﻿Two new genera and one new species of the tribe Adeshini (Hymenoptera, Braconidae, Braconinae) from India and South Africa

﻿Abstract Two new genera and one new species of the Braconinae tribe Adeshini are described and illustrated: Crenuladesha Ranjith & Quicke, gen. nov., type species Adeshanarendrani Ranjith, 2017, comb. nov. from India, and Protadesha Quicke & Butcher, gen. nov., type species Protadeshaintermedia Quicke & Butcher, sp. nov. from South Africa. The former lacks the mid-longitudinal propodeal carina characteristic of the tribe, and the latter displays less derived fore wing venation with two distinct abscissae of vein 2CU. A molecular phylogenetic analysis is included to confirm their correct placement. Since neither of the two new genera displays all of the characters given in the original diagnosis of the Adeshini a revised diagnosis is provided, as well as an illustrated key to the genera.

Two new genera and one new species of the tribe Adeshini (Hymenoptera, Braconidae, Braconinae) from India and South Africa

Introduction
The subfamily Braconinae (Hymenoptera: Braconidae) is a diverse, species rich and morphologically distinct group of wasps with a cosmopolitan distribution (Shaw and Huddleston 1991). Until now, the subfamily included more than 2500 species belonging to 188 genera (Ranjith et al. 2016a, b). Most of the generic diversity is restricted to tropical regions (Quicke 2015) especially those of the Old World. The biology of braconines ranges from being strictly parasitoid, by far the commonest biology, though a few cases of entomophytophagy (Ranjith et al. 2016b), obligate phytophagy (Flores et al. 2005;Perioto et al. 2011) and obligate predation within plant galls are also known (Ranjith et al. 2022). Most braconines are idiobiont ectoparasitoids of immature hosts which principally are moderately concealed, such as gall inducers and leaf rollers or miners, or deeply concealed such as wood-borers (Shaw and Huddleston 1991;Quicke 1993;Ranjith et al. 2016b). Exceptionally, members of the Braconini subtribe Aspidobraconina are endoparasitoids within butterfly pupae (van Achterberg 1984;Quicke 1987) and this distinctive way of development was recently also reported in the apparently closely-related genus Crinibracon Quicke (Gupta et al. 2016).
The tribal classification of the Braconinae is confused, and has not been fully investigated. Historically, several tribes were erected based almost entirely on the West Palaearctic fauna (Quicke 1987). van Achterberg (1989) effectively synonymized several previously recognised tribes (see Quicke 1988bQuicke , 1990Quicke et al. 1996), into the Aphrastobraconini (previously restricted to a small group of genera with aberrant wing venation, by defining them based on a ventrally elongate scapus. Formal changes in the systematic status of most of the remaining tribes have been superseded by molecular research (Belshaw et al. 2001) which supports two major clades, the Aphrastobraconini and the Braconini, and with the two genera of Coeloidini nested between them, though not as a monophyletic group. A detailed molecular study of relationships in preparation confirms that the Adeshini also constitute a valid separate tribe.
The tribe Adeshini was erected by van Achterberg (1983) within the Braconinae to accommodate two genera, Adesha Cameron and Adeshoides van Achterberg, which display a unique apomorphic condition of their fore wing venation with respect to the remainder of the subfamily, both having vein 2CU not differentiated into two abscissae and 2cu-a arising interstitially with m-cu (in the terminology of van Achterberg (1983) CUla arises at the same level as 2-CU1 and consequently having vein CUlb, if present, much longer than 3-CU1) (Quicke 1986).
Including those described in this paper, the Adeshini includes 16 described species classified within nine genera (van Achterberg 1983; Quicke 1986Quicke , 1988Quicke and Ingram 1993;Quicke and Polaszek 2000;Wang et al. 2006;Liu et al. 2007;Ranjith et al. 2017;Oanh et al. 2023). The known species are entirely from the Old World, being distributed from Africa, through tropical Asia to Australia (Yu et al. 2016). The majority of the genera are known from only one or just a few specimens, and three previously described genera, Adeshoides, Aneuradesha and Indadesha are still monotypic (Quicke 1986;Quicke and Polaszek 2000). Further, most genera are currently known from more limited geographic regions, but Africadesha Quicke has a disjunct distribution comprising the Afrotropical and Australian regions (Quicke 1986;Quicke and Ingram 1993) and Furcadesha Quicke is recorded from tropical Asia to the Australia (Quicke 1986;Quicke and Ingram 1993;Liu et al. 2007).
Biology is known for four species, but based on these, they appear to be specialized parasitoids of hispine chrysomelid beetles. Adesha albolineata Cameron has been reared from the pest coconut leaf-miner, Promecotheca cumingi Baly (Quicke and Polaszek 2000), Aneuradesha harleyi Quicke from Asmangulia cuspidata Maulik, a minor pest of sugar-cane (Quicke and Polaszek 2000), and one species of Furcadesha has been reared from Dactylispa sp. (Jalala, unpublished), both on monocotyledonous host plants.
Here we describe two new, morphologically aberrant, genera of Adeshini to make their names available. We use DNA sequences from one mitochondrial (cytochrome oxidase I, COI) and one nuclear gene (28S) to confirm the placement of both new genera within the Adeshini. A revised tribe diagnosis and an illustrated key to all genera are presented. Further, we provide photographic portfolios of all of the adeshine genera.

Phylogenetic analysis
A molecular data matrix comprising 37 Braconinae taxa plus one member of the Exothecinae was assembled from published and newly-generated sequences. Our taxon sampling was based on covering a reasonable amount of the taxonomic diversity of the Braconini and Aphrastobraconini. All genera were represented by both 28S D2 or D2+D3 sequences except for one Adesha specimen, and we included data from all available sequenced Adeshini specimens which collectively represented seven of the nine known genera.
The length-variable 28S sequences were aligned according to the secondary structure model of Gillespie et al. (2005) and our interpretation is provided in Suppl. material 1. The confidently alignable 28S bases were treated as either pairing or not-pairing (Butcher et al. 2014;Quicke et al. 2016), and the three codon positions of CO1 were also treated as separate partitions for analysis. Sequences were analyzed on RAxML (Stamatakis, 2014), using a GTR+G rate model with five data partitions (three cytochrome oxidase codons plus pairing and non-pairing 28S bases) and conducting a combined ML search and rapid bootstrap using the '-f a' option and 100 runs.

Phylogenetic results
The Adeshini were recovered monophyletic with 100% bootstrap support as well as in individual analyses of the single gene data (not shown), but nested among the representative Braconini (as was the single included representative of the Coeloidini (Fig. 1). These three groups together formed a clade in the ML tree but with only 15% bootstrap support. The 10 included genera of Aphrastobraconini were recovered as a monophyletic clade with 100% bootstrap support (Fig. 1).
Both of the new genera described below were recovered in the Adeshini. Crenuladesha gen. nov. was recovered as sister group to Spinadesha with 100% bootstrap support. In neither of the single analaysis or combined trees was Protadesha gen. nov. recovered basally within the tribe (Fig. 1).

Taxonomy Adeshini van Achterberg
Adeshini van Achterberg, 1983: 175;Quicke 1987: 99;Quicke 1988a: 265. Diagnosis. Distinguished from other Braconinae by having the first abscissa of fore wing vein 2CU (the normally transverse vein 2CUa) short or completely absent, with 2CUb often arising at the same level of 1CUb OR if below the level of 1CUb, then vein 2cu-a present AND veins 2CU distinct AND the former not longer than vein 2cu-a. Additionally, scapus shorter ventrally than dorsally in lateral view; hind wing vein 1r-m very short, approximately as long as R, and oblique, the basal cell consequently being very narrow; hind wing posteriorly emarginate and setosity of posterior margin of hind wing very long compared with that of the wing membrane; first metasomal tergite movably connected to the second tergite; ovipositor shorter than the metasoma, at least with a distinct, pre-apical, dorsal angulation and ventral serrations.
Diagnosis. Differs from all other genera of Adeshini in having a complete, partially crenulate, mid-longitudinal propodeal groove, other genera having at least a partial, and usually complete, mid-longitudinal carina. In addition, unlike other members of the tribe the basal lobe of the claws is distinctly pectinate, and the dorsal valve of the ovipositor has a distinct pre-apical angulation in lateral profile, rather than being smoothly rounded.
Description. Head. Scapus shorter ventrally than dorsally in lateral aspect. Etymology. From the combinations of Latin "crenula" meaning notched and Adesha type genus of the tribe, in reference to the crenulate propodeal groove.
Remarks. The distinct and complete malar suture and crenulated mid-longitudinal groove on the propodeum are putative autapomorphies of Crenuladesha Ranjith & Quicke, gen. nov. In addition to that the presence of postero-lateral semicircular emarginations of the fifth metasomal tergite displays some affinities with the Adesha, Furcadesha and Indadesha. In common with Adesha, Aneuradesha, Furcadesha and Spinadesha are the presence of a mid-longitudinal groove on the frons, smooth vertex, mesosternum and pleural suture and exhibit a lesser character sharing with Africadesha, Aneuradesha and Protadesha on the basis of a single character; smooth mesosternum.   Diagnosis. Differs from all other genera of Adeshini in having fore wing vein 2CU with a clearly-differentiated and sub-transverse basal abscissa (2CUa).

Etymology. From
Greek proto meaning first and the genus name Adesha in reference to the potentially earlier form of modification of the fore wing venation in relation to that of other Adeshini.

Protadesha intermedia
Ovipositor sheathes approximately as long as hind basitarsus. Coloration. Body and legs uniformly dark honey brown except: three small dark marks, one on each lateral lobe and a smaller less distinct one medio-anteriorly on the medial lobe of the mesoscutum; tarsi of all legs which are darker brown. Wings with hyaline membrane and brown venation; pterostigma yellowish medially darker around its borders. It should be noted that the mounted holotype was chemically extracted for its DNA and is, as a consequence, is now somewhat duller than the fresh specimen would have been. Fig. 4A shows the specimen prior to DNA extraction.

Distribution. South Africa (Gauteng).
Etymology. The species name intermedia is Latin meaning in between, referring to the position of fore wing vein 2CUb relative to 1CU.
Adesha acuta Quicke, 1986 Material examined. Holotype ♀, Malaysia, Pahang, Kuala Lipis, 29.v.1926, coll. H.M. Pendlebury (NHMUK). Distribution. Malaysia (Pahang), Pakistan (Nargis et al. 2020  We have seen numerous other specimens of Spinadesha from Thailand and Borneo all of which are morphologically very similar. Oanh et al. (2023) provided a key to the five species described to date based largely on coloration and slight differences in the profiles of the denticulations on the posterior margin of the 5 th metasomal tergite.

Morphological plasticity
Although with only weak bootstrap support, the combined analysis (Fig. 1) and each individual gene analysis (not shown) recovered both Crenuladesha gen. nov. and Protadesha gen nov. in derived positions within the Adeshini. This suggests that the presence of a sub-transverse fore wing vein 2CU1b in Protadesha is likely a character reversal. The arrangement of veins forming the distal part of the 1 st subdiscal cell with 1CU and 2CU at the same level as in other Adeshini occurs in several other groups of Braconidae, e.g., some Hormiinae, the rogadine tribe Facitorini, a few Doryctinae and Alysiinae. In all these cases it is restricted to small-bodied species/genera. The longitudinal propodeal groove of Crenuladesha seems likely to have resulted from increasing depth of the often crenulate border of the mid-longitudinal carina, finally with the central carina being lost. During such an evolutionary transition, the function of the structure (presumably propodeal strengthening) would be maintained.
Even though many of the genera are represented by one or two species, both Furcadesha and Africadesha have particularly wide distributions. Several species, based on morphology, also appear to have wide distributions, for example, Adesha albolineata (Borneo to Pakistan) and A. acuta (Peninsular Malaysia to Pakistan). Further, two specimens of Adesha sp., one from Thailand the other from the island of Sumatra (Indonesia) for which DNA sequence data are available, had barcodes belonging to the same BIN (AAG7566) and thus are most probably conspecific.