rainfall at Vioolsdrift
Data type: image, graph
Brief description: Average rainfall Vioolsdrift
File: oo_4625.png