Corresponding author: Simon van Noort (
Academic editor: M. Yoder
The liopterid subfamily
The liopterid subfamily
Biologically the family is still poorly known with knowledge restricted to a couple of published records of association through rearing: two species of
During visits to European museums to image types of Afrotropical
Specimens were point mounted on black, acid-free card for examination (using a Wild M-5 stereomicroscope with incandescent and fluorescent light sources), photography and long term preservation. Images were acquired using the EntoVision® multiple-focus imaging system. This system comprises a Leica® M16 microscope with a JVC® KY-75U 3-CCD digital video camera attached that fed image data to a notebook computer. The program Cartograph® 5.6.0 was then used to merge an image series (representing typically 10-15 focal planes) into a single in-focus image. Lighting was achieved using techniques summarized in
Morphological terminology follows that of
Identification keys were produced in three formats to facilitate accessibility by a range of end-users (
Natural History Museum, London. Curator: David Notton
Natural History Museum, Paris. Curator: Claire Villemant
National Museums of Kenya, Nairobi. Curator: Martha Gikunga
Robert Copeland personal collection, Nairobi.
Royal Museum for Central Africa, Tervuren. Curator: Eliane de Coninck
Iziko South African Museum, Cape Town. Curator: Simon van Noort
South African National Collection of Insects. Curator: Ros Urban.
National Museum of Natural History, Washington DC. Curator: Matt Buffington.
The
This subfamily includes two genera,
Medium-sized to very small cynipoids. Very small species look superficially like cynipids, but careful attention to the relative size of the metasomal terga will help separate
Online interactive keys to genera of Afrotropical
The genus is represented in all biogeographical regions except for the Western Palaearctic and Australia (
The type female of
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1 | Mesoscutum smooth, shiny with only remnants of transverse costae. Median mesoscutal impression present, reaching halfway to anterior margin. The two scutellar foveae each with four subcarinae creating a transverse row of 10 longitudinally elongate subfoveae. Latero-ventral margin of pronotum angled where it meets lateral pronotal carina. F1 of antenna shorter than F2 | |
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– | Mesoscutum foveate-reticulate, or transversely carinate with foveae set in rows, or with rough discontinuous transverse costae produced into irregularly raised and slightly backward pointing teeth. Median mesoscutal impression, if present, obscured by sculpture. Scutellar foveae usually two in number, or less frequently subdivided into a maximum of 6 subfoveae. Latero-ventral margin of pronotum evenly curved. F1 of antenna equal to or longer than F2 | 2 |
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2 | Pronotal crest gradually raised, medially forming a conspicuous slightly backward pointing ridge-like tooth. Mesoscutum with rough discontinuous transverse costae produced into irregularly raised and slightly backward pointing teeth. Speculum longitudinally costate. Median propodeal area not delimited by lateral propodeal carinae, posteriorly foveate-reticulate | |
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– | Pronotal crest flat. Mesoscutum foveate-reticulate or with continuous transverse carinae with foveae set in rows. Speculum smooth, rarely longitudinally costate. Median propodeal area distinctly delimited by lateral propodeal carinae, posteriorly not foveate-reticulate |
We erect this new species-group to accommodate a single species that is likely to be a Madagascan endemic. The island is known for its high degree of endemism (
Behara (
Madagascar (currently only known from Behara).
Named after André Seyrig (1897–1945) collector of the type series and prolific collector of many other faunal and floral taxa from Madagascar. Noun in the genitive case.
Belongs to the newly erected
This species, along with
FEMALE. Length 5.2–8.0 mm (holotype: 6.3 mm ). Head, antennae (except for terminal segments which are darker), pronotum, legs and proximal third of metasoma brownish-orange; rest of mesosoma dark brown; rest of metasoma orange (
Metanotal-propodeal complex strongly excavated, excavations bordered by strong carinae. Metepisternum dorsally excavated with pubescence, medially polished with indications of minor rugose remnants, ventrally pubescent. Dorsellum with two strong medial foveae; laterally strongly excavated with fine pubescence in lateral depressions. Lateral propodeal carina present; median longitudinal propodeal carina well-defined, crossed by wrinkled transverse and longitudinal carinae extending onto nucha (
MALE. Length 4–5.5 mm. Very similar to female except for abdominal petiole, which is much more elongate, T2 as long as wide in dorsal view and twice as long as high in lateral view. Tergites 6–8 each with a dorsal terminal area of setiferous punctures, rest of tergite polished.
Historically this taxon has been recognized variously at generic (
Previously the species-group was only known from the African mainland with three described species (
This species–group is characterized in females by excavations (spiracular peritremata) on the terminal portion of T8 associated with the spiracle (
1 | Forewing with RS+M vein arising at or very close to base of basal vein ( |
2 |
– | Forewing with RS+M vein arising at basal third or mid-way up basal vein ( |
3 |
2 | Head and mesosoma black ( |
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– | Head pronotum, mesoscutum and scutellum reddish-brown ( |
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3 | Body length about 4 mm. Body colour entirely dark ( |
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– | Body length about 6–10 mm. Head and mesosoma dark, metasoma yellow to brown ( |
4 |
4 | Face evenly curved in lateral view. Genae ventrally strongly expanded posteriorly. Median mesopleural impression distinct. Lower mesopleuron densely punctate with pubescence. Apical teeth of metatibia rounded apically; first metatarsomere 2.0× length of remaining 4 metatarsomeres combined |
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– | Face distinctly raised medially and curved inward ventrally in lateral view ( |
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Cameroon.
Madagascar.
Named after the type locality. Noun in apposition.
Belongs to the
There is the possibility that this species may be the male of, and hence conspecific, with
See note under
MALE. Length 2.7 mm. Head and mesosoma black; metasoma reddish-brown; antennae (except for three terminal segments which are darker), are orangish-brown (
Mesosoma dorsally with scattered long pubescent. Anterior flange and plate of pronotum uniformly areolete-rugulose and pubescent. Pronotum dorsomedially distinctly raised, in lateral view distinctly higher than anterior margin of mesoscutum (
Rs+M of forewing nebulous, arising from the base of basal vein (
FEMALE. Unknown.
Kenya.
Democratic Republic of Congo, Uganda.
Belongs to the
This species along with
See note under
FEMALE. Length 4.5 mm. Head, antennae (except for terminal segments which are darker), mesosoma (except for propodeum and mesopleuron which are black) reddish-brown; legs and metasoma dark brown. Wings clear (
Mesosoma dorsally with scattered long pubescent (
Rs+M of forewing defined, arising from the base of basal vein (
MALE. Unknown.
1 | Head compressed longitudinally; occiput not concave in dorsal view. Mesoscutum densely foveate, without transverse carinae |
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– | Head not compressed longitudinally; occiput distinctly concave in dorsal view. Mesoscutum more or less foveate-reticulate, with or without transverse carinae | 2 |
2 | Median frontal carina absent. T6 of female metasoma the largest and T8 distinctly exposed. Median propodeal area without a strong transverse carina | 3 |
– | Median frontal carina present (may only be weakly represented between toruli). T6 of female metasoma not always the largest; if T6 the largest, then T8 is not exposed. Median propodeal area usually with a strong transverse carina. Occasionally variations occur, but never come in combination of features as the above collate | 7 |
3 | Forewing entirely clear | 4 |
– | Forewing at least ferruginous in marginal cell | 5 |
4 | Antennae of female with 11 flagellomeres; apical flagellomere less than twice as long as subapical flagellomere. Pronotal crest medially not raised into a triangular process. Metasoma black |
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– | Antennae of female with 10 flagellomeres; apical flagellomere longer than twice as long as subapical flagellomere. Pronotal crest medially raised into a small but distinct triangular process. Metasoma brown |
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5 | Antennal scrobes longitudinally carinate in upper part and glabrous in lower part. Upper mesopleuron glabrous, smooth, shiny. Metasoma black |
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– | Antennal scrobes longitudinally carinate entirely. Upper mesopleuron foveate to rugose. Metasoma brown | 6 |
6 | Vertex longitudinally carinate laterally. Pronotal crest medially raised into a small, distinct rounded triangular process. Scutellar foveae without submedian carinae |
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– | Vertex foveate-reticulate entirely without longitudinal carination. Pronotal crest smoothly flat, without triangular process. Scutellar foveae subdivided by distinct submedian carinae |
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7 | Distance between posterior ocelli at most twice as large as the distance between posterior ocellus and eye. Metasomal T5 of female normal, T6 the largest; T7 exposed, almost entirely covering T8 | 8 |
– | Distance between posterior ocelli at least three times the distance between posterior ocellus and eye. Metasomal T5 of female dorsally expanded, being the largest (at least so dorsally); T7 is largely or entirely covered by T6; T8 exposed | 22 |
8 | Female flagellum distinctly thicker toward apex; median flagellomeres not or slightly constricted toward ends. Antennae with distal flagellomeres 1–3 black, contrasting to the rest, which are yellow-orange | 9 |
– | Female flagellum not distinctly thicker toward apex; median flagellomeres distinctly constricted toward ends. Antennae yellow or gradually becoming somewhat darker toward apex, but never with contrasting colors between distal and proximal flagellomeres | 13 |
9 | Wings with infuscate patch covering submarginal and marginal cells and extending distally slightly beyond vein RS ( |
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– | Wings clear | 10 |
10 | Metacoxae ventrally expanded to form a triangular lobular process |
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– | Metacoxae ventrally not expanded to form a triangular lobular process | 11 |
11 | Antennae with apical 3 flagellomeres black ( |
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– | Antennae with either the apical or apical 2 flagellomeres dark-brown to black. Antennal scrobes heavily and densely punctate with longitudinal carinae posteriorly | 12 |
12 | Female flagellum with terminal two segments usually dark brown; T9 without dense brush of setae |
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– | Female flagellum usually with only terminal segment darkened (occasionally blending into penultimate segment); female T9 with a distinct brush of setae present, engulfing ovipositor ( |
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13 | Dorsal surface of head between toruli and posterior margin of lateral ocelli mostly smooth, shiny with lateral carina of antennal scrobe bound on both sides by smooth areas, and sub-confluent with genal carina ( |
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– | Dorsal surface of head between toruli and posterior margin of lateral ocelli sculptured or with diagonal carina, except for antennal scrobes, which may be smooth; lateral carina of antennal scrobe interrupted on vertex and not joining genal carina | 14 |
14 | Ocellar plate of head not defined by lateral carinae, and without a small, triangular glabrous area beneath anterior ocellus |
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– | Ocellar plate of head well defined by lateral carinae, with a small, triangular glabrous area beneath anterior ocellus | 15 |
15 | Vertex with distinct longitudinal carination | 16 |
– | Vertex without distinct longitudinal carination | 20 |
16 | Median frontal carina almost extending to clypeus. Ocellar plate either with a row of relatively uniform, large foveae along the lateral carinae delimiting the plate or delimited with two parallel carinae | 17 |
– | Median frontal carina not or slightly extending in lower face. Ocellar plate delimited only by a simple lateral carina | 18 |
17 | Ocellar plate with a row of relatively uniform, large foveae along the lateral carinae delimiting the plate |
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– | Ocellar plate delimited only by two simple parallel lateral carinae ( |
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18 | Lateral surface of pronotum longitudinally costate in lower part. Lateral propodeal carinae medially strongly curved. Nucha dorsally longitudinally carinate |
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– | Lateral surface of pronotum without longitudinal costa in lower part. Lateral propodeal carinae nearly parallel, if strongly curved then nucha dorsally glabrate | 19 |
19 | Lateral propodeal carinae medially strongly curved ( |
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– | Lateral propodeal carinae nearly parallel |
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20 | Median frontal carina not distinctly extending in lower face. Head and mesosoma black. Rs and Rs+M veins of forewing distinct and brown in color |
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– | Median frontal carina distinctly extending to middle of lower face. Head and mesosoma brown to dark brown. Rs and Rs+M veins of forewing reduced and pale in color | 21 |
21 | Antennal scrobes not distinctly depressed. Mesoscutum foveate-reticulate. Head, pronotum, mesoscutum, and mesoscutellum dark brown. Antenna yellow. Lateral occipital carinae well developed and crestlike. Median propodeal area glabrate |
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– | Antennal scrobes distinctly depressed. Mesoscutum transversely costate with foveae set in between. Head, pronotum, mesonotum, and mesoscutellum reddish brown. Antenna black, except basal two segments yellow. Lateral occipital carinae not crestlike. Median propodeal area areolate-reticulate |
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22 | Wings slightly and evenly tinted |
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– | Wings with large brown to dark brown macula | 23 |
23 | Upper mesopleuron foveate-rugose |
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– | Upper mesopleuron glabrous | 24 |
24 | Vertex diagonally carinate entirely or heavily punctate medially and diagonally carinate laterally | 25 |
– | Vertex generally glabrate to glabrous with sparse punctures, and sometimes slightly rugose laterally | 26 |
25 | Vertex entirely diagonally carinate. Forewing clear basally and distally with a broad medial smoky band across the marginal and submarginal cells ( |
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– | Vertex heavily punctate medially and diagonally carinate laterally. Forewing clear in distal two-fifths and ferruginous basally. Metasomal T7 of female distinctly exposed |
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26 | Lateral propodeal carinae straight and percurrent; medial propodeal area with one straight and percurrent longitudinal carina |
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– | Lateral propodeal carinae discontinuous, medially interrupted by a large fovea; medial propodeal area with more than one non percurrent longitudinal carinae | 27 |
27 | Forewing ferruginous only medially, and clear both proximately and distally. Metasomal T7 of female only slightly exposed, only 1:15 as long as T6 as measured medially on lateral sides |
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– | Forewing clear only distally, and ferruginous in basal two-thirds. Metasomal T7 distinctly visible, about 1:2.5as long as T6 as measured along middle of lateral sides |
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South Africa.
Named after the Alexandria Forest, which now forms part of the Greater Addo Elephant National Park.
Belongs to the
FEMALE. Length 2–2.5 mm. Head, mesosoma and metasoma dark brown; antennae and legs light yellow; terminal segment of antennae dark brown (
Metanotal-propodeal complex strongly excavated, excavations bordered by strong carinae. Metepisternum dorsally excavated with pubescence, medially polished with indications of minor rugose remnants, ventrally pubescent. Dorsellum with two strong medial foveae; laterally strongly excavated with fine pubescence in lateral depressions. Lateral propodeal carina present; median longitudinal propodeal carina well-defined, crossed by wrinkled transverse and longitudinal carinae extending onto nucha. Rs+M of forewing defined, but nebulous where it arises from basal vein at posterior third (
MALE. Unknown.
Angola.
Democratic Republic of Congo.
Central African Republic.
Named after the village Bayanga.
Belongs to the monophyletic
FEMALE (
MALE (
Cameroon.
Democratic Republic of Congo.
Uganda.
South Africa.
Zimbabwe.
Central African Republic.
Named after the Dzanga-Sangha special forest reserve, which forms part of the Dzanga-Ndoki National Park. Noun in apposition.
Belongs to
FEMALE (
MALE (
South Africa.
South Africa, Zimbabwe.
Namibia
Cameroon
Democratic Republic of Congo
Democratic Republic of Congo, Kenya.
Central African Republic, Democratic Republic of Congo.
“Matele” is BaAka for tattoo. The BaAka pygmies, who live in the forests of Cameroon, Central African Republic, Congo, Democratic Republic of Congo and Gabon, use sap from the plant
Belongs to the
FEMALE. Length 1.9 mm. Head and dorsal mesosoma blackish-brown; lateral mesosoma, metasoma, and coxae dark brown; femora lighter brown, tibiae and tarsi yellowish-brown (
South Africa.
Democratic Republic of Congo.
Uganda, Kenya.
The rainforest at Kanyawara, the area around MUBFS (Makerere University Biological Field Station) where the holotype was collected, is classified by foresters as Parinari forest, distinguished on photo aspect maps by the large spreading crowns of
Belongs to the
FEMALE. Length 2–2.5 mm. Head, mesosoma black; metasoma, coxae and femora (in part) dark brown; antennae and rest of legs light yellow; terminal segment of antennae dark brown (
Metanotal-propodeal complex strongly excavated, excavations bordered by strong carinae. Metepisternum dorsally excavated with pubescence, medially polished, ventrally pubescent. Dorsellum with two strong medial foveae; laterally strongly excavated with fine pubescence in lateral depressions. Lateral propodeal carina present, curved medially; median longitudinal propodeal carina well-defined, crossed by wrinkled transverse and longitudinal carinae extending onto nucha. Rs+M of forewing defined, but nebulous where it arises from basal vein at posterior third (
MALE. Unknown.
South Africa.
Burundi.
Named after the type locality along the Ruvubu River in the Ruvubu National Park.
Belongs to the
FEMALE. Length 3 mm. Head and mesosoma black, metasoma, coxae, femora dark brown; tibiae and tarsi light brown. Wings clear; weakly infuscate in marginal and submarginal cells extending towards apex slightly past vein RS (
There is a damaged specimen in the Royal Museum for Central Africa (RMCA) collection (Congo Belge: P.N.A., Escarpem. De Kabasha, 1500m, 14.xii.1934, G.F. de Witt: 919;
Rwanda.
South Africa.
South Africa.
Democratic Republic of Congo.
South Africa.
1F:
Previously only known from the holotype female (
MALE. Length 1.5mm. Very similar to female, except for longer abdominal petiole, 1.54× as wide as long in dorsal view, as long as high in lateral view. Tergite 5 laterally expanded and by far the largest tergite, latero-medially 1.5× longer than all other tergites combined. First flagellomere 1.5× length of second, slightly wider distally, excavated laterally.
The Holotype was collected while feeding at flowers of
Kenya, South Africa, Tanzania.
Democratic Republic of Congo.
The generic richness within the
Besides inferred association with
The
Largely as a result of recent comprehensive targeted inventory surveys (but also as a result of unearthing previously sampled specimens in historical collections) the species richness of the Afrotropical
SVN was funded by South African National Research Foundation grants: GUN 2068865; GUN 61497; GUN 79004; GUN 79211; GUN 81139. MLB was funded by the Systematic Entomology Lab, USDA/ARS. The expedition to the Central African Republic was supported by funds from the World Wildlife Fund (US). The Ugandan Wildlife Authority and UNCST provided permits to conduct research in Uganda. We thank Dr. Richard Bagine, Director of Research, Kenya Wildlife Service for granting R. S. Copeland permission to sample in Kenyan National Parks and Reserves. Dr. Helida Oyieke, Head of Collections and Research, National Museums of Kenya, was instrumental in granting permission to loan insects for study. We thank Mattias Forshage (Swedish Museum of Natural History) for his careful review of the manuscript including numerous helpful suggestions and István Miko (Pennsylvania State University) for formulating the URI table. We also acknowledge MorphBank (
List of morphological terms. File format: Adobe PDF file (pdf). doi: