Research Article |
Corresponding author: Ralph S. Peters ( r.peters@leibniz-zfmk.de ) Academic editor: Miles Zhang
© 2021 Christoph Braun, Ralph S. Peters.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Braun C, Peters RS (2021) Twelve new species of Dipara Walker, 1833 (Hymenoptera, Chalcidoidea, Pteromalidae, Diparinae) from Kenya, with a key to the Afrotropical species. ZooKeys 1067: 101-157. https://doi.org/10.3897/zookeys.1067.72395
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Based on 261 female specimens of the genus Dipara Walker, 1833 from leaf litter samples of the Kakamega Forest in Kenya, we describe the following twelve new species: Dipara andreabalzerae sp. nov., Dipara corona sp. nov., Dipara fastigata sp. nov., Dipara kakamegensis sp. nov., Dipara lux sp. nov., Dipara nigroscutellata sp. nov., Dipara nyani sp. nov., Dipara reticulata sp. nov., Dipara rodneymulleni sp. nov., Dipara sapphirus sp. nov., Dipara tenebra sp. nov., and Dipara tigrina sp. nov. For Dipara albomaculata (Hedqvist, 1963) and Dipara nigrita Hedqvist, 1969, we give new distribution records. We examined the available type material of all described Dipara species from the Afrotropical mainland, i.e., Dipara albomaculata (Hedqvist, 1963), Dipara machadoi (Hedqvist, 1971), Dipara maculata (Hedqvist, 1963), Dipara nigrita Hedqvist, 1969, Dipara pallida (Hedqvist, 1969), Dipara punctulata (Hedqvist, 1969), Dipara saetosa (Delucchi, 1962), Dipara straminea (Hedqvist, 1969), Dipara striata (Hedqvist, 1969), and Dipara turneri Hedqvist, 1969. We provide figures, descriptions, and diagnoses of the newly described species and figures and diagnoses of the ten known species as well as an identification key to all species of the Afrotropical mainland.
Morphometry, parasitoid wasps, taxonomy
In this study, we contribute to the taxonomic knowledge of the Afrotropical fauna of the genus Dipara Walker, 1833, with the first alpha-taxonomic treatment of this group and region in 50 years. We describe twelve new species and diagnose and key all new and previously described Afrotropical mainland species. Dipara belongs to the subfamily Diparinae within the chalcidoid family Pteromalidae (Heraty et al. 2012). The genus shows a cosmopolitan distribution (
The early taxonomic work on Dipara was confounded by the strong sexual dimorphism in this group. Males are usually macropterous and have filiform antennae. Females can range from macropterous to apterous and have clavate antennae. Additionally, males tend to be extremely similar even across different genera while females show a lot of interspecific morphological variation (
A peculiar characteristic of Diparinae females is their intraspecific variation in the wing form with macropterous and brachypterous specimens being found in the same species (
There is a severe lack of information about the biology of Dipara species. One of their main habitats is supposed to be leaf litter (
So far, ten species of Dipara have been described from the Afrotropical mainland, with a distribution ranging from the Democratic Republic of Congo to South Africa, including Dipara albomaculata (Hedqvist, 1963), Dipara machadoi (Hedqvist, 1971), Dipara maculata (Hedqvist, 1963), Dipara nigrita Hedqvist, 1969, Dipara pallida (Hedqvist, 1969), Dipara punctulata (Hedqvist, 1969), Dipara saetosa (Delucchi, 1962), Dipara straminea (Hedqvist, 1969), Dipara striata (Hedqvist, 1969) and Dipara turneri Hedqvist, 1969 (
We based our work on an extraordinary series of 261 female Dipara specimens from the Kakamega Forest in Kenya. Collection of the specimens was done in the framework of the BIOTA (BIOdiversity Monitoring Transect Analysis in Africa) East Africa project (
With the description of twelve new Dipara species from the leaf litter of Kakamega Forest in Kenya we can show that the species diversity of the genus has not been sufficiently studied and the true diversity of Afrotropical Dipara, and presumably other Diparinae, has been underestimated. Since our very much geographically limited study already more than doubles the number of known species, we expect that numerous additional species of Afrotropical Dipara still await discovery and description. This study may serve as a starting point for future in-depth investigations, including thorough taxonomic revisions of the Afrotropical Diparinae, Chalcidoidea or, more generally, parasitoid wasp fauna.
In the following, abbreviations are given of the museums where the material used in this study is stored. The abbreviations will be used throughout the text.
The terminology is based on
A total of 261 female Dipara specimens from the Kakamega Forest in Kenya were examined. They were collected in 2007 and 2008 using Winkler extraction of a 1 m² leaf litter sample in multiple transects throughout the Kakamega Forest (
Digital imaging was done with a Keyence VHX-2000 digital microscope. For images of the dorsal and lateral habitus and the head the VHX-J250 objective (250–2500×) was used. The images were stacked and edited in brightness, coloration and contrast using the Keyence internal software. Further editing of figures was done with Microsoft Power Point 2010. For the images for the morphometric measurements the Keyence VH-Z20R objective (20–200×) with a magnification of 200× was used. For the body length and the gaster length magnifications of 100× or 150× were used if the character did not fit into an image with 200× magnification. After calibration, measurements were done using ImageJ 1.53a. Characters used for morphometric measurements are given and explained in Table
List of morphometric characters with abbreviations and definitions (character definitions are based on
Abbr. | Character | Definition |
---|---|---|
scp.l | Scape length | Length of scape exclusive of radicle, outer aspect |
pdl.l | Pedicel length | Length of pedicel, outer aspect |
pdl.b | Pedicel breadth | Breadth of pedicel, outer aspect |
pdl.flg | Pedicel + flagellum | Combined length of pedicel plus flagellum, outer aspect |
clv.l | Clava length | Length of clava, outer aspect |
clv.b | Clava breadth | Breadth of clava, outer aspect |
tor.d | Toruli diameter | Greatest diameter of right torulus, outer aspect |
ant.d | Antennae distance | Greatest distance between outer edges of toruli |
ant.eye | Distance from antennal insertion to eye | Distance between center of insertion point of antennae and level of ventral margin of the eyes measured straight down from insertion point of antennae |
eye.b | Eye breadth | Greatest breadth of eye, lateral view |
eye.h | Eye height | Greatest length of eye height, lateral view |
mspl.l | Malar space length | Distance between the point where malar sulcus enters mouth margin and malar sulcus enters lower edge of eye, lateral view |
hea.h | Head height | Distance between lower edge of clypeus and lower edge of anterior ocellus, frontal view |
upf.l | Upper face length | Distance between lower edge of toruli and lower edge of anterior ocellus |
hea.b | Head breadth | Greatest breadth of head, dorsal view |
eye.d | Eye distance | Shortest distance between eyes, dorsal view |
pol.l | POL | Shortest distance between posterior ocelli, dorsal view |
ool.l | OOL | Shortest distance between posterior ocellus and eye margin, dorsal view |
prn.l | Pronotum length | Length of pronotum along median line from anterior edge of pronotum collar to anterior edge of mesoscutum |
prn.b | Pronotum breadth | Greatest breadth of pronotum, dorsal view |
msc.b | Mesoscutum breadth | Greatest breadth of mesoscutum just in front of level of tegula, dorsal view |
msc.l | Mesoscutum length | Length of mesoscutum along median line from posterior edge of pronotum to posterior edge of mesoscutum, dorsal view |
mss.l | Mesosoma length | Length of mesosoma along median line from anterior edge of pronotum collar to posterior edge of nucha, dorsal view |
sctl.l | Mesoscutellum length | Length of mesoscutellum along median line from posterior edge of mesoscutum to posterior edge of mesoscutellum, dorsal view |
ppd.l | Propodeum length | Length of propodeum measured along median line from anterior edge to posterior edge of nucha, dorsal view |
fm3.l | Metafemur length | Length of metafemur, from distal end of trochanter to tip of metafemur, measured along midline, outer aspect |
fm3.b | Metafemur breadth | Greatest breadth of metafemur, outer aspect |
ptl.l | Petiole length | Length of petiole measured along median line, from posterior edge of nucha to posterior edge of petiole, dorsal view |
ptl.b | Petiole breadth | Greatest breadth of petiole, outer aspect, dorsal view |
gst.l | Gaster length | Length of gaster along median line from posterior edge of nucha to tip of ovipositor sheath, dorsal view |
The range of the morphometric measurements is given in the species description with the value for the holotype in parentheses. If more than five specimens were present, five specimens were used for the morphometric measurements, and their respective collection numbers are given in parentheses at the beginning of the taxonomic treatment (see below). If five or less specimens were available, all specimens were used. In a few cases the number of specimens used for a certain measurement varies from the total number of specimens used, either because the measured character was not visible in some specimens or because more specimens were used for the in-depth morphometric analysis, using a subset of the characters (see below). In these cases, the collection numbers of specimens used are given in parentheses directly after the respective measurement (see Suppl. material
Shape categories for morphometric measurements and ratios in the species descriptions.
Character | Categories | |||
---|---|---|---|---|
Body length (in µm) | small | medium | large | |
< 2000 | 2000–3000 | > 3000 | ||
Head shape in frontal view (head breadth/head height) | round | oval | ||
< 1.31 | > 1.31 | |||
Mesosoma shape in dorsal view (head breadth/mesoscutum breadth) | robust | of medium breadth | slender | |
< 1.20 | 1.20–1.50 | > 1.50 | ||
Antennae distance (antennae distance/torulus diameter) | close | far apart | ||
< 1.33 | > 1.33 | |||
Distance from antennal insertion to eye (distance from antennal insertion to eye/torulus diameter) | same level as eyes | short | long | |
0 | < 1.1 | > 1.1 | ||
Pronotum shape in dorsal view (pronotum breadth/pronotum length) | large and elongated | of medium length | short and slim | |
< 2.5 | 2.5–3.5 | > 3.5 | ||
Petiole length (petiole length/petiole breadth) | short | medium | long | very long |
< 1.5 | 1.5–2.0 | 2.0–2.5 | > 2.5 | |
Gaster length (gaster length/mesosoma length) | short | medium | long | |
< 1.20 | 1.20–2.0 | > 2.0 |
Two putative species were found that clearly differed in the wing form, with one being macropterous and the other being brachypterous, but they were otherwise very similar with no obvious qualitative characters found to separate them. Since wing form might vary within species (
As suggested by
The morphometric analysis of specimens of the two morphologically similar putative species showed that they can be reliably separated (Fig.
Shape analysis of D. kakamegensis sp. nov. (circles) and D. nyani sp. nov. (triangles) using the characters highlighted in bold in Table
Based on the results of the scree graph (not shown), only the first and second principal component (PC) were relevant for the further analysis of shape. The results of the shape PCA (Fig.
The LDA ratio extractor (
The allometry ratio spectrum (Fig.
The separating ratios were used for the diagnoses of the two species in the descriptions below.
Examination of the posterior notal wing process (pnwp) in the newly described species showed that it was present and pointed in all cases. Figure
Dorsal view of a part of the mesosoma showing the posterior notal wing process (red) of A D. nyani sp. nov. (macropterous) B D. kakamegensis sp. nov. (brachypterous with medium sized wings), C D. andreabalzerae sp. nov. (brachypterous with medium sized wings) and D D. nigroscutellata sp. nov. (brachypterous with small wings). The former two are very similar but can be separated by morphometric analysis. Scale bar: 100 µm.
Dipara
Tricoryphus
Apterolelaps
Alloterra
Trimicrops
Parurios
Epilelaps
Pseudipara
Uriolelaps
Hispanolelaps
Pseudiparella
Emersonia
Grahamisia
Afrolelaps
Pondia
Diparomorpha
Female (taken from
The Diparinae genus key by
1 | Notauli present (Figs |
2 |
– | Notauli absent | D. machadoi (Hedqvist, 1971) |
2(1) | Petiole with thicker and longer bristle anterio-laterally, reaching gt1 (Figs |
3 |
– | Petiole with one or multiple thin and shorter setae laterally | 4 |
3(2) | Vertex and propodeum smooth (Fig. |
D. pallida (Hedqvist, 1969) |
– | Vertex reticulate; propodeum subcarinate (Fig. |
D. punctulata (Hedqvist, 1969) |
4(2) | Head and mesosoma never completely black, usually dark brown or lighter, sometimes bright yellowish brown; if head and mesosoma partly dark brown to black, then pro- and metacoxa white (Fig. |
5 |
– | Head and mesosoma black; coxae dark brown (Figs |
D. nigrita Hedqvist, 1969 |
5(4) | Face with one or two transverse dark brown to black stripes, reaching from one eye to the other, sometimes interrupted or fainter in interantennal and supraclypeal area (Figs |
6 |
– | Face without distinct dark stripes, uniformly colored or sometimes with diffuse darker coloration (Figs |
16 |
6(5) | Face with one transverse dark brown to black stripe, reaching from one eye to the other at the level of the ventral margin of the eyes (Figs |
7 |
– | Face with two transverse dark brown to black stripes at the level of toruli and at the level of the ventral margin of the eyes, enclosing a stripe of brighter coloration (Figs |
12 |
7(6) | Median and lateral area of mesoscutum with distinct transverse broad black stripe (Figs |
8 |
– | Lateral areas of mesoscutum with two black spots (Figs |
9 |
8(7) | Macropterous, fore wings reaching gt7; petiole distinctly longer than wide (Fig. |
D. corona sp. nov. |
– | Brachypterous, fore wings reaching slightly beyond petiole; petiole slightly wider than long (Fig. |
D. turneri Hedqvist, 1969 |
9(7) | Petiole short, < 1.5× as long as wide (Figs |
10 |
– | Petiole very long, > 2.5× as long as wide (Figs |
11 |
10(9) | Propodeum medially reticulated between carinae (Fig. |
D. tigrina sp. nov. |
– | Propodeum without reticulation between carinae; petiole with three pairs of small setae laterally (visible in dorsal view) (Fig. |
D. striata (Hedqvist, 1969) |
11(9) | Body brown to dark brown; vertex smooth (Fig. |
D. tenebra sp. nov. |
– | Body yellowish brown; vertex reticulate (Fig. |
D. lux sp. nov. |
12(6) | Gastral tergites smooth (Fig. |
13 |
– | Gastral tergites reticulate (Fig. |
D. reticulata sp. nov. |
13(12) | Lateral area of mesoscutum with two black spots; petiole long, < 2.1× as long as wide (Figs |
14 |
– | Mesoscutum without distinct black coloration; petiole very long, 2.53–2.80× as long as wide (Fig. |
D. rodneymulleni sp. nov. |
14(13) | Mesocoxa and petiole bright yellowish brown (Figs |
15 |
– | Mesocoxa and petiole white | D. maculata (Hedqvist, 1963) |
15(14) | Brachypterous, fore wing reaching middle of gt1; mesoscutellum smaller, mesosoma length 3.90–4.86× mesoscutellum length (Fig. |
D. kakamegensis sp. nov. |
– | Macropterous, fore wings reaching gt7; mesoscutellum larger, mesosoma length 3.43–3.83× mesoscutellum length (Fig. |
D. nyani sp. nov. |
16(5) | Mesoscutellum black (Figs |
17 |
– | Mesoscutellum not black (14C, 23C) | 21 |
17(16) | Propodeum medially smooth and laterally transversely carinate; gt1 without bristles (Figs |
18 |
– | Propodeum completely smooth; gt1 with a pair of large bristles dorso-anteriorly (Figs |
19 |
18(17) | Gaster brown to yellowish brown; anterior part of mesoscutellum and frenum forming an angle of 120–125° in lateral view (Fig. |
D. andreabalzerae sp. nov. |
– | Gaster dark brown; anterior part of mesoscutellum and frenum forming an angle of 90° in lateral view (Fig. |
D. fastigata sp. nov. |
19(17) | Body brown to dark brown; lateral area of mesoscutum completely black (Figs |
20 |
– | Body yellowish brown to brown; lateral area of mesoscutum not completely black with small yellowish brown area on its most lateral part (Fig. |
D. nigroscutellata sp. nov. |
20(19) | Vertex smooth; clava white; pro- and metacoxa white (Fig. |
D. albomaculata (Hedqvist, 1963) |
– | Vertex reticulate between ocelli, rest smooth; clava dark brown; pro- and metacoxa with proximal 1/3 brown and rest yellowish brown (Fig. |
D. saetosa (Delucchi, 1962) |
21(16) | Only slight metallic tint on black parts of the mesoscutum; lateral areas of mesoscutum with two black spots; gt1 with a pair of large bristles dorso-anteriorly (Fig. |
D. straminea (Hedqvist, 1969) |
– | Strong blue metallic tint on the following areas: vertex between ocelli, pronotum laterally, median area of mesoscutum posteriorly between notauli, lateral area of mesoscutum, mesoscutellum; lateral areas of mesoscutum without black spots; gt1 without a pair of large bristles dorso-anteriorly (Fig. |
D. sapphirus sp. nov. |
Holotype
Kenya • 1 ♀; Kakamega Forest, Kenya; 00°14'22.9N, 34°51'21E; 1594 m a.s.l.; 24 Jul. 2007; Hita-Garcia, F. leg.; Winkler extraction; Transect 12;
Female (specimens used for morphometric measurements:
Body brown to yellowish brown (Fig.
Size: small to medium sized, body length 1619–2183 (1809) µm.
Coloration: body brown to yellowish brown (Fig.
Head: head round, 1.26–1.31× (1.31) wider than high (Fig.
Mesosoma: pronotum large and elongated, 1.99–2.14× (2.02) as wide as long, substrigate, with a pair of setae close to the posterior edge (Fig.
Metasoma: petiole short, 1.25–1.44× (1.29) longer than wide in dorsal view, costate-rugose, with lateral pair of large white setae visible in dorsal view (Fig.
Dipara andreabalzerae is similar to D. albomaculata, D. fastigata, D. nigroscutellata, and D. saetosa in having a black mesoscutellum while the general body coloration is not black. Dipara andreabalzerae differs from D. albomaculata, D. nigroscutellata and D. saetosa in different propodeum sculpture. It differs from D. fastigata in body coloration, which is much darker in D. fastigata and the more obtuse angle formed by the anterior part of the mesoscutellum and the frenum in lateral view.
Unknown.
As the first author, I dedicate this species to my mother, Andrea Balzer, who sadly passed away in 2017.
Habitat: Leaf litter.
Host: Unknown.
Kenya.
Holotype Kenya • 1 ♀; Kakamega Forest, Kenya; 00°14'52.3N, 34°52'5.3E; 1607 m a.s.l.; 21 Aug. 2007; Hita-Garcia, F. leg.; Winkler extraction; Transect 18;
Female. Broad dark brown stripe across head from one eye to the other below toruli (Fig.
Size: medium sized, body length 2346 µm.
Coloration: body brown to orangish brown (Fig.
Head: head oval, 1.33× wider than high (Fig.
Mesosoma: pronotum of medium length, 3.01× wider than long, substrigate, with a pair of bristles medially close to the posterior edge (Fig.
Metasoma: petiole short, 1.20× longer than wide in dorsal view, anterior quarter constricted and rugose, rest costate, with lateral pair of large white setae visible in dorsal view (Fig.
Dipara corona is similar to D. turneri in having a distinct transverse broad black stripe on the median and lateral areas of the mesoscutum. In other not completely black species the black spots on the mesoscutum are restricted to the lateral area.
Dipara corona differs from D. turneri in the wing form and in the different petiole shape. The petiole is distinctly longer than wide in D. corona and slightly wider than long in D. turneri. Other differences include the body coloration, the shape of the mesoscutellum and the shape of the metacoxa.
Unknown.
Named after the Latin word corona for crown because of the raised and shiny part between the ocelli in frontal view, and additionally as a reference to the pandemic in 2020 and the following years caused by SARS-CoV-2, also known as the Corona virus.
Habitat: Leaf litter.
Host: Unknown.
Kenya.
Holotype
Kenya • 1 ♀; Kakamega Forest, Kenya; 00°14'6.1N, 34°52'9.2E; 1605 m a.s.l.; 28 Aug. 2007; Hita-Garcia, F. leg.; Winkler extraction; Transect 23;
Female. Body brown to dark brown (Fig.
Size: small sized, body length 1946 µm.
Coloration: body brown to dark brown (Fig.
Head: head round, 1.25× wider than high (Fig.
Mesosoma: pronotum large and elongated, 1.93× wider than long, with a transverse carina, anteriorly of carina substrigate and posteriorly smooth, with a pair of setae posterio-medially (Fig.
Metasoma: petiole medium, 1.6× longer than wide in dorsal view, reticulate, with two pairs of setae laterally visible in dorsal view (Fig.
Dipara fastigata is similar to D. andreabalzerae, D. albomaculata, D. nigroscutellata and D. saetosa in having a black mesoscutellum while the general body coloration is not black. Dipara fastigata differs from D. albomaculata, D. nigroscutellata and D. saetosa in different propodeum sculpture. It differs from D. andreabalzerae in general body coloration, which is much lighter in D. andreabalzerae, and the 90° angle formed by the anterior part of the mesoscutellum and the frenum in lateral view.
Unknown.
Named after the Latin adjective fastigatus for pointed or sharp. The name refers to the raised mesoscutellum.
Habitat: Leaf litter.
Host: Unknown.
Kenya.
Holotype
Kenya • 1 ♀; Kakamega Forest, Kenya; 00°21'4.9N, 34°51'41.1E; 1602 m a.s.l.; Hita-Garcia, F. leg.; Winkler extraction; Transect 1;
Female (specimens used for morphometric measurements:
Body bright yellowish brown (Fig.
Size: small to medium sized, body length 1483–2227 (2027) µm.
Coloration: body bright yellowish brown (Fig.
Head: head round to oval, 1.25–1.63× (1.63) wider than high (Fig.
Mesosoma: pronotum of medium length, 2.85–3.37× (3.37) as wide as long (Fig.
Metasoma: petiole short to medium, 1.15–1.72× (1.15) as long as wide in dorsal view, reticulate-rugose, with lateral pair of large white setae visible in dorsal view (Fig.
The bristles on the forewing can vary from five to 15. This variation is found in the bristles at the tip of the wing while along the edges there are constantly five bristles. In some specimens there are just a few larger bristles at the tip and in others there can be up to ten small bristles at the tip. The number of bristles can vary between left and right wing in one specimen. The surface sculpture of the median part of the propodeum can vary from rugose to smooth.
Dipara kakamegensis is very similar to D. nyani. It differs from D. nyani in the following characters: D. kakamegensis is brachypterous and the mesoscutellum is smaller relative to the mesosoma length, based on the morphometric analysis (Fig.
Unknown.
Named after the collecting locality.
Habitat: Leaf litter.
Host: Unknown.
Kenya.
Holotype
Kenya • 1 ♀; Kakamega Forest, Kenya; 00°18'13.4N, 34°48'16E; 1554 m a.s.l.; 20 Jun. 2007; Hita-Garcia, F. leg.; Winkler extraction; Transect 5;
Female. Body yellowish brown (Fig.
Size: medium sized, body length 2243–2772 (2772) µm.
Coloration: body yellowish brown (Fig.
Head: head oval, 1.31–1.34× (1.34) wider than high, reticulate except for interantennal area smooth (Fig.
Mesosoma: pronotum of medium length, 3.28–3.37× (3.37) as wide as long, reticulate, with two pairs of setae close to posterior edge (Fig.
Metasoma: petiole very long, 2.50–2.61× (2.50) longer than wide in dorsal view, areolate-rugose, with lateral pair of large white setae visible in dorsal view (Fig.
Dipara lux is similar to D. corona, D. machadoi, D. striata, D. tenebra, D. tigrina and D. turneri in having one dark brown to black stripe across the face. Dipara lux is different from D. machadoi in having distinct notauli, which are lacking in D. machadoi. It differs from D. corona, D. striata, D. turneri and D. tigrina in having a very long petiole. Dipara lux and D. tenebra are very similar in body shape and differ in their body coloration which is much brighter in D. lux and in the surface sculpture of the head. They share the otherwise unique character of having a dense brush of setae close to the proximal end of the marginal vein on the fore wing.
Unknown.
Named after the Latin word lux for light, in contrast to D. tenebra which looks very similar but has a darker coloration.
Habitat: Leaf litter.
Host: Unknown.
Kenya.
Holotype
Kenya • 1 ♀; Kakamega Forest, Kenya; 00°19'49.9N, 34°52'16.1E; 1580 m a.s.l.; 07 Aug. 2007; Hita-Garcia, F. leg.; Winkler extraction; Transect 15;
Female (specimens used for morphometric measurements:
Body yellowish brown to brown (Fig.
Size: small sized, body length 1653–2015 (1815) µm.
Coloration: body yellowish brown to brown (Fig.
Head: head round, 1.19–1.26× (1.21) wider than high (Fig.
Mesosoma: pronotum large and elongated, 1.48–1.65× (1.48) as wide as long, strigate, with two or three pairs of setae laterally close to the posterior edge (Fig.
Metasoma: petiole short, 0.98–1.16× (1.06) as long as wide, costate-rugose, with lateral pair of large white setae visible in dorsal view (Fig.
The bristles on the gt1 and the tip of the forewing can sometimes be missing. In this case the pit where the bristles are supposed to be is still visible.
Dipara nigroscutellata is similar to D. andreabalzerae, D. albomaculata, D. fastigata, and D. saetosa in having a black mesoscutellum while the general body coloration is not black. Dipara nigroscutellata differs from D. andreabalzerae and D. fastigata in having a pair of large bristles dorso-anteriorly on the gt1. It differs from D. albomaculata and D. saetosa in the general body coloration, which is much brighter in D. nigroscutellata and in the coloration of the lateral area of the mesoscutum. In D. nigroscutellata the lateral area is laterally yellowish brown and in D. albomaculata and D. saetosa the lateral area is completely black.
Dipara nigroscutellata is similar to D. straminea in sharing the bristles on the gt1 and in propodeum sculpture. It differs from D. straminea in having a black mesoscutellum.
Unknown.
Named after the black mesoscutellum.
Habitat: Leaf litter.
Host: Unknown.
Kenya.
Holotype
Kenya • 1 ♀; Kakamega Forest, Kenya; 00°19'45.7N, 34°52'2.8E; 1573 m a.s.l.; 17 Aug. 2007; Hita-Garcia, F. leg.; Winkler extraction; Transect 21;
Female. Body bright yellowish brown (Fig.
Size: small to medium sized, body length 1696–2064 (2037) µm (specimens used for measurement:
Coloration: body bright yellowish brown (Fig.
Head: head round, 1.25–1.30× (1.30) wider than high (Fig.
Mesosoma: pronotum short and slim, 3.84–5.79× (5.79) as wide as long, reticulate (Fig.
Metasoma: petiole medium to long, 1.78–2.05× (2.02) (specimens used for measurement:
Dipara nyani is very similar to D. kakamegensis. It differs from it in the following characters: D. nyani is macropterous and the mesoscutellum is larger relative to the mesosoma length, based on the results of the morphometric analysis (Fig.
Unknown.
Named after the word for monkey in the national language of Kenya, Swahili, because of the dorsal black dots and the mesoscutellum which resemble the face of a monkey.
Habitat: Leaf litter.
Host: Unknown.
Kenya.
Holotype
Kenya • 1 ♀; Kakamega Forest, Kenya; 00°14'6.1N, 34°52'9.2E; 1605 m a.s.l.; 28 Aug. 2007; Hita-Garcia, F. leg.; Winkler extraction; Transect 23;
Female. Gastral tergites reticulate (Fig.
Size: medium sized, body length 2303–2927 (2303) µm.
Coloration: vertex and upper face brown to orangish brown, lower face yellowish brown (Fig.
Head: head oval, 1.35–1.42× (1.35) wider than high (Fig.
Mesosoma: pronotum large and elongated, 1.78–2.38× (1.78) wider than long, substrigate, with two transverse rows of setae on posterior 1/2 (Fig.
Metasoma: petiole short, 1.20–1.31× (1.24) wider than long, rugose, with four pairs small white setae laterally (Fig.
Dipara reticulata is similar to D. kakamegensis, D. maculata, D. nyani, and D. rodneymulleni in having transverse stripes across the face. Dipara reticulata is different form all other Dipara species in having reticulated gastral tergites. In all other species the gastral tergites are smooth.
Unknown.
Named for the reticulated gastral tergites.
Habitat: Leaf litter.
Host: Unknown.
Kenya.
Holotype
Kenya • 1 ♀; Kakamega Forest, Kenya; 00°19'36N, 34°52'14.6E; 1570 m a.s.l.; 28 Jun. 2007; Hita-Garcia, F. leg.; Winkler extraction; Transect 6;
Female. Face with two dark brown stripes at the level of the ventral margin of the eye, interrupted in supraclypeal area, and at the level of the toruli (Fig.
Size: medium to large, body length 2718–3397 (3397) µm.
Coloration: body brown (Fig.
Head: head round, 1.22–1.29× (1.29) (specimens used for measurement:
Mesosoma: pronotum short and slim, 3.29–3.81× (3.74) as wide as long, strigulate-reticulate, sparsely setose (Fig.
Metasoma: petiole very long, 2.53–2.80× (2.79) longer than wide, with anterior 2/3 rugose and rest carinate, with lateral pair of large white setae visible in dorsal view (Fig.
Dipara rodneymulleni shares the stripes across the face with D. maculata, D. nyani, D. kakamegensis, and D. reticulata and but other than that has a completely different morphology and coloration. The most obvious characters to distinguish D. rodneymulleni are the very long petiole and the absence of black spots on the lateral areas of the mesoscutum.
Male. Unknown.
Named after professional skateboarder Rodney Mullen who revolutionized street skating like no other, reflecting the first author’s lifelong passion for skateboarding.
Habitat: Leaf litter.
Host: Unknown.
Kenya.
Holotype
Kenya • 1 ♀; Kakamega Forest, Kenya; 00°13'15.5N, 34°53'24.7E; 1597 m a.s.l.; 23 Aug. 2007; Hita-Garcia, F. leg.; Winkler extraction; Transect 22;
Female (specimens used for morphometric measurements:
Strong blue metallic tint on the following areas: vertex between ocelli, pronotum laterally, median area of mesoscutum posteriorly between notauli, lateral area of mesoscutum and mesoscutellum (Fig.
Size: small to medium sized, body length 1667–2432 (2081) µm.
Coloration: body dark brown (Fig.
Head: head round, 1.26–1.30× (1.29) wider than high (Fig.
Mesosoma: pronotum mostly short and slim, 3.30–3.60× (3.60) wider than long, medially and around posterior margin smooth, laterally reticulate, with some setae close to the posterior edge (Fig.
Metasoma: petiole short to medium, 1.31–1.67× (1.60) longer than wide in dorsal view, costate-rugose, with lateral pair of large white setae visible in dorsal view (Fig.
In body shape, D. sapphirus is similar to D. lux and D. tenebra but can be distinguished from all other Dipara species by having a very distinct blue metallic tint on the following body parts: vertex between ocelli, pronotum laterally, median area of mesoscutum posteriorly between notauli, lateral area of mesoscutum and mesoscutellum (Fig.
Unknown.
Named after sapphires for the blue metallic tint.
Habitat: Leaf litter.
Host: Unknown.
Kenya.
Holotype
Kenya • 1 ♀; Kakamega Forest, Kenya; 00°27'10.6N, 34°51'48.7E; 1676 m a.s.l.; 19 Jun. 2007; Hita-Garcia, F. leg.; Winkler extraction; Transect 4;
Female. Body brown to dark brown (Fig.
Size: medium sized, body length 2293–2474 (2293) µm.
Coloration: body brown to dark brown (Fig.
Head: head oval, 1.33–1.37× (1.33) wider than high (Fig.
Mesosoma: pronotum of medium length, 3.11–3.12× (3.11) wider than long, substrigate, with row of setae close to the posterior edge (Fig.
Metasoma: petiole very long, 2.51–2.77× (2.51) longer than wide in dorsal view, costate-rugose, with lateral pair of large setae visible in dorsal view (Fig.
Dipara tenebra is similar to D. corona, D. lux, D. machadoi, D. striata, D. tigrina, and D. turneri and in having one dark brown to black stripe across the face. Dipara tenebra is different from D. machadoi in having distinct notauli, which are lacking in D. machadoi. It differs from D. corona, D. striata, D. tigrina, and D. turneri in having a very long petiole. Dipara tenebra and D. lux are very similar in body shape and differ in their body coloration which is much darker in D. tenebra and in the surface sculpture of the head. They share the otherwise unique character of having a dense brush of setae close to the proximal end of the marginal vein on the fore wing.
Unknown.
Named after the Latin word tenebra for darkness, in contrast to D. lux which looks very similar but is much lighter in coloration.
Habitat: Leaf litter.
Host: Unknown.
Kenya.
Holotype
Kenya • 1 ♀; Kakamega Forest, Kenya; 00°21'4.4N, 34°51'41.1E; 1602 m a.s.l.; 05 Jun. 2007; Hita-Garcia, F. leg.; Winkler extraction; Transect 2;
Diagnosis. Female. Propodeum laterally smooth, medially distinctly subcarinate with reticulation between carinae, carinae extending to nucha (Fig.
Size: medium sized, body length 2329 µm.
Coloration: body yellowish brown to brown (Fig.
Head: head round, 1.24× wider than high, entirely reticulate (Fig.
Mesosoma: pronotum large and elongated, 1.95× as wide as long, reticulate, with two rows of small setae close to posterior margin (Fig.
Metasoma: petiole short, 1.37× as long as wide in dorsal view, with at least six pairs of small white setae laterally visible in dorsal view, subcarinate (Fig.
Dipara tigrina is similar to D. corona, D. lux, D. machadoi, D. striata, D. tenebra, and D. turneri in having one dark brown to black stripe across the face. It differs from D. corona, D. lux, D. machadoi, D. tenebra, and D. turneri in the propodeum sculpture. The propodeum sculpture is similar in D. punctulata and D. striata. They show a very distinct surface sculpture with a striated subcarinate pattern extending to the nucha. Dipara tigrina differs from D. punctulata and D. striata in having more setae laterally on the propodeum and in having a reticulated pattern medially between the carinae on the propodeum.
The only available specimen of this species has an irregular black spot on the propodeum. This spot is considered an aberration and thus is not part of the species description.
Male. Unknown.
Named after the Latin adjective tigrinus for the tiger-like stripes on the gaster.
Habitat: Leaf litter.
Host: Unknown.
Kenya.
Afrolelaps albomaculata
Grahamisia albomaculata
Dipara albomaculata
Paratype Angola • 1 ♀; Mabete, Caungula; 20. Jul. 1962; A. de Barros Machado leg.; NHMUK013455574.
Kenya • 5 ♀; Kakamega Forest, Kenya; 00°22'43.7N, 34°41'57.3E; 1452 m a.s.l.; 25 Aug. 2008; Hita-Garcia, F. leg.; Winkler extraction; Transect 35;
Female. Body brown to dark brown (Fig.
The holotype of D. albomaculata is supposed to be stored at the
Dipara albomaculata is similar to D. andreabalzerae, D. fastigata, D. nigroscutellata and D. saetosa in having a black mesoscutellum while the general body coloration is not black. It differs from D. andreabalzerae and D. fastigata in having a pair of bristles dorso-anteriorly on the gt1. It differs from D. nigroscutellata in the general body coloration, which is much darker and in the coloration of the lateral area of the mesoscutum. In D. albomaculata the lateral area is completely black and D. nigroscutellata has a small yellowish brown area on its most lateral part. The differences to D. saetosa can be found in the smooth vertex and the white pro- and metacoxa.
Dipara albomaculata is similar to D. straminea in sharing the bristles on the gt1 and the propodeum sculpture. It differs from D. straminea in having a black mesoscutellum.
Additional specimens from this species were found in the Kakamega Forest in Kenya and the distribution is updated accordingly.
Angola; Kenya.
Diparomorpha machadoi
Dipara machadoi
Female. Notauli absent.
The holotype of D. machadoi is supposed to be stored at the
Afrolelaps maculata
Grahamisia maculata
Dipara maculata
Female. Face with two transverse stripes of dark brown coloration just at the level of toruli and at the level of the ventral margin of the eye, enclosing a stripe of pale yellowish white coloration; mesocoxa and petiole white.
The holotype of D. maculata is supposed to be stored at the
Dipara nigrita Hedqvist, 1969: 195.
Holotype
Democratic Republic Of Congo • 1 ♀; Mount Kabobo, Terr. Albertville, Hte. Kiymbi; 1700 m a.s.l; Oct. 1958; N. Leleup leg.; “Humus en forêt”;
Kenya • 1 ♀; Kakamega Forest, Kenya; 00°14'22.9N, 34°51'21E; 1594 m a.s.l.; 24 Jul. 2007; Hita-Garcia, F. leg.; Winkler extraction; Transect 12;
Female. Head and mesosoma black, coxae dark brown (Figs
Dipara nigrita was originally described as brachypterous (Fig.
Macropterous individuals have fully developed wings with the fore wings reaching the gt7 (Fig.
Dipara nigrita is the only species which shows a completely black coloration of the head and mesosoma. Darker specimens of D. albomaculata sometimes have a partly very dark brown to black head and mesosoma but never completely black. Additionally, the coxa of D. albomaculata are white in contrast to the dark brown coxa of D. nigrita.
Additional specimens from the species were found in the Kakamega Forest in Kenya and the distribution is updated accordingly.
Democratic Republic of Congo; Kenya.
Pondia pallida
Dipara pallida
Holotype South Africa • 1 ♀; Port St. John, Pondoland; Jan. 1924; R.E. Turner leg.; NHMUK013455580.
Dipara pallida is similar to D. punctulata in having a large bristle anterio-laterally on the petiole. They differ in the surface sculpture of the vertex and the propodeum.
Pondia punctulata
Dipara punctulata
Holotype South Africa • 1 ♀; Port St. John, Pondoland; Jan. 1924; R.E. Turner leg.; NHMUK013455579.
Female. Vertex reticulate (Fig.
Dipara punctulata is similar to D. pallida in having a large bristle anterio-laterally on the petiole. They differ in the surface sculpture of the vertex and the propodeum.
Grahamisia saetosa
Dipara saetosa
Holotype
Tanzania • 1 ♀; Tanganyika Terr., Mt. Oldeani, versant Est; 2350–2500 m a.s.l.; 6.–9. Jun. 1957;
Female. Vertex reticulate between ocelli, rest smooth (Fig.
Dipara saetosa is similar to D. albomaculata, D. nigroscutellata, and D. straminea in having a pair of large bristles dorso-anteriorly on the gt1. It differs from D. straminea in having a black mesoscutellum. In contrast to D. nigroscutellata the lateral area of the mesoscutum is completely black. Differences to D. albomaculata can be found in the reticulation between the ocelli and the coloration of the clava and the pro- and metacoxa.
Grahamisia straminea
Dipara straminea
Holotype
Democratic Republic Of Congo • 1 ♀; Kivu, Terr. Mwenga, S.-O. Tombwe, Luiko; 2100 m a.s.l.; Jan. 1952; N. Leleup leg.; “Récolté dans l’humus”;
Female. Mesoscutellum yellowish brown (Fig.
Dipara straminea is similar to D. albomaculata, D. nigroscutellata, and D. saetosa in having a pair of large bristles dorso-anteriorly on the gt1. It differs from them by having a yellowish brown mesoscutellum.
Grahamisia striata
Dipara striata
Holotype South Africa • 1 ♀; Cape Province, Somerset East; 1.–26. Jan. 1931; R.E. Turner leg.; NHMUK013455578.
Dipara striata is similar to D. corona, D. lux, D. machadoi, D. tenebra, D. tigrina, and D. turneri in having one dark brown to black stripe across the face. It differs from D. corona, D. lux, D. machadoi, D. tenebra, and D. turneri in the propodeum sculpture. The propodeum sculpture is similar in D. punctulata and D. tigrina. They show a very distinct striated subcarinate pattern extending to the nucha. Dipara striata differs from D. punctulata in lacking a large bristle anterio-laterally on the petiole. Dipara striata differs from D. tigrina in having less setae laterally on the petiole and in lacking reticulation on the propodeum.
Dipara turneri
Holotype South Africa • 1 ♀; Port St. John, Pondoland; 6.–25. Feb. 1924; R.E. Turner leg.; NHMUK013455576.
Diagnosis. Female. Broad dark brown stripe across head from one eye to the other below toruli (Fig.
Dipara turneri is similar to D. corona in having a distinct transverse broad black stripe on the median and lateral area of the mesoscutum. In other not completely black species the black spots on the mesoscutum are restricted to the lateral area.
Dipara turneri differs from D. corona in the wing form and in the petiole shape. The petiole is longer than wide in D. corona and wider than long in D. turneri.
Our results confirm that there is still a lot of undiscovered diversity within Microhymenoptera and the genus Dipara in particular (
Some of the species descriptions in this study are based on so far unparalleled series of Dipara specimens. While most of the previously described Dipara species are known only from the holotype or just a few specimens, D. kakamegensis, for example, is described from 108 specimens and D. nigroscutellata from 86 specimens. These large series allowed for an advanced insight into intraspecific variation of Dipara species. The characters used for the species descriptions and diagnoses were found to be consistent among the large series, which gave us some confidence in delimiting species using the same characters in species with less specimens available. Our insights reveal that in most cases Afrotropical Dipara species seem to be reliably distinguishable by comparatively simple morphological characters of females like color patterns, surface sculpture or the number and position of setae or bristles. However, the intraspecific variation of the wing form of Diparinae females (
While shedding more light on the species diversity of Dipara their biology remains largely unknown. All specimens were found in the leaf litter confirming that this might be their preferred habitat (
Adding information on the biology, taxonomy, and distribution of species, is a crucial task. We still have only very limited knowledge on the biodiversity on this planet. We are aware, though, that we are facing presumably unprecedented biodiversity loss, especially through habitat destruction, and that this is one of the most pressing problems of our time (
Thanks to Hannes Baur for support with the morphmetric analysis and to Mircea-Dan Mitroiu for valuable information on Diparinae. We thank Natalie Dale-Skey (
Table S1
Data type: Xslx file.
Explanation note: Measurements for the morphometric analysis of Dipara kakamegensis sp. nov. and Dipara nyani sp. nov. See Table
Table S2
Data type: Xslx file.
Explanation note: Morphometric measurements for the descriptions of the newly described Dipara species. See Table
Table S3
Data type: Xslx file.
Explanation note: Shape ratios for the descriptions of the newly described Dipara species. See Table