Revision of the species of Lytopylus from Area de Conservación Guanacaste, northwestern Costa Rica (Hymenoptera, Braconidae, Agathidinae)

Abstract Thirty two new species of Lytopylus (Agathidinae) are described with image plates for each species: Lytopylus alejandromasisi sp. n., Lytopylus alfredomainieri sp. n., Lytopylus anamariamongeae sp. n., Lytopylus angelagonzalezae sp. n., Lytopylus cesarmorai sp. n., Lytopylus eddysanchezi sp. n., Lytopylus eliethcantillanoae sp. n., Lytopylus ericchapmani sp. n., Lytopylus gahyunae sp. n., Lytopylus gisukae sp. n., Lytopylus guillermopereirai sp. n., Lytopylus gustavoindunii sp. n., Lytopylus hartmanguidoi sp. n., Lytopylus hernanbravoi sp. n., Lytopylus hokwoni sp. n., Lytopylus ivanniasandovalae sp. n., Lytopylus johanvalerioi sp. n., Lytopylus josecortesi sp. n., Lytopylus luisgaritai sp. n., Lytopylus mariamartachavarriae sp. n., Lytopylus miguelviquezi sp. n., Lytopylus motohasegawai sp. n., Lytopylus okchunae sp. n., Lytopylus pablocobbi sp. n., Lytopylus robertofernandezi sp. n., Lytopylus rogerblancoi sp. n., Lytopylus salvadorlopezi sp. n., Lytopylus sangyeoni sp. n., Lytopylus sarahmeierottoae sp. n., Lytopylus sergiobermudezi sp. n., Lytopylus sigifredomarini sp. n., and Lytopylus youngcheae sp. n. A dichotomous key and a link to an electronic, interactive key are included. All specimens were reared from Lepidoptera larvae collected in Area de Conservación Guanacaste (ACG) and all are associated with ecological information including host caterpillar, collection date, eclosion date, caterpillar food plant, and locality. Neighbor-joining and maximum likelihood analyses of the barcode region of the mitochondrial cytochrome c oxidase subunit I gene (COI DNA barcode) were conducted to aid in species delimitation.

Although this article appears to be the second taxonomic revision of Lytopylus Forster, 1862 from Area de Conservación Guanacaste (ACG), the revision by Sharkey et al. (2011) employed the name Lytopylus in error and later Sharkey et al. (2016) transferred all of species described under Lytopylus in that paper to Aerophilus Szépligeti, 1902. This work focuses on specimens of Lytopylus reared from Lepidoptera larvae collected by Drs Janzen and Hallwachs and the team of ACG parataxonomists since 1978 in the ACG (Janzen et al. 2009. All are associated with ecological information including host caterpillar, collection date, eclosion date, caterpillar food plant, and locality. COI mitochondrial DNA barcodes for most specimens are deposited in the Barcode of Life Datasystem (BOLD) (http://www.boldsystems.org) (Hebert et al. 2003) and are equally available at http://janzen.sas.upenn. edu. The sequence data are publicly available through the Public Data Portal of BOLD (http://www.boldsystems.org/index.php/Public_BINSearch?searchtype=records).
We include an image plate for each species, a traditional identification key and a digital web-based interactive key; both have illustrations of morphological characters (https://www.dropbox.com/s/j9xongce1qrav5j/Revised%20Lytopylus%20Interac-tive%20key.zip?dl=0). A diagnoses and descriptions are provided for thirty-two new species and one previously described species.

Species concepts
We use Mayr's (1969) biological species concept, i.e., a species consists of a group of natural populations that are reproductively isolated from other groups. Because insect taxonomists usually work with dead specimens, delimitation of insect species is based on methods that indirectly infer reproductive isolation rather than direct observation, i.e., similarity in morphological, molecular (COI DNA sequences), geography, and host use data if recorded.
For the specimens which DNA sequences were not available in BOLD, DNA was extracted from individual legs at University of Kentucky (UKY) with Qiagen DNeasy Blood and Tissue Kit following the manufacturer's animal tissue protocol (Qiagen Inc., Chatsworth, California, USA).
COI was amplified from extracted DNA using the forward primer mlCOIintF (Leray et al. 2013) and reverse primer jgHCO2198 (Geller et al. 2013). Unique 9 bp tags, designed using Barcode Generator (available from http://comailab.genomecenter. ucdavis.edu/index.php/Barcode_generator) were attached to the primers so that each sequence could be traced to its parent specimen by the unique combination of tags. PCR was performed using Takara reagents consisted of 10X buffer, 2.5 μM nucleotides, 1 μM of each primer, 0.125 U Takara Ex Taq, 2 μL template DNA and enough dd H 2 O for a total reaction volume of 25 μL. We followed the "touchdown" thermal cycling protocol for these primers as outlined in Leray et al. (2013).
COI PCR DNA products, in addition to those from BOLD, were sequenced on an Illumina MiSeq system at the UKY Genomics Core Laboratory.

DNA assembly and phylogenetic analysis
Individual directional reads were downloaded from BOLD (produced by Sanger sequencing) and were edited and assembled using Geneious Pro (v. 6.1.6;Drummond et al. 2010) with the default settings. Edited sequences were stored in the NEXUS file format. The three sequences produced by NGS at UKY were included in the file of edited sequences. NGS sequencing data was assembled using PEAR (Zhang et al. 2013) and demultiplexed using custom Phython scripts. Among all bidirectional reads from each specimen, the 1 st and 2 nd most numerous reads were manually retrieved from the output file. The sequences were then queried against the GenBank nucleotide library using NCBI BLAST (https://blast.ncbi.nlm.nih.gov/ Blast.cgi) and those that were highly similar to Lytopylus specimens were retained. Finally, three COI sequences were exported from the FASTQ file and added to the file of edited sequences. The multiple sequence alignment was assembled on the MAFFT server (http://www.ebi.ac.uk/Tools/msa/mafft/; v. 7; Katoh et al. 2013) using the default settings.
A NJ tree (Saitou and Nei 1987) was constructed by using PAUP* (v. 4.0β10; Swofford 2003) using the p-distance setting. ML analyses were performed using Garli (v. 2.01;Zwickl 2006). For ML, the data were partitioned by codon position for COI (three partitions). We applied the most complex model available (GTR+I+G; Rodriguez et al. 1990) to each partition as per recommendations of Huelsenbeck and Rannala (2004) for likelihood-based analyses. Garli applies separate parameter estimates to each partition. A 20-replicate ML analysis was performed using default settings. Additionally, a ML bootstrap analysis (minimum 500 replicates) was conducted to assess nodal support (Garli, default settings). The COI data set analyzed herein is available from the senior author upon request.

Host use
Besides the notes included here additional data can be accessed at http://janzen.sas. upenn.edu (Janzen et al. 2009).

Species delimitation
The NJ tree and the tree of highest log-likelihood from 20 ML search reps in Fig. 1 and Fig. 2 were based solely on COI. These trees were constructed solely to assist in the delimitation of species. Molecular species concepts were initially based on the NJ tree and were compared to the best ML tree with COI data. The 2% genetic distance cut-off, which has been a conventional threshold for species delimitation using COI barcodes (Jones et al. 2011) and has been used in the Barcode Index Numbers (BINs) (http://www.barcodinglife.org/index.php/Public_BarcodeIndexNumber_Home), was used to cluster putative species (Smith et al. 2013). Morphological and host use data were then employed to make final decisions when genetic distances between putative molecular species were near the 2% threshold or below it, as is necessary for other groups of insects (e.g., Janzen et al. 2017).

Species delimitation
The NJ tree and the highest log-likelihood ML tree with COI data both suggest twenty-eight molecular species, and twenty-eight putative species were clustered using the 2% genetic distance cut-off.
Before running the molecular analyses I.K. and M.S. independently sorted the specimens to morphospecies and had error rates of 62% and 54% respectively based on our final species delimitations. All possible types of errors were discovered, i.e., clumping, splitting, and both clumping and splitting (mixing the members of two or more species). In contrast the molecular species concepts matched with our final species delimitations at 96.6%. The implications for previous taxonomic Figure 1. The NJ tree of the COI DNA barcode region for twenty-nine of the thirty-three Lytopylus species treated here. Triangles represent collapsed clades; their lengths (measured horizontally) represent the distance from the most basal node to the apex of the longest branch. The number of specimens in each triangle is given in parentheses following the species name. The node labeled with a red "A" is discussed in the text. treatments of braconids (and other speciose small tropical insects) based solely on morphology are dire.
The sole incongruity between molecular species concepts and final species concepts concerned L. sigifredomarini and L. guillermopereirai. The genetic variation between these two species was 0.4%, and the ML tree grouped them together (Fig. 2, node A). We, however, delimited them as separate species because they are morphologically distinct in their strikingly different color patterns, and the NJ tree recovered these two species as monophylic sister taxa. (Fig. 1, node A). In addition, they attack different species of host caterpillars with different feeding niches. Tree of highest log-likelihood from 20 ML search reps of the COI data set. Terminals with bold-faced type indicate species described herein. ML bootstrap values appear above the branches. Triangles represent collapsed clades; their lengths (measured horizontally) represent the distance from the most basal node to the apex of the longest branch. The number of specimens in each triangle is given in parentheses following the species name. The node labeled with a red "A" is discussed in the text.
The four species (L. alejandromasisi, L. ivanniasandovalae, L. josecortesi, L. mariamartachavarriae) for which genetic data were not available were delimited using morphological and host data.  areola of propodeum with well-defined margins. Anterior transverse carina of propodeum reaching the lateral margin. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.1 times longer than wide.

Systematics
Female. Unknown. Etymology. Lytopylus alejandromasisi is named in honor of Alejandro Masis in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Biology. Reared one time from Gelechiidae "same as 93-SRNP-3345.1" feeding on very new leaves of Bursera tomentosa (Burseraceae) in ACG dry forest at 280 m elevation.
Description. Holotype: female. Body length 4.4 mm. Fore wing length 5.1 mm. Fore wing mostly infuscated. Pronotum entirely pale (yellow to orange). Scutellar sulcus with one median longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum not reaching the lateral margin. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.1 times longer than wide. Ovipositor about same length as body.
Males. Similar to holotype except for face. Face usually paler than holotype. Etymology. Lytopylus alfredomainieri is named in honor of Alfredo Mainieri in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Biology. Reared five times from two species of Olethreutes (Olethreutinae, Tortricidae) leaf-tiers feeding on mature leaves of Meliosma glabrata (Sabiaceae) in ACG cloud forest edge at 1220 to 1276 m elevation.
Description. Holotype: female. Body length 6.2 mm. Fore wing length 5.5 mm. Fore wing mostly infuscated. Scutellar sulcus lacking longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum reaching the lateral margin. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.5 times longer than wide. Ovipositor about same length as body.
Male. Unknown. Etymology. Lytopylus anamariamongeae is named in honor of Ana Maria Monge in recognition of her participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Biology. Reared one time from Antaeocerconota Janzen433 (Depressariidae) a leaftier feeding on mature leaves of Inga punctata (Fabaceae) in ACG dry forest -rain forest ecotone at 540 m elevation.

Male. Unknown.
Etymology. Lytopylus angelagonzalezae is named in honor of Angela González Grau in recognition of her participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Biology. Reared two times from Anacampsis Janzen353 (Anacampsinae, Gelechiidae) feeding on two species of Rutaceae in ACG dry forest -rain forest ecotone at 280 to 825 m elevation.
Type material. Diagnosis. Fore wing mostly infuscated; scutellar sulcus with one median longitudinal carina; median tergites entirely melanic; lateral tergites one and two entirely white.
Description. Holotype: female. Body length 5.7 mm. Fore wing length 5.4 mm. Fore wing mostly infuscated. Scutellar sulcus with one median longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum absent. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.5 times longer than wide. Ovipositor slightly longer than body.
Male. Fore wing with a slight yellow tinge. Body color pattern similar to holotype, but slightly lighter.
Etymology. Lytopylus cesarmorai is named in honor of Cesar Mora in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Biology. Reared two times from Stenoma BioLep82 (Depressariidae) feeding on mature leaves of Apeiba membranacea (Malvaceae) in ACG rain forest at 527 m elevation.
Biology. Reared 48 times from seven species of dichomeridine Gelechiidae feeding on seven species of mature leaves of Malvaceae, Violaceae, Rubiaceae, Asteraceae, and Fabaceae growing in ACG rain forest at 240 to 645 m elevation.
Description. Holotype: female. Body length 6.0 mm. Fore wing length 5.5 mm. Fore wing with one black band. Scutellar sulcus with one median longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum absent. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.2 times longer than wide. Ovipositor about same length as body.
Males. Similar to holotype except for fore legs color. Fore legs usually less melanic. Etymology. Lytopylus eddysanchezi is named in honor of Eddy Sánchez in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Biology. Reared 11 times from one species leaf-tier in the Depressariidae, feeding on mature leaves of Meliosma glabrata (Sabiaceae) in ACG rain forest at 540 to 645 m elevation.
Type material. Diagnosis. Fore wing mostly infuscated; mesoscutum entirely melanic; anterior transverse carina of propodeum absent; median areola of propodeum narrow and not closed posteriorly; median areola length 11x its width, lateral longitudinal carinae of median tergite 1 well-defined; median tergites entirely reddish orange.
Female. Unknown. Etymology. Lytopylus eliethcantillanoae is named in honor of Elieth Cantillano in recognition of her participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Type material. Diagnosis. Fore wing with two black bands; pronotum anteriorly black and posteriorly pale.
Description. Holotype: female. Body length 5.8 mm. Fore wing length 5.7 mm. Fore wing with two black bands. Scutellar sulcus lacking longitudinal carina. Anterior transverse carina of propodeum absent. Median areola of propodeum with welldefined margins. Median areola of propodeum narrow. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.7 times longer than wide. Ovipositor slightly longer than body.
Male. Body color similar to holotype. Body length slightly shorter than holotype. Etymology. Named in honor of Dr Eric G. Chapman, research analyst in the Department of Entomology at the University of Kentucky, for his kindly advice on molecular systematics and phylogenetics.
Biology. Reared five times but only from the leaf-tier Stenoma adytodes (Depressariidae) feeding on mature leaves of Pouteria juruana (Sapotaceae) at the intersection of the ACG dry forest and rain forest ecosystems at 722 m elevation.
Description. Holotype: female. Body length 7.1 mm. Fore wing length 6.4 mm. Fore wing mostly infuscated. Scutellar sulcus with one median longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum reaching the lateral margin. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.4 times longer than wide. Ovipositor slightly longer than body.
Males. Similar to holotype. Etymology. Lytopylus gahyunae is named in honor of Gahyun Park, wife of the first author.  Biology. Reared 43 times from six species of Antaeotricha (40) and Stenoma (2) (Depressariidae) feeding on mature leaves of 3 species of Guarea and 1 of Trichilia (Meliaceae) in ACG rain forest at 380 to 620 m elevation.  Fig. 13 Diagnosis. Vertex of head mostly pale; fore wing mostly infuscated with a quadrate second submarginal cell; mesoscutum entirely pale (yellow to orange); median areola of propodeum length 15x its width; anterior transverse carina of propodeum not reaching the lateral margin; median tergites entirely pale (yellow to orange).
Description. Holotype: female. Body length 5.0 mm. Fore wing length 5.0 mm. Fore wing mostly infuscated with a quadrate second submarginal cell. Scutellar sulcus with one median longitudinal carina. Median areola of propodeum length 15x its width with well-defined margins. Anterior transverse carina of propodeum not reaching the lateral margin. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.2 times longer than wide. Ovipositor longer than metasoma, but shorter than body.
Male. Unknown. Etymology. Lytopylus gisukae is named in honor of Gisuk Lee, mother-in-law of the first author.
Type material.  Diagnosis. Apical flagellomeres brown not distinctly paler than subapical flagellomeres; vertex of head entirely pale; fore wing mostly infuscated with a triangular second submarginal cell; mesoscutum entirely pale (yellow to orange); median areola spindleshaped; anterior transverse carina of propodeum absent; median tergites entirely pale (yellow to orange).
Description. Holotype: female. Body length 4.8 mm. Fore wing length 4.5 mm. Fore wing mostly infuscated. Scutellar sulcus with one median longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum absent. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.3 times longer than wide. Ovipositor slightly longer than body.
Male. Unknown. Etymology. Lytopylus gustavoindunii is named in honor of Gustavo Induni in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Biology. Reared 12 times from two species of palm-feeding (Geonoma, Chamaedorea) Depressariidae (Stenoma Janzen142 and Stenoma Janzen284) in the understory of ACG rain forest from 645-742 m elevation. Diagnosis. Fore wing with one black band; mid tibia black basally and distally, yellow at mid-length. Description. Holotype: female. Body length 4.3 mm. Fore wing length 4.3 mm. Fore wing with one black band. Scutellar sulcus with one median longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum absent. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.3 times longer than wide. Ovipositor longer than metasoma, but shorter than body.
Male. Unknown. Etymology. Lytopylus hartmanguidoi is named in honor of Hartman Guido in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Biology. Reared five times from three species leaf-tiers in the Depressariidae, feeding on mature leaves of Hiraea reclinata (Malpighiaceae) at the intersection of the ACG dry forest and rain forest ecosystems at 540 m elevation.
Female. Unknown. Etymology. Lytopylus hernanbravoi is named in honor of Hernan Bravo in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Description. Holotype: male. Body length 5.8 mm. Fore wing length 5.0 mm. Fore wing with a slight yellow tinge. Scutellar sulcus with four longitudinal carinae. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum absent. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 as long as wide.
Female. Unknown. Etymology. Lytopylus ivanniasandovalae is named in honor of Ivannia Sandoval in recognition of her participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Biology. Reared one time from Dichomerus Janzen703 (Dichomeridinae, Gelechiidae) tying and feeding on mature leaves of Neurolaena lobata (Asteraceae) in ACG rain forest at 660 m elevation.  Diagnosis. fore wing mostly infuscated; pronotum mostly yellow; mesoscutum mostly pale (yellow to orange); anterior transverse carina of propodeum reaching the lateral margin; median tergites mostly pale posterior tergum black.
Description. Holotype: female. Body length 4.9 mm. Fore wing length 4.6 mm. Fore wing mostly infuscated. Scutellar sulcus with one median longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum reaching the lateral margin. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 0.9 times longer than wide. Ovipositor longer than metasoma, but shorter than body.

Males.
Occiput usually more melanic. Median tergites usually mostly pale with posterior terga black.
Etymology. Lytopylus johanvalerioi is named in honor of Johan Valerio in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Biology. Reared six times from two species of Cerconota leaf-tiers in the Depressariidae, feeding on mature leaves of three species of Inga (Fabaceae) in ACG rain forest at 540-645 m elevation.
Male. Similar to holotype. Etymology. Lytopylus josecortesi is named in honor of José Cortés in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Description. Holotype: female. Body length 6.7 mm. Fore wing length 6.3 mm. Fore wing mostly infuscated. Scutellar sulcus lacking longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum reaching the lateral margin. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 as long as wide. Ovipositor longer than metasoma, but shorter than body.
Male. Unknown. Variation. Paratype propodeum mostly pale. Etymology. Lytopylus luisgaritai is named in honor of Luis Garita in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Description. Holotype: female. Body length 4.4 mm. Fore wing length 4.0 mm. Fore wing hyaline. Scutellar sulcus with three longitudinal carinae. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum reaching the lateral margin. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 0.9 times longer than wide. Ovipositor longer than metasoma, but shorter than body.
Male. Similar to holotype, but median tergites mostly pale with three posterior terga melanic.
Etymology. Lytopylus mariamartachavarriae is named in honor of María Marta Chavarría in recognition of her participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Description. Holotype: female. Body length 5.1 mm. Fore wing length 4.9 mm. Fore wing mostly infuscated. Scutellar sulcus with one median longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum not reaching the lateral margin. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.1 times longer than wide. Ovipositor longer than metasoma, but shorter than body.
Males. Body length usually shorter than holotype. Median tergites mostly melanic. Variation. Female anterior head varies from mostly pale to mostly melanic. Etymology. Lytopylus miguelviquezi is named in honor of Miguel Viquez in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica. Biology. Reared 58 times from the Dichomeris designatella complex (21), gel-Janzen01 Janzen179 (13), and gelJanzen01 Janzen485 (16), all leaf tying dichomeridine Gelechiidae feeding on mature leaves of two species of Erythroxylum (Erythroxylaceae) and two species of Rinorea (Violaceae) in ACG rain forest-dry forest ecotone, and rain forest at 109 to 540 m elevation.
Type material.  Diagnosis. Vertex of head entirely yellow; fore wing mostly infuscated with a quadrate second submarginal cell; mesoscutum mostly or entirely pale (yellow to orange); median areola of propodeum kite-shaped; anterior transverse carina of propodeum not reaching the lateral margin; median tergites entirely pale (yellow to orange).
Description. Holotype: female. Body length 4.9 mm. Fore wing length 4.9 mm. Fore wing mostly infuscated. Scutellar sulcus with one median longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum not reaching the lateral margin. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 as long as wide. Ovipositor longer than metasoma, but shorter than body.
Males. Vertical of head and occiput usually mostly melanic. Body length usually shorter than holotype. Median tergites mostly melanic.
Variation. Female occiput varies from entirely pale to mostly pale. Male hind femur color varies from mostly pale to black and pale with a similar percentage of each color.
Etymology. Lytopylus motohasegawai is named in honor of Motohiro Hasegawa in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Biology. Reared 36 times from gelJanzen01 Janzen28, a leaf-tier in the Gelechiidae feeding on mature leaves of two species of Roupala (Proteaceae) in ACG rain forest at 415 to 740 m elevation.
Description. Holotype: female. Body length 6.8 mm. Fore wing length 6.3 mm. Fore wing mostly infuscated. Scutellar sulcus with one median longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum absent. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.5 times longer than wide. Ovipositor about same length as body. Males. Similar to holotype except for median tergite color. Median tergites usually mostly pale with posterior three terga melanic.
Etymology. Lytopylus okchunae is named in honor of Okchun Kim, grandmother of the first author.
Male. Unknown. Etymology. Lytopylus pablocobbi is named in honor of Pablo Cobb in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica. Biology. Reared one time from elachJanzen01 Janzen640 (Depressariidae), a stenomine leaf-tier feeding on mature foliage of Bunchosia odorata (Malpighiaceae) in ACG dry forest -rain forest ecotone at 722 m elevation.
Type material.  Biology. Reared only one time and from the leaf-tier Stenoma Janzen687 (Depressariidae) feeding on mature leaves of Pouteria exfoliata (Sapotaceae) at the intersection of the ACG dry forest and rain forest ecosystems at 540 m elevation.
Description. Holotype: female. Body length 5.5 mm. Fore wing length 5.3 mm. Fore wing mostly infuscated. Scutellar sulcus with one median longitudinal carina. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum reaching the lateral margin. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 as long as wide. Ovipositor longer than metasoma, but shorter than body.
Male. Unknown. Etymology. Named in honor of Sangyeon Park, father-in-law of the first author. Biology. Reared one time from elachJanzen01 Janzen847 (Depressariidae) as a leaf-tier feeding on mature leaves of Senegalia tenuifolia (Fabaceae) in ACG rain forest at 527 m elevation. Male. Unknown. Etymology. Named in honor of Sarah Meierotto, graduate student in the Department of Entomology at the University of Kentucky, for her assistance.
Description. Holotype: male. Body length 3.9 mm. Fore wing length 3.6 mm. Fore wing hyaline. Fore wing RS+Ma more complete. Scutellar sulcus with four  longitudinal carinae. Median areola of propodeum with well-defined margins. Anterior transverse carina of propodeum reaching the lateral margin. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.1 times longer than wide.

Female. Unknown.
Variation. Male mesoscutum varies from less melanic to mostly pale. Male propodeum varies bicolored to entirely pale.
Etymology. Lytopylus sergiobermudezi is named in honor of Sergio Bermúdez in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Female. Unknown. Etymology. Lytopylus sigifredomarini is named in honor of Sigifredo Marín in recognition of his participation in the collaborative development of the ICE-ACG geothermal project of Pailas II, northwestern Costa Rica.
Biology. Reared three times from Antaeotricha Janzen224 (Stenomatinae, Depressariidae) feeding on mature leaves of Hirtella media (Chrysobalanaceae) in ACG rain forest at 410 to 620 m elevation.
Description. Holotype: female. Body length 8.9 mm. Fore wing length 8.1 mm. Fore wing with one black band. Scutellar sulcus with one median longitudinal carina. Anterior transverse carina of propodeum not reaching the lateral margin. Median areola of propodeum with well-defined margins. Lateral longitudinal carinae of median tergite 1 well-defined. Median syntergite 2+3 1.5 times longer than wide. Ovipositor slightly longer than body.
Males. Body length usually shorter than holotype. Hind femur varies from mostly pale to mostly melanic.
Etymology. Named in honor of Youngche Choi, mother of the first author. Biology. Reared five times from two species of stenomatine Depressariidae leaftiers feeding on mature leaves of Calophyllum brasiliense (Calophyllaceae) in ACG rain forest at 540 to 740 m elevation.  . This study has been supported by the Government of Canada through its ongoing support of Genome Canada, the Biodiversity Institute of Ontario, and the Ontario Genomics Institute (2008-0GI-ICI-03)(MAS), and by a Discovery Grant from Natural Sciences and Engineering Research Council of Canada (MAS). Funding was also provided by Hatch projects KY008041 and KY008065 (to MJS). Many thanks to HIC lab member Sarah Meierotto for their expertise and time in editing this manuscript. The information reported in this paper (No. 17-08-081) is part of a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director.