Orientocardiochiles, a new genus of Cardiochilinae (Hymenoptera, Braconidae), with descriptions of two new species from Malaysia and Vietnam

Abstract For the first time in 21 years, a new genus of cardiochiline braconid wasp, Orientocardiochiles Kang & Long, gen. nov. (type species Orientocardiochiles joeburrowi Kang, sp. nov.), is discovered and described. This genus represents the ninth genus in the Oriental region. Two new species (O. joeburrowi Kang, sp. nov. and O. nigrofasciatus Long, sp. nov.) are described and illustrated, and a key to species of the genus, with detailed images, is added. Diagnostic characters of the new genus are analyzed and compared with several other cardiochiline genera to allow the genus to key out properly using an existing generic treatment. The scientific names validated by this paper and morphological data obtained from this project will be utilized and tested in the upcoming genus-level revision of the subfamily based on combined morphological and molecular data.

Orientocardiochiles Kang & Long, gen. nov. represents the 17 th genus in the world, the ninth genus in the Oriental region, and the fifth genus in both Malaysia and Vietnam.
Females of the subfamily are known as solitary koinobiont endoparasitoids of lepidopteran larvae, laying only one egg in each host and allowing the host to continue its development while parasitized. Larvae of Pyralidae and Noctuidae, among other families, are typically hosts and include important crop pests such as the tobacco budworm, Heliothis virescens (Fabricius, 1777), and cotton bollworm, Heliocoverpa armigera (Hübner, 1808) (Huddleston and Walker 1988). Of the 13 cardiochiline species of Vietnam's fauna, one species, Cardiochiles philippensis Ashmead, 1905, is reported as an endoparasitoid reared from larvae of the rice leaf-folder, Cnaphalocrocis medinalis (Guenée, 1854) (Lepidoptera: Pyralidae) (Long and Belokobylskij 2003).

Materials and methods
The type specimens for the present work were provided by the Braconidae Collection of the Institute of Ecology and Biological Resources (IEBR: Ha Noi, Vietnam), the Hymenoptera Institute (HIC: 116 Franklin Ave., Redlands, California, USA), and Museums Victoria (MVMA: Melbourne, Victoria, Australia). Other materials were borrowed from HIC and Illinois Natural History Survey (INHS: Champaign, Illinois, USA). All HIC material will be deposited in the Canadian National Collection of Insects (CNC: Ottawa, Ontario, Canada), including the holotype of Orientocardiochiles joeburrowi Kang sp. nov. and the holotype of Orientocardiochiles nigrofasciatus Long, sp. nov. is housed in IEBR.
The morphological terminology mostly follows Dangerfield et al. (1999) andvan Achterberg (1993). Morphological terminology can also be checked at the Hymenoptera Ontology website (http://portal.hymao.org/projects/32/public/ontology/). Terms for sculpture are based on Harris (1979), and wing vein terminology mostly follows the modified Comstock-Needham system (van Achterberg 1993). Definitions of the morphological measurements used in this study are mostly based on van Achterberg (1988).
For the specimen of O. joeburrowi sp. nov., morphological analysis was conducted using a Leica MZ75 stereomicroscope. Color habitus images were captured using a Visionary Digital BK Plus imaging system (Dun, Inc.), equipped with a Canon EOS 5DS R DSLR camera. Images were stacked in Zerene Stacker v. 1.04 (Zerene Systems LLC.). All images were made by IK and edited in Adobe Photoshop CS 6 (Adobe Systems, Inc). Body parts of the specimen were also measured using Adobe Photoshop CS 6 (Adobe Systems, Inc).
For the specimen of O. nigrofasciatus sp. nov., morphological analysis was conducted using an Olympus SZ61 binocular microscope; measurement were carried out using an Olympus SZ40 binocular microscope; the photographs were produced by KDL with a Sony 5000 digital camera attached to a Nikon SMZ 800N binocular microscope at IEBR and processed with Adobe Photoshop CS5 to adjust the size and background.
The key to species of Orientocardiochiles gen. nov. and descriptions of the two species are based on females. Distribution maps were produced using QGIS 3.10.0 (QGIS Development Team 2019). Google satellite maps were downloaded via the QuickMapServices plugin.
Diagnosis (based on all the members of the genus). Body large and stout, finely sculptured, whitish to yellow pale in color with black spots and stripes. Head in dorsal view transverse. Antenna 41-or 43-segmented. Eyes sparsely pilose. Clypeus with distinct suture and two clypeal tubercles present apically. Malar suture present. Mandible bidentate and angularly bent ventrally. Mouthparts (the length of galea and glossa) short. Maxillary palpus 5-or 6-segmented. Labial palpi 4-segmented. Notauli deep, crenulate, meeting posteriorly in deep smooth area. Scutellar sulcus curved, with 5+ crenulae. Scutellum more or less elevated medially, without carina laterally and apically. Propodeal areola completely developed and kite-shaped or elongated pentagonal. Epicnemial carina absent. Mesopleuron mostly smooth; precoxal sulcus well-defined and crenulate, not reaching posterior margin. Metapleuron rugulose. Mesosternal sulcus finely crenulate. Hind tibia without apical projection; inner tibial spur distinctly longer than outside spur, subequal to half of hind basitarsus. Tarsal claws pectinate. Forewing with elongated pterostigma; vein r reach-ing at apical fourth of pterostigma; SR1 sharply angled at basal fourth; basal fourth of vein SR1 almost perpendicular to apical vein 3-SR. Vein 1a present as a spectral short trace; 1 st discal cell in forewing rather short compared to first submarginal cell. Second submarginal cell elongated. First subdiscal (brachial) cell broad. M+CU in hind wing distinctly shorter than 1-M. Hind wing with 6 hamuli. T1 widened apically, with lateral suture clearly defined throughout. T2 mostly rugose except for plateau-like projection (Figs 2D, 5G); plateau-like projection of T2 present at anteromedial base. T3 entirely smooth. Hypopygium sharply pointed at apex, median longitudinal area evenly sclerotized or largely desclerotized medially throughout; median enfold of hypopygium present or absent. Ovipositor sheath longer than metasoma, pointed at apex, and with short setae throughout.
Distribution. Oriental (Malaysia, Vietnam). Biology. Unknown. Etymology. The name for the genus refers to Cardiochiles from the Oriental region. From "orientum" (Latin for the eastern region) and the generic name "Cardiochiles Nees, 1819." Gender: masculine.
Male. Unknown. Etymology. Named in honor of Joseph Lee Burrow, the world-class college football quarterback for the LSU Tigers and the 2019 Heisman Trophy winner.

Distribution.
Orientocardiochiles joeburrowi sp. nov., is known from only one female specimen collected from Wang Kelian, Malaysia, which is near the Thailand-Malaysia border (Fig. 4A, B).

Character discussion
Members of Orientocardiochiles Kang & Long, gen. nov., Austerocardiochiles Dangerfield, Austin, & Whitfield, 1999, Hansonia Dangerfield, 1996, Heteropteron Brullé, 1846, and Wesmaelella Spinola, 1851 share the presence of lateral sutures on the first metasomal tergum that are clearly defined throughout the length of the tergum (Figs  1D, 2D, 5G). According to the most recent complete genus-level phylogeny of the subfamily based on morphological characters (Dangerfield et al. 1999), three monophyletic clades are separated based on the presence or absence and length of eye setae.
The basal clade of Dangerfield et al. (1999) is composed of members of Heteropteron and Wesmaelella that have glabrous eyes (Fig. 7B). The members of these genera do not possess clypeal tubercles, nor a median areola on propodeum (Fig. 7A, B). Also absent is the lens-shaped area, or plateau-like projection on the second metasomal tergum. Based on the characters mentioned, Orientocardiochiles gen. nov. cannot be placed in the basal clade.
Clade A of Dangerfield et al. (1999) contains Hansonia and other genera that have short and sparse eye setae as in Orientocardiochiles gen. nov. (Figs 2F, 5C, D). Members of Austerocardiochiles and other genera in clade B mostly have long and dense eye setae (Fig. 1F), unlike Orientocardiochiles gen. nov. Based on the eye setae character, Orientocardiochiles gen. nov. can be placed in clade A. However, Orientocardiochiles gen. nov. could be included in the clade B if the short eye setae character is an independently developed character, as has occurred in one of the basal genera of the clade B, Asiacardiochiles Telenga, 1955. Based on the hypopygial median fold of O. nigrofasciatus sp. nov. (Fig. 5N), placing Orientocardiochiles gen. nov. in clade B is more probable than clade A because members of two genera in clade B, Austerocardiochiles and Cardiochiles, possess the hypopygial median fold (Fig. 1C).

Distribution and diversity
As mentioned in the Introduction, members of Cardiochilinae are distributed worldwide. Regarding the genus-level diversity, the Australasian region has the highest diversity. Ten cardiochiline genera have been recorded from the Australasian region (Dangerfield et al. 1999;Yu et al. 2016). As a result of this work, we confirmed that Cardiochilinae exhibits the second highest genus-level diversity in the Oriental region. Nine genera, including the introduced genus Toxoneuron (Say, 1836), are recorded from the region. Five genera have been recorded from Malaysia and Vietnam in each country. More genera will be likely to be found in Malaysia and Vietnam after further collecting.

Future directions
In the past two decades, molecular data combined with morphological data has been widely utilized to improve resolution of the species-, genus-, tribe-, and subfamily-level relationships of Braconidae. However, a genus-level phylogeny of Cardiochilinae based on molecular data is lacking. This shows the necessity of a novel phylogeny. Therefore, IK is in process of conducting a new genus-level revision of Cardiochilinae to elucidate the genus-level relationships of the subfamily using a novel phylogeny based on combined molecular and morphological data.