Research Article |
Corresponding author: Fernando Farache ( fhafarache@gmail.com ) Academic editor: Michael S. Engel
© 2015 Fernando Farache, Jean-Yves Rasplus.
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:
Farache FHA, Rasplus J-Y (2015) Conidarnes, a new oriental genus of Sycophaginae (Hymenoptera, Agaonidae) associated with Ficus section Conosycea (Moraceae). ZooKeys 539: 119-145. https://doi.org/10.3897/zookeys.539.6529
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The sycophagines are strictly associated with two subgenera of Ficus L. (Moraceae), namely Sycomorus and Urostigma. They mostly oviposit through the fig wall and lay their eggs within the fig flowers, being either gall-makers or parasitoids of other fig wasps. In this contribution, a new genus of Sycophaginae, Conidarnes Farache & Rasplus, gen. n., is described with seven new species: Conidarnes achterbergi Farache & Rasplus, sp. n.; Conidarnes bergi Farache & Rasplus, sp. n.; Conidarnes laevis Farache & Rasplus, sp. n.; Conidarnes santineloi Farache & Rasplus, sp. n.; Conidarnes subtectae Farache & Rasplus, sp. n.; Conidarnes sulcata Farache & Rasplus, sp. n.; and Conidarnes sumatranae Farache & Rasplus, sp. n. Illustrations, morphological diagnoses, dichotomous keys and multi-entry online keys to species are provided. Conidarnes species strictly occur in the oriental region, and their distribution does not overlap with the distribution of the two other genera belonging to the same clade. Due to their relative rarity, we encourage extensive sampling of Conosycea figs to improve our knowledge of the genus.
Chalcidoidea , taxonomy, fig, mutualism, non-pollinating fig wasp, gall maker
Recent phylogenetic analyses of the Chalcidoidea have retrieved Sycophaginae as sister to the pollinating Agaoninae (
There are about 60 described species of Sycophaginae that occur in all tropical and subtropical regions. The subfamily Sycophaginae was retrieved as a monophyletic assemblage and divided into three main clades each of which may warrant tribal status (
A first clade—sister to the remaining Sycophaginae—that only includes Eukoebelea Ashmead species associated with Ficus subsection Malvanthera in Australasia.
A clade that includes species of large and early gallmakers, belonging to three genera: i) the Australasian genus Pseudidarnes Girault associated with Ficus subsection Malvanthera, ii) the Neotropical Anidarnes Bouček associated with Ficus section Americana, and iii) a few species associated with Ficus section Conosycea that cannot be placed in any existing genus which requires the establishment of the new genus described here.
Sister to the previous clade, a highly diversified clade composed of the New World Idarnes Walker associated with Ficus section Americana, and the Old World Sycophaga mostly associated with Ficus subgenus Sycomorus, but also including two species associated with Ficus subgenus Urostigma section Urostigma.
We have recently reviewed the genera Anidarnes and Pseudidarnes with the description of nine new Anidarnes and six new Pseudidarnes species (
Specimen handling and imaging follow
Images were produced with a Leica M16 lens and a JVC KY-75U 3CCD digital camera. Cartograph v5.6.0 (Microvision, Evry, France) software was used for focus stacking.
Type and specimen depositories, and their respective curators are:
CBGP France, Montpellier. Centre de Biologie pour la Gestion des Populations (Emmanuelle Artige).
Conidarnes santineloi Farache & Rasplus, sp. n.
Antennae with 13–14 antennomeres (one or two anelli), including a stub or nipple-like terminal flagellomere. Funicular segments slightly longer than wide to transverse. Antennae inserted at the middle line of compound eyes or below. Toruli contiguous. Clypeal margin bilobed. Malar sulcus absent. Petiole very short, transverse. Ovipositor sheaths without a median constriction and depigmentation.
Females. Size and colour. Body length 1.5–4.0 mm. Length of the ovipositor sheaths 0.4–6.4 mm. Body colour variable. Antennae mostly yellow, sometimes with orange or brown tinges. Head and mesosoma brown to black, usually with green, blue and orange metallic lustre. Legs yellow to brown. Coxae sometimes concolorous with mesosoma. Wings hyaline, sometimes medially infuscate in males. Metasoma usually brown black, sometimes yellow.
Head. Antenna with 13 or 14 antennomeres (including a stub or nipple-like terminal antennomere), usually with two anelli but sometimes with a single anellus (antennal formula 11263 or 11163). Terminal antennomere (i.e. a nipple-like thirteenth or fourteenth antennomere) sometimes conspicuous. Funicular segments slightly longer than wide to transverse. Face sculpture usually reticulate, sometimes slightly engraved. Upper face sometimes smooth. Antennae inserted at the middle line of compound eyes or below. Toruli contiguous, distance between toruli always smaller than one torulus diameter. Clypeal margin bilobed. Malar sulcus absent.
Mesosoma. Pronotum and mesonotum sculpture variable. Pronotum longer than high in lateral view. Notauli usually complete, but incomplete in Conidarnes laevis sp. n. Mesoscutellar-axillar complex with straight or incurved axillular grooves and transverse frenal sulcus, forming a square mesoscutellum (an apomorphy of Sycophaginae). Mesoscutellum trapezoidal, wider near frenal sulcus and narrower near transscutal articulation. Propodeum transverse. Wings with short and sparse pilosity. Postmarginal vein inconspicuous, stub-like. Marginal vein at least as long as stigmal vein.
Metasoma. Petiolate, petiole very short, transverse. Margin of eighth gastral tergite deeply sinuate, A-like, with thumbnail-like medial flap (epipygium) and with a peg-like cercus arising from the membrane on either side of the epipygium (apomorphy of Sycophaginae). Length of the ovipositor sheaths varying from 0.3× (about as long as the hind tibia) to nearly twice as long as body. Ovipositor sheaths without a median constriction and depigmentation.
Males. Similar to females but usually slender and shorter. Exhibiting different coloration, the mesosoma sometimes mostly yellow. Wings sometimes medially infuscate.
The generic name is masculine and derived from Idarnes Walker, 1843, in the same manner than other sycophagine genera (Pseudidarnes Girault, 1927 and Anidarnes Bouček, 1993) and is associated to the prefix Con used for Conosycea, the host plant section of the included species. The origins of the name Idarnes were discussed in
1 | Notauli incomplete (Fig. |
laevis sp. n. |
– | Notauli complete. Face entirely reticulate | 2 |
2 | Sculpture of mesoscutum and mesoscutellum mostly smooth, lateral lobes of mesoscutum engraved reticulate (Fig. |
sulcata sp. n |
– | Sculpture of mesoscutum and mesoscutellum reticulate, lateral lobes of mesoscutum reticulate. Scrobe without a median longitudinal sulcus | 3 |
3 | Antenna with one anellus (Fig. |
santineloi sp. n. |
– | Antenna with two anelli. Funicular segments mostly as long as wide or slightly longer than wide. | 4 |
4 | Ovipositor approximately 1.5× as long as body (Fig. |
achterbergi sp. n. |
– | Ovipositor as long as body or shorter. Coloration of metasoma homogeneously brown | 5 |
5 | Ovipositor nearly as long as body (Fig. |
subtectae sp. n. |
– | Ovipositor nearly 0.5× as long as body (Figs |
6 |
6 | Antennae inserted near the middle line of compound eyes (Fig. |
sumatranae sp. n. |
– | Antennae inserted at the lower line of compound eyes (Fig. |
bergi sp. n. |
♀: MALAYSIA: SE. Sabah: nr. Danum Valley Field C. El. C., 4.96° 117.69°, 400m, 21–25.III.1978, v. Achterberg C, Malaise trap 7 (CBGP).
Metasoma ventrally yellow, dorsally dark brown. Antenna with two anelli. Antennae inserted at the lower line of compound eyes. Mesoscutum and mesoscutellum sculpture reticulate. Length of the ovipositor sheaths 1.7× body length.
Female. Size and colour. Body length 3.8 mm. Length of the ovipositor sheaths 6.4 mm. Antennae yellow orange. Head and mesosoma with metallic lustre, mostly green and blue. Head dorsally more orange. Legs yellow. Metasoma ventrally yellow, dorsally dark brown.
Head. Scape 4.8× as long as wide. Antenna with two anelli. Proximal anellus longer than distal anellus. Funicular segments mostly as long as wide or slightly longer than wide. Terminal antennomere conspicuous. Antennae inserted at the lower line of compound eyes. Supraclypeal area shorter than clypeus and narrow. Face sculpture reticulate. Scrobe without a median longitudinal sulcus.
Mesosoma. Pronotum sculpture reticulate. Pronotum elongated, nearly twice as long as high in lateral view. Mesoscutum and mesoscutellum sculpture reticulate. Notauli complete. Frenal sulcus crenulated. Metascutellum long, rectangular to trapezoidal. Anterior margin of propodeum slightly crenulated. Propodeum sculpture reticulate. Propodeum without a median line.
Metasoma. Length of the ovipositor sheaths 1.7× body length.
Male. Unknown.
The species is dedicated to our colleague and renowned specialist of Hymenoptera, Kees van Achterberg who collected the holotype.
Unknown
♀: INDONESIA: Java: Gunung Tjibodas, –6.88° 106.65°, 530m, 19.XI.1954, van der Vecht J., ex F. involucrata, Wiebes Coll. n° 114 (
Paratypes. 8♀, 8♂: INDONESIA: Java: Gunung Tjibodas, –6.88° 106.65°, 530m, 19.XI.1954, van der Vecht J., ex F. involucrata, Wiebes Coll. n° 114, 19.XI.1954, de Gunst JH, ex F. involucrata, Wiebes Coll. n° 116 & 5103 (5♀, 5♂
Antennae inserted at the lower line of compound eyes. Supraclypeal area shorter than clypeus. Supraclypeal area narrow. Mesoscutum and mesoscutellum sculpture reticulate. Prosternal posterior margin medially acute. Propodeum without a median line. Length of the ovipositor sheaths 0.46× body length.
Female. Size and colour Body length 2.8 mm. Length of the ovipositor sheaths 1.3 mm. Head and mesosoma black, slightly green. Metallic lustre faint. Antennae and legs yellow, coxae concolorous with mesosoma. Metasoma brown.
Head. Scape 5× as long as wide. Antenna with two anelli. Proximal anellus longer than distal anellus. Funicular segments mostly as long as wide or slightly longer than wide. Terminal antennomere inconspicuous. Antennae inserted at the lower line of compound eyes. Supraclypeal area shorter than clypeus, narrow. Face sculpture reticulate. Scrobe without a median longitudinal sulcus.
Mesosoma. Pronotum sculpture reticulate. Pronotum elongated, nearly twice as long as high in lateral view. Prosternal posterior margin medially acute. Mesoscutum and mesoscutellum sculpture reticulate. Notauli complete. Frenal sulcus crenulated. Metascutellum short, inconspicuous. Anterior margin of propodeum crenulated. Propodeum sculpture reticulate. Propodeum without a median line.
Metasoma. Length of the ovipositor sheaths 0.46× body length.
Male. Similar to female, except the following characters: Head and mesosoma darker. Legs browner. Pedicel and funicular segments more elongated. Antenna more setose. Pronotum more elongated.
The specific name is a tribute to our colleague and friend Kees Berg (2 July 1934–31 August 2012), for his excellent and unparalleled work on the taxonomy of fig trees.
Reared from syconia of Ficus involucrata Blume.
♀: INDONESIA: E. Kalimantan: Kutai Nature Reserve, 0.37° 117.27°, 5m, 1978, Leighton, ex F. kerkhoveni, Wiebes Coll. n° 3950 (
Head, mesosoma, and metasoma mostly brown. Upper face smooth; lower face reticulate. Mesoscutum and mesoscutellum sculpture mostly smooth. Notauli incomplete. Frenal sulcus smooth. Length of the ovipositor sheaths 1× body length.
Female. Size and colour. Body length 1.7 mm. Length of the ovipositor sheaths 1.7 mm. Antennae yellow. Head dark brown, with metallic green lustre, mostly at the lower face. Mesosoma and metasoma brown. Legs proximally brown, tibiae and tarsi yellow.
Head. Scape 3.5× as long as wide. Antenna with two anelli. Proximal anellus shorter than distal anellus. Funicular segments mostly transverse. Terminal antennomere inconspicuous. Antennae inserted at the lower line of compound eyes. Supraclypeal area shorter than clypeus, or inconspicuous, narrow. Upper face smooth; lower face reticulate. Scrobe without a median longitudinal sulcus.
Mesosoma. Pronotum sculpture mostly smooth, slightly engraved. Pronotum elongated, nearly twice as long as high in lateral view. Mesoscutum and mesoscutellum sculpture mostly smooth. Notauli incomplete. Frenal sulcus smooth. Metascutellum short, inconspicuous. Anterior margin of propodeum smooth. Propodeum sculpture smooth. Propodeum without a median line.
Metasoma. Length of the ovipositor sheaths 1× body length.
Male. Unknown.
The specific name refers to the smooth body sculpture observed in this species.
Reared from syconia of Ficus kerkhovenii Koord. & Valeton.
This species presents unique characters, such as a smooth body with no sculpture and an elongated mesosoma. These characters are mostly associated to galler fig wasps that enters the syconium through the ostiole (
♀: BRUNEI: Temburong National Park, 4.554° 115.157°, 100m, 25.XI.1996, Rasplus J.Y., ex F. pallescens, n° JRAS00114 (CBGP).
Paratypes. 9♀, 5♂: same locality and information as holotype (7♀, 3♂ CBGP; 1♀, 1♂
Antenna with one anellus. Funicular segments mostly transverse. Mesoscutum and mesoscutellum sculpture reticulate. Prosternal posterior margin not medially acute. Propodeum with a depressed median line. Length of the ovipositor sheaths 0.3× body length.
Female. Size and colour. Body length 1.6 mm. Length of the ovipositor sheaths 0.45 mm. Head, mesosoma, and metasoma black, slightly green. Metallic lustre faint. Antennae and legs yellow, forecoxae brown.
Head. Scape 3.5× as long as wide. Antenna with one anellus. Funicular segments mostly transverse. Terminal antennomere inconspicuous. Antennae inserted just below the middle line of compound eyes. Supraclypeal area higher than clypeus, wide. Face sculpture reticulate. Scrobe without a median longitudinal sulcus.
Mesosoma. Pronotum sculpture reticulate. Pronotum not elongated, 1.5× as long as wide in lateral view or less. Prosternal posterior margin not medially acute. Mesoscutum and mesoscutellum sculpture reticulate. Notauli complete. Frenal sulcus crenulated. Metascutellum long, rectangular to trapezoidal. Anterior margin of propodeum smooth. Propodeum sculpture slightly reticulate to smooth. Propodeum with a depressed median line.
Metasoma. Length of the ovipositor sheaths 0.3× body length.
Male. Similar to female except the following characters: Mesosoma and metasoma yellow. Mesoscutum, axillae, axillulae, and dorsal metasoma partially brown black, with faint metallic lustre. Legs completely yellow. Pedicel and funicular segments slender. Antenna more setose. Pronotum slender.
The specific name is dedicated to our friend and colleague Rodrigo Augusto Santinelo Pereira due to his excellent and pioneering work on fig wasps and Ficus in Brazil.
Collected from syconia of Ficus pallescens L., the form with small leaves (see
♀: BRUNEI: Temburong, Kuala Belalong, 4.538° 115.159°, 105m, 23.IV.1997, Hossaert-Mckey M., ex F. subtecta, n° JRAS00117 (CBGP).
Antenna with two anelli. Funicular segments mostly as long as wide or slightly longer than wide. Antennae inserted just below the middle line of compound eyes. Supraclypeal area wide. Mesoscutum and mesoscutellum reticulate. Length of the ovipositor sheaths 0.9× body length.
Female. Size and colour. Body length 1.8 mm. Length of the ovipositor sheaths 1.6 mm. Antennae yellow. Head and mesosoma black, with faint blue, green, and orange metallic lustre. Legs mostly yellow distally. Coxae almost concolorous with body. Femora yellow brown. Metasoma dark brown.
Head. Scape 4.8× as long as wide. Antenna with two anelli. Proximal anellus longer than distal anellus. Funicular segments mostly as long as wide or slightly longer than wide. Terminal antennomere conspicuous. Antennae inserted just below the middle line of compound eyes. Supraclypeal area higher than clypeus, and wide. Face sculpture reticulate. Scrobe without a median longitudinal sulcus.
Mesosoma. Pronotum sculpture reticulate. Pronotum elongated, nearly twice as long as high in lateral view. Mesoscutum and mesoscutellum sculpture reticulate. Notauli complete. Frenal sulcus crenulated. Metascutellum long, rectangular to trapezoidal. Anterior margin of propodeum crenulated. Propodeum sculpture reticulate. Propodeum with a depressed median line.
Metasoma. Length of the ovipositor sheaths 0.9× body length.
Male. Unknown.
The specific name refers to the host fig species.
Reared from syconia of Ficus subtecta Corner
♀: CHINA: Yunnan: Luxi county, Mang Hai town, 24.53° 102.54°, 1750m, 28.IV.2006, Rasplus J.Y.; Peng Y.Q. & Yang D.R., ex F. altissima, n° JRAS01616_04 (CBGP).
Paratypes. 7♂: CHINA: Yunnan: Cheng Zhi village, 21.92° 101.24°, 540m, 6♂, 9.IV.2002, Gu H.Y. & Rasplus J.Y., ex F. altissima, n° JRAS0875 (5 ♂ CBGP; 1♂
Scrobe with a median longitudinal sulcus, extending from median ocellus to interantennal area. Mesoscutum and mesoscutellum sculpture mostly smooth. Lateral area of the mesoscutum mostly engraved reticulate. Propodeum sculpture smooth, slightly engraved alutaceous near spiracles. Propodeum without a median line. Length of the ovipositor sheaths 1× body length.
Female. Size and colour. Body length 2.7 mm. Length of the ovipositor sheaths 2.6 mm. Antennae yellow. Head with metallic lustre, mostly green, slightly orange and blue. Mesosoma mostly brown, with faint metallic lustre, green and blue. Legs mostly brown, tarsal segments and foretibia yellow. Metasoma dark brown.
Head. Scape 4.6× as long as wide. Antenna with two anelli. Proximal anellus nearly as long as distal anellus. Funicular segments mostly transverse. Terminal antennomere conspicuous. Antennae inserted at the lower line of compound eyes. Supraclypeal area inconspicuous. Face sculpture mostly reticulate, smooth near scrobe. Scrobe with a median longitudinal sulcus, extending from median ocellus to interantennal area.
Mesosoma. Pronotum sculpture alutaceous, engraved. Pronotum elongated, nearly twice as long as high in lateral view. Prosternal posterior margin medially acute. Mesoscutum and mesoscutellum sculpture mostly smooth. Lateral area of the mesoscutum mostly engraved reticulate. Notauli complete. Frenal sulcus smooth. Metascutellum as long as frenum, smooth, and not well delimited laterally. Anterior margin of propodeum smooth. Propodeum sculpture smooth, slightly engraved alutaceous near spiracles. Propodeum without a median line.
Metasoma. Length of the ovipositor sheaths 1× body length.
Male. Similar to female except the following characters: Mesosoma and metasoma mostly yellow. Axillulae, metanotum, and propodeum mostly brown. Some metasomal segments slightly brown dorsally. Scape and pedicel shorter, funicular segments more transverse. Antenna more setose.
The specific name refers to the longitudinal sulcus separating the scrobal cavity in this species.
Reared from syconia of Ficus altissima Blume.
Conidarnes sulcata was included in the phylogenetic analyses by
♀: INDONESIA: Sulawesi: Pattunuang, –5.059° 119.718°, 180m, 27.VIII.2007, Rasplus J.Y., ex F. sumatrana, n° JRAS02085_0202 (CBGP).
Paratypes. 1♂: same locality and information as holotype, n°JRAS02085_0201 (CBGP).
Antennae inserted near the middle line of compound eyes. Funicular segments mostly as long as wide or slightly longer than wide. Mesoscutum and mesoscutellum sculpture reticulate. Propodeum with a reticulate median line, slightly striate, and thicker near anterior margin. Length of the ovipositor sheaths 0.4× body length.
Female. Size and colour. Body length 1.9 mm. Length of the ovipositor sheaths 0.8 mm. Antennae yellow. Head and mesosoma black, with green and blue metallic lustre. Legs mostly yellow, forecoxae concolorous with body. Hindcoxae proximally concolorous with body. Metasoma dark brown.
Head. Scape 3.5× as long as wide. Antenna with two anelli. Proximal anellus longer than distal anellus. Funicular segments mostly as long as wide or slightly longer than wide. Terminal antennomere inconspicuous. Antennae inserted near the middle line of compound eyes. Supraclypeal area higher than clypeus and narrow. Face sculpture reticulate. Scrobe without a median longitudinal sulcus.
Mesosoma. Pronotum sculpture reticulate. Pronotum elongated, nearly twice as long as high in lateral view. Mesoscutum and mesoscutellum sculpture reticulate. Notauli complete. Frenal sulcus crenulated. Metascutellum long, rectangular to trapezoidal. Anterior margin of propodeum crenulated. Propodeum sculpture reticulate. Propodeum with a reticulate median line, slightly striate, and thicker near anterior margin.
Metasoma. Length of the ovipositor sheaths 0.4× body length.
Male. Similar to female, but slightly smaller.
The specific name refers to the host species.
Reared from syconia of Ficus sumatrana Miq.
Conidarnes sumatranae was included in several phylogenetic analyses (
5♂: INDONESIA: E. Kalimantan: Kutai Nature Reserve, 0.37° 117.27°, 1978, Bingham M., ex F. sundaica Bl. v. beccariana (King) det. Corner, Wiebes Coll. N°3543 (
Female. Unknown.
Male. Size and colour. Body length 3.1 mm. Antennae yellow orange. Head and mesosoma mostly black, with metallic blue lustre. Legs mostly dark brown, proximally darker. Metasoma brown.
Head. Scape 5.3× as long as wide. Antennae inserted just below the middle line of compound eyes. Supraclypeal area shorter than clypeus and narrow. Face sculpture reticulate. Scrobe with a median longitudinal sulcus, extending from median ocellus to interantennal area.
Mesosoma. Pronotum sculpture alutaceous, engraved. Pronotum elongated, nearly twice as long as high in lateral view. Mesoscutum and mesoscutellum sculpture reticulate. Notauli complete. Frenal sulcus crenulated. Metascutellum long, rectangular to trapezoidal. Anterior margin of propodeum crenulated. Propodeum sculpture slightly reticulate to smooth. Propodeum without a median line.
This species was reared from Ficus sundaica Blume v. beccariana (King).
We have examined only males, but they clearly belong to an undescribed species. Since we described Conidarnes species mostly based on females, we prefer not to describe this species until more specimens are found.
In this study, we describe a new oriental genus of Sycophaginae that includes seven new species. Conidarnes can easily be assigned to Sycophaginae due to the presence of a square mesoscutellum and the morphology of the terminal gastral tergites/epipygium, which are synapomorphies of the subfamily (
Phylogenetically, Conidarnes is nested within a clade including Pseudidarnes and Anidarnes. Species belonging to this clade are large gall inducers (
Conidarnes species are restricted to the Oriental region. Only one species was sampled in continental Asia (C. sulcata, from Xishuangbanna in southwest China), whereas all other species were sampled in the insular region of Southeast Asia: five species in Borneo (C. achterbergi, C. laevis, C. santineloi, C. subtectae, and an undescribed species ex Ficus sundaica), one in Java (C. bergi) and one in Sulawesi (C. sumatranae).
The distribution of Conidarnes does not overlap with distribution of the two other genera belonging to the same clade. Indeed, Anidarnes is restricted to America (
Another characteristic of Sycophaginae species belonging to the clade of large gall inducers is that they are rare and globally difficult to sample (
We are indebted to Simon van Noort for the assistance in the development of online keys. For assistance in imaging fig wasps we thank Gunther Fleck. We are also grateful to Frédérique Bakker for providing specimens from
Description characters and HAO codes for Conidarnes
Data type: List of morphological characters
Explanation note: List of characters and states, with HAO portal codes. http://portal.hymao.org/projects/32/public/ontology
LUCID Key for Conidarnes
Data type: Key to species
Explanation note: Digital multi-entry key to species of Conidarnes in LUCID format: http://www.lucidcentral.com