Review of the world species of Exoryza ( Hymenoptera , Braconidae , Microgastrinae ) , with description of five new species

The world species of the genus Exoryza (Hymenoptera, Braconidae, Microgastrinae) are revised. Ten species are recognized, including five new, authored by Fernandez-Triana: mariabustosae, richardashleyi, ritaashleyae, rosamatarritae and yeimycedenoae. The species Dolichogenidea oryzae Walker, 1994 is considered as related to Exoryza – although is not formally transferred to that genus. A dichotomous key to all species, descriptions and illustrations are provided. The only region where the genus is not yet recorded is Australasia, but this may be a collecting artifact. Host caterpillars (Lepidoptera) include species within Choreutidae, Depressariidae, Gelechiidae, and Pyraloidea – all but Pyraloidea are new host records. The status of Exoryza is questioned (it may only represent a species-group within the genus Dolichogenidea) but it is retained as a valid genus until a comprehensive phylogenetic study of Microgastrinae is available.


Introduction
The braconid subfamily Microgastrinae contains more than 60 genera (Yu et al. 2012), many of them lacking comprehensive revisions and keys to their species.Exoryza is a rarely collected genus (Mason 1981, Valerio et al. 2004), and it is poorly represented in collections.It was described by Mason (1981) to include two species of Apantelini (sensu Mason 1981) with broad and heavily sculptured metasomal tergites 1 -2, and a coarsely rugose and areolate propodeum.Three other species have since been described (Song and Chen 2003, Valerio et al. 2004, Rousse and Gupta 2013), expanding the known distribution of the genus to all continents but Australia.
Here we review Exoryza, as it occurs in Area de Conservación Guanacaste (ACG), northwestern Costa Rica (Janzen et al. 2009, Janzen andHallwachs 2011), as part based on Kimura 2-parameter to visually demonstrate the variation present within and between each species in the DNA barcode locus is presented in Figure 40.All information for the sequences (including GenBank accessions) associated with each individual specimen can be retrieved from the Barcode of Life Data System (BOLD) (Ratnasingham and Hebert 2007) using the persistent DOI dx.doi.org/10.5883/DS-ASEXORYZ[requested April 15, 2016 but not yet active].
The new ACG species described below received patronyms honoring the teachers and supporters of a biodiversity conservation event and school child nature awareness competition, conducted by the Programa de Educación Biológica (PEB) of ACG in the last half of 2015 (Kazmier 2015).
The status of Exoryza as a valid genus has been questioned by many authors (e.g., Valerio et al. 2004, Rousse and Gupta 2013, Fernandez-Triana et al. 2014).Mason (1981) characterized Exoryza as having T1 and T2 heavily sculptured, and propodeum coarsely rugose (with areola present but obscured by propodeum sculpture).The distinction between it and Dolichogenidea is particularly difficult (e.g., Fernandez-Triana et al. 2014), since many species of Dolichogenidea have a sculptured propodeum (with or without areola), and T1 is occasionally sculptured (although not as strongly as in Exoryza).T2 is also more or less broadly trapezoidal in shape in Exoryza, without the more or less sinuate hind margin many Dolichogenidea species have.All known sequences (DNA barcodes) of Exoryza cluster together in a NJ tree containing more than 50 genera and 17,518 sequences of world species of Microgastrinae (see Appendix S2 in Smith et al. 2013, page 41, sequences 4019-4041); but they are within a larger cluster containing Dolichogenidea (and a few specimens of Parapanteles and Apanteles which were most likely misidentified and are actually Dolichogenidea).With no recent, comprehensive phylogenetic study of Microgastrinae available, we consider it best to maintain Exoryza as a valid genus for the time being.

Diversity and host data
Five species of Exoryza were previously known (Mason 1981, Song and Chen 2003, Valerio et al. 2004, Rousse and Gupta 2013), and five new species are described below for a total of 10 species worldwide.Dolichogenidea oryzae Walker, 1994 could probably be included within Exoryza as well -based on the examined specimens (including holotype), host data (Chilo spp.) and distribution (West Africa), that species fits well within the generic concept used in the present paper.However, we have refrained from transferring D. oryzae to Exoryza due to the possibility that in the future Exoryza becomes a synonym of Dolichogenidea -see comments in previous paragraphs.But we include D. oryzae in the dichotomous key for Exoryza species provided below.
Exoryza seems to be almost cosmopolitan, with one species recorded from the Nearctic, six Neotropical, one Afrotropical, one Oriental, and one species found in both the Oriental and Eastern Palaearctic regions (Table 1).So far the only region where the genus has not been recorded is Australasia, but this may be a collecting artifact.We anticipate that additional species will be found as more collecting and study of world collections advances.

Exoryza mariabustosae
Male.As female but metacoxa and metatibia entirely dark brown to black, and T1-T2 narrower and smaller.Biology.Reared from Gelechiidae (an undetermined species with interim name 'gelJanzen01 Janzen319').
Etymology.Named in honor of María Margarita Bustos González for her enthusiasm in teaching her La Garita Vieja students to understand and protect the wild nature that occurs in her homeland.
Description.Detailed descriptions and images of the species available in Mason (1981) and Valerio et al. (2004).
Comments.Valerio et al. (2004) mentioned that the holotype of this species was in the CNC.However, the NMNH is the institution actually storing the type -see the original description for details (Mason 1981: 41).Exoryza monocavus Valerio & Whitfield, 2004 Exoryza monocavus Valerio & Whitfield, 2004: 3. Original description.
Description.A detailed description and images of the species are available in Valerio et al. (2004).
Male.As female, with slightly smoother sculpture.
Etymology.Named in honor of Richard Ashley from Huacas, Nicoya, for his serious enthusiasm for understanding and protecting the wild nature that occurs all around him, and specifically for causing 20 Guanacaste school children and their teachers to begin to share that enthusiasm.ACG, Sector Cacao, Sendero Cima, 1,460m, 10.93328, -85.45729, 18.xii.2008, coll. D. Janzen & W. Hallwachs.DNA voucher code: DHJPAR0031500.
Etymology.Named in honor of Rita Ashley from Huacas, Nicoya, for her serious enthusiasm for understanding and protecting the wild nature that occurs all around her, and specifically for causing 20 Guanacaste school children and their teachers to begin to share that enthusiasm.Paratypes.4 #F (CNC, NMNH).Costa Rica, Alajuela, ACG.DNA voucher codes: DHJPAR0033751, DHJPAR0049364, DHJPAR0052979, DHJPAR0053807.
Etymology.Named in honor of Rosa Iris Matarrita Díaz for her enthusiasm in teaching her Colonia Bolaños students to understand and protect the wild nature that occurs in her homeland.
Comments.Although the holotype was not examined, we studied two CNC specimens that were compared with the holotype by Mason (Mason 1981, Valerio et al. 2004).(Ratnasingham and Hebert 2007) for all barcoded specimens from each of the five species of Exoryza in ACG.Tip labels include species name, specimen accession number, sequence length and host species.
Etymology.Named in honor of Yeimy Cedeño Solís for her enthusiasm in encouraging her Area de Conservacion Tempisque community to understand and protect the wild nature that occurs in her homeland.

Figure 40 .
Figure 40.Neighbor-Joining (NJ -Saitou and Nei 1987) tree based on Kimura 2-parameter distances (K2P -Kimura 1980) made using BOLD(Ratnasingham and Hebert 2007) for all barcoded specimens from each of the five species of Exoryza in ACG.Tip labels include species name, specimen accession number, sequence length and host species.