Revision of the Mesoamerican species of Calolydella Townsend (Diptera: Tachinidae) and description of twenty-three new species reared from caterpillars in Area de Conservación Guanacaste, northwestern Costa Rica

Abstract Background Twenty-three new species of the genus Calolydella Townsend, 1927 (Diptera: Tachinidae) are described, all reared from multiple species of wild-caught caterpillars across a wide variety of families (Lepidoptera: Crambidae; Erebidae; Geometridae; Hesperiidae; Lycaenidae; Nymphalidae; Pieridae; Riodinidae; and Sphingidae). All caterpillars were collected within Area de Conservación Guanacaste (ACG), in northwestern Costa Rica. This study provides a concise description of each new species using morphology, life history, molecular data, and photographic documentation. In addition to the new species, we also provide a generic redescription and revised key to species of the genus Calolydella from Central and South America. New information The following 23 new species of Calolydella are described by Fleming and Wood: C. adelinamoralesae sp. n., C. alexanderjamesi sp. n., C. argentea sp. n., C. aureofacies sp. n., C. bicolor sp. n., C. bifissus sp. n., C. crocata sp. n., C. destituta sp. n., C. discalis sp. n., C. erasmocoronadoi sp. n., C. felipechavarriai sp. n., C. fredriksjobergi sp. n., C. inflatipalpis sp. n., C. interrupta sp. n., C. nigripalpis sp. n., C. omissa sp. n., C. ordinalis sp. n., C. renemalaisei sp. n., C. susanaroibasae sp. n., C. tanyadapkeyae sp. n., C. tenebrosa sp. n., C. timjamesi sp. n., C. virginiajamesae sp. n. Lydella frugale Curran, 1934 is proposed as a new synonym of Pygophorinia peruviana Townsend, 1927, syn. n., under the combination Calolydella frugale (Curran, 1934), comb. n.


Introduction
The New World genus Calolydella Townsend, 1927 (Tachinidae: Blondeliini) was erected for one male and several female specimens collected by Townsend himself at Itaquaquecetuba (now a suburb of São Paolo), Brazil and described as Calolydella geminata Townsend, 1927. When Wood (1985 revised the tribe Blondeliini, he synonymized three other genera under Calolydella: Olindopsis Townsend, 1927, Pygophorinia Townsend, 1927, and Prodexodes Townsend, 1927. These synonymies raised the number of species in the genus from one to 13 (Wood 1985).
Here, we describe 23 new species of Calolydella reared from wild-caught caterpillars collected as part of the ongoing inventory of Lepidoptera larvae (and their parasitoids) in the terrestrial portion of Area de Conservación Guanacaste (ACG), an area of 120,000 hectares in northwestern Costa Rica . Each new species is distinguished by morphological differences, differences in CO1 (cox1 or cytochrome oxidase 1) gene sequences, and comparison (by DMW and AJF) with previously described species of Calolydella. Although many of the species of Calolydella are common throughout the Neotropics (Wood 1985), none of the previously known species have been found in ACG to date. Individual and comparative details of the parasitization ecology of these flies will be treated in later papers, once the overall knowledge of the caterpillar-attacking tachinids found in ACG is more complete (Smith et al. 2006, Smith et al. 2007, Smith et al. 2008, Smith et al. 2012, Fleming et al. 2014a, Fleming et al. 2014b, Fleming et al. 2015a, Fleming et al. 2015b, Fleming et al. 2015c, Fleming et al. 2015d, Fleming et al. 2016a, Fleming et al. 2016b).
All DHJPARxxxxxxx-coded tachinids had one leg removed for DNA barcoding and couriered to the Center for Biodiversity Genomics of the Biodiversity Institute of Ontario (BIO) at the University of Guelph (Canada). All associated data and successful barcodes are permanently and publicly deposited in the Barcode of Life Data System (BOLD) (Ratnasingham and Hebert 2007), and a select set of these data are subsequently migrated to GenBank. Each barcoded specimen also receives accession numbers from the Barcode of Life Data System (BOLD) and GenBank, respectively. The dynamic nature of the inventory means that it is continually adding new specimens; newly added Calolydella specimens can be found by searching for the genus in BOLD.
All inventoried specimens discussed herein were collected under Costa Rican government research permits issued to DHJ; the Tachinidae samples were exported under permit by DHJ from Costa Rica to their final depository in the CNC. Tachinid identifications for the inventory are done by DHJ in coordination with a) visual inspection of morphology by AJF and DMW, b) DNA barcoding by MAS and BIO, and c) databasing and association with host caterpillars by DHJ and WH through the inventory itself.
The date of capture of fly specimens is the date of eclosion of the fly and not the date of capture of the caterpillar. The fly eclosion date is much more representative of the time when that species is on the wing than is the date of capture of the parasitized caterpillar. The "collector" is the parataxonomist who collected the caterpillar, rather than the person who later retrieved the newly eclosed fly and processed it by freezing, pinning, labeling and oven-drying. The biology and parasitization rates of individual tachinid species will be the subject of later papers. The holotypes of the new species described herein are deposited in the Diptera collection of the CNC, Ottawa, Canada.

Imaging
Habitus and terminalia photographs were taken using the methods outlined in Fleming et al. (2014a). Raw image files were first processed with Adobe Photoshop CS6, then digitally stacked to produce a final composite image using Zerene Stacker Software v1.04.
Adult flies were dissected following standard practice (O'Hara 1983). The morphological terminology follows Cumming and Wood (2009). If only one male was available, it was designated as the holotype and not subjected to dissection.

Interim names for undescribed host species
Names of undescribed host species follow a standardized, interim naming system used for taxonomic units considered to be distinct species and identified by a combination of DNA barcode, morphology, and host-plant information. The interim names are given in the format "Eois Janzen52", where the species epithet is composed of the name of the taxonomist who first singled out the species and a number. This prevents confusion with already described species while maintaining traceability of each undescribed species within the ACG project. dilation weak or absent; palps normally slender and yellow, some species with black or inflated palps; postpedicel of similar length in both sexes; arista minutely pubescent, slightly thickened at base. Thorax: prosternum setose; proepisternum bare; postpronotum with 2-4 setae in a narrow triangle or a straight line (rarely inner basal seta absent); acrostichal setae 1-3:3-4; dorsocentral setae 1-3:2-3; intra-alar setae 1-3:2-3; supra-alar setae 1-2:3; katepisternal setae 2-3 [Wood (1985) described the genus as having three katepisternal setae; however, we show that this trait can vary in females]; lateral scutellar setae shorter than subapical setae and curved medially; fore tibia with one posterior seta; mid tibia with one anterodorsal seta; wings ranging from smoky gray to pale translucent brown; wing vein R with 2-7 setulae dorsally at base, these sometimes extending to crossvein R-M. Abdomen: ST1+2 with one pair of median marginal setae, mid-dorsal depression not extending to hind margin of syntergite; 3rd and 4th abdominal tergites with 1-2 pairs of discal setae. Abdominal tergites marginally banded with gold pollinosity; in some species banding interrupted by a dark median stripe; male abdomen elongated, conical, tapering to a sharp point; in females, abdomen rounded and downwardly curved; in some species the coloration and pattern of pollinosity are sexually dimorphic.

Diagnosis
Calolydella are slender-bodied flies characterized by an abdomen longer than it is wide, with bold abdominal markings of yellow-gold pollinose stripes on a black background. According to Wood (1985), they lack distinctive chaetotactic characters and are difficult to characterize to genus except by their general appearance. Wood (1985) suggested the presence of only two pairs of presutural acrostichal setae as a diagnostic character; however, our current examination of this trait suggests that it varies among species, as does the rest of the thoracic chaetotaxy. Sternite 5 with only short hair-like setae, all more or less equal in size. Sternites 6 and 7 of female larger and more heavily sclerotized.
Female (Fig. 1d, e, f). Length: 5-7mm. As male, except for the following characters: fronto-orbital plate 1.6X as wide as in male; 2:3 acrostichal setae; wing vein R with 6-7 setulae extending to crossvein R-M, and ground color of abdomen appearing slightly darker than in male.

Diagnosis
Calolydella adelinamoralesae can be distinguished from all other species of Calolydella by the following combination of traits: parafacial all silver, frontal setae extending to base of postpedicel, anatergite bare, and T4 with 1-2 pairs of discal setae.

Etymology
The specific epithet is in honor of Adelina Morales Chaves of La Cruz, Guanacaste, Costa Rica, in recognition of the many years she dedicated to sorting ACG Malaise trap samples, many of which contained Calolydella flies.
Female: not known at this time.

Diagnosis
Calolydella alexanderjamesi can be distinguished from all other species of Calolydella by the following combination of traits: parafacial mostly gold, fronto-orbital plate with small black setulae interspersed among frontal setae, anatergite bare, and thoracic pollinosity gold on both dorsal and lateral surfaces.

Etymology
The specific epithet is in honor of Alexander James of Levittown, Pennsylvania, in recognition of the moral and family support of his mother, Tanya Dapkey, in her efforts curating and preparing ACG parasitoid flies for DNA barcoding.

Calolydella argentea
Male: not known at this time.

Diagnosis
Calolydella argentea can be distinguished from all other species of Calolydella by the following combination of traits: parafacial and fronto-orbital plate entirely silver, frontal setae extending to base of postpedicel with intermingled small setulae, 3:3 acrostichal setae, anterior 2/3 of abdominal tergites with silver pollinose bands, anatergite with three or more hair-like setae, and wing vein R with 6-7 setulae, almost reaching to crossvein R-M.

Etymology
The specific epithet is derived from the Latin adjective "argentum", meaning silver, in reference to its entirely silver parafacial and fronto-orbital plate, a feature unique to this species.
Male: not known at this time.

Diagnosis
Calolydella aureofacies can be distinguished from all other species of Calolydella by the following combination of traits: parafacial mostly gold, female palps black basally, anatergite with three or more hair-like setae, and scutellar discal setae absent.

Etymology
The specific epithet is derived from the Latin adjective "aurum", meaning gold, and " facies", meaning face, in reference to its gold parafacial and fronto-orbital plate.
Male: not known at this time.

Diagnosis
Calolydella bicolor can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate entirely gold, parafacial entirely silver, anatergite with three or more hair-like setae, and gold abdominal pollinose bands interrupted by a median dark stripe.

Etymology
The specific epithet is derived from the Latin adjective "bicolor", meaning two-toned, in reference to its gold fronto-orbital plate and silver parafacial.
Female (Fig. 7d, e, f). Length: 5mm. As male, except for the following characters: fronto-orbital plate 3X as wide as in male; frontal vitta not tapered, apically 1.2X as wide as in male; two katepisternal setae.

Diagnosis
Calolydella bifissus can be distinguished from all other species of Calolydella by the following combination of characters: two bold thoracic vittae, abdominal pollinosity concolorous on both dorsal and ventral surfaces, anatergite with a small tuft of hair-like setae, and abdominal ground color dark brown-orange.

Etymology
The specific epithet is derived from the Latin adjective "bifissus", meaning cleft or cloven, in reference to cleft nature of the cerci, which are fused basally, a character state unique to this species.

4+5
Female: not known at this time.

Diagnosis
Calolydella crocata can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate mostly gold-pollinose but with some silver pollinosity and sparsely setulose, thorax with four vittae visible only under certain angles of light, anatergite bare, and abdomen with transverse marginal pollinose bands interrupted by a thin dark median stripe.

Etymology
The specific epithet is derived from the Latin adjective "crocatum", meaning yellow, in reference to the gold pollinosity present over most of the body of the new species.
Female (Fig. 11d, e, f). Length: 5-6mm. As male, except for the following characters: fronto-orbital plate 2X as wide as in male; and T3 with one pair of median marginal setae and one pair of discal setae.

Diagnosis
Calolydella destituta can be distinguished from all other species of Calolydella by the following combination of traits: anatergite with a small tuft of setae, pollinose bands on

Etymology
The specific epithet is derived from the Latin adjective "destituta", meaning absent or lacking, in reference to the absence of median marginal setae on T3, a character state unique to this species.
Female: not known at this time.

Diagnosis
Calolydella discalis can be distinguished from all other species of Calolydella by the following combination of traits: parafacial mostly gold, fronto-orbital plate with a single a b c Figure 13.
a: habitus in dorsal view b: head in frontal view c: habitus in lateral view row of small black setulae outside of the frontal setae, anatergite with a small tuft of three or more hair-like setae, and T3 and T4 each with two pairs of discal setae.

Etymology
The specific epithet refers to the two pairs of discal setae on T3 and T4.

Diagnosis
Calolydella erasmocoronadoi can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate bare, up to 50% silver pollinose, thoracic vittae visible only under certain angles of light, anatergite with three or more setae arranged in a small tuft, wing vein R with 6-7 setulae, and abdomen with uninterrupted transverse marginal pollinose bands.

Diagnosis
Calolydella felipechavarriai can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate almost bare, anatergite with at most two hair-like setae, and T4 with one pair of discal setae.

Etymology
The specific epithet is in honor of Luis Felipe Chavarria Diaz of ACG, Guanacaste, Costa Rica, in recognition of his high quality accounting, photography and administrative support of the Guanacaste Dry Forest Conservation Fund (www.gdfcf.or g) and of the ACG parataxonomists who collected and reared the caterpillars from which the new species of ACG Calolydella described here emerged.

Distribution
Costa Rica, ACG, Alajuela Province, Palomo, 96m.  Description Male (Fig. 19a, b, c). Length: 7-9mm. Head (Fig. 19b): frontal setae extending to base of postpedicel; fronto-orbital plate gold, with a single row of fine setulae outside of frontal setae; parafacial silver throughout. Thorax (Fig. 19a, c): gold on both dorsal and lateral surfaces; outermost two thoracic vittae, twice as wide as innermost two; postpronotum with three setae; 3:3 acrostichal setae; 3:3 dorsocentral setae; 2:3 intraalar setae; 2:3 supra-alar setae; three katepisternal setae; anatergite with three or more hair-like setae, often in a small tuft; scutellar discal setae situated as wide apart as subapical scutellar setae. Wing vein R4+5 with 6-7 setulae dorsally, reaching from base almost to crossvein R-M. Abdomen (Fig. 19a): ground color dark brown-orange, with uninterrupted transverse marginal pollinose bands; abdominal pollinosity gold dorsally, silver ventrally, with an orange spot lateroventrally at base of ST1+2; T3 with one pair of median marginal setae and two pairs of discal setae; T4 with a row of marginal setae and one pair of discal setae. Terminalia (Fig. 20): sternite 5 ( Fig. 20c) with two small lobes and a wide U-shaped median cleft, 0.47X the length of the sternite from lobe to apex; inner margin covered by dense pollinosity, appearing darker than surrounding cuticle; entire lobe of sternite with short setae, of varying lengths. Cerci (Fig. 20b), in dorsal view, separated by a narrow gap not widened at apex; each cercus very slightly tapered from its already narrow base; cercus long, slender and with an upward bend when viewed laterally; setose along its entire length. Surstylus (Fig. 20a) slender and digitiform with a strong divot at tip, giving it an overall bent look; appearing 0.2X shorter than cercus when viewed laterally; with short setae along entire length; tip of surstylus slightly curved inwards when viewed dorsally.

Diagnosis
Calolydella fredriksjobergi can be distinguished from all other species of Calolydella by the following combination of characters: parafacial mostly gold, anatergite with three or more hair-like setae arranged in a small tuft, two pairs of discal setae on T3 and one pair of discal setae on T4.

Etymology
The specific epithet is in honor of Fredrik Sjoberg of Runmaro Island, Sweden, in recognition of his wonderful book "The Fly Trap" (Sjöberg 2014, translation by Thomas Teal); a book that explains much about Malaise traps, which have captured so many ACG Tachinidae.

Ecology
Calolydella fredriksjobergi has been reared six times from six different species belonging to various families of Lepidoptera (Nymphalidae; Erebidae; and Crambidae), in rain forest, cloud forest, and dry forest ecosystems.

a b
c d e f Figure 19. Description Male (Fig. 21a, b, c). Length: 5-9mm. Head (Fig. 21b, e): frontal setae extending beyond base of postpedicel; fronto-orbital plate gold, sparsely setulose throughout; parafacial almost entirely gold (75-90% coverage). Thorax (Fig. 21a, c, d, f): pollinosity gold on both dorsal and lateral surfaces; or gold on dorsal surface and silver laterally (more than 50% coverage); thorax with outermost two vittae twice as wide as innermost two; postpronotum with three setae; 3:3 acrostichal setae; 3:3 dorsocentral setae; 2:3 intra-alar setae; 2:3 supra-alar setae; three katepisternal setae; anatergite with three or more hair-like setae, often in a small tuft; scutellar discal setae situated as wide apart as subapical scutellar setae. Wing vein R with at most 2-3 small setulae dorsally at base. Abdomen (Fig. 21a, d): ground color dark brown-orange; with uninterrupted transverse marginal pollinose bands; pollinosity gold dorsally, silver ventrally, and with an orange spot lateroventrally at base of ST1+2; T3 with one pair of median marginal setae and two pairs of discal setae; T4 with two pairs of discal setae. Terminalia (Fig.  22): sternite 5 ( Fig. 22c) with two small lobes and a wide U-shaped median cleft, 0.37X the length of the sternite from lobe to apex; inner margin covered by dense pollinosity, appearing darker than surrounding cuticle; entire lobe of sternite with sparse short setae, of varying lengths. Cerci (Fig. 22b), in dorsal view, separated by a narrow gap widening at apex; slender and straight; cercus, in lateral view, long, very slightly tapered from its already narrow base, setose along its basal half. Surstylus (Fig. 22a) subequal to length of cercus, turned slightly inward at tip, appearing as a small apical lobe when viewed dorsally; surstylus slender and digitiform when viewed laterally; with short setae along entire length; tip of surstylus not curved inwards when viewed dorsally.

Diagnosis
Calolydella inflatipalpis can be distinguished from all other species of Calolydella by the following combination of traits: fine setulae interspersed among frontal setae, palps normal in male but significantly inflated in female, anatergite with three or more hair-like setae arranged in a small tuft, and T3 and T4 each with two pairs of discal setae.

Etymology
The specific epithet is derived from the Latin adjective "inflatum", meaning inflated, and "palpus", meaning palp, in reference to the inflated, bulbous palpi of females of this species.

Diagnosis
Calolydella interrupta can be distinguished from all other species of Calolydella by the following combination of traits: parafacial at least 50% silver pollinose, two bold thoracic vittae, abdominal pollinosity interrupted by median stripe, anatergite with three or more setae arranged in a small tuft, and T3 with a complete row of marginal setae.

Etymology
The specific epithet is derived from the Latin adjective "interruptum", meaning breach or interruption, in reference to dark median stripe breaking up the gold marginal pollinosity on the abdomen of this species.
Female: not known at this time.

Etymology
The specific epithet is derived from the Latin adjective "nigrum", meaning black, and " palpus", meaning palp, in reference to the black palpi of males of this species.

Diagnosis
Calolydella omissa can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate gold, parafacial silver, abdomen with uninterrupted transverse silver pollinose bands across all tergites, anatergite bare (rarely setose), and T3 without discal setae.

Etymology
The specific epithet is derived from the Latin adjective "omissa", meaning dropped or absent, in reference to the absence of discal setae on T3, a character state unique to this species.

Diagnosis
Calolydella ordinalis can be distinguished from all other species of Calolydella by the following combination of characters: parafacial mostly gold, fronto-orbital plate with setulae sparsely interspersed among frontal setae, anatergite bare, and a complete row of discal setae on T4.

Etymology
The specific epithet is derived from the Latin adjective "ordinarius", meaning ordinary or complete, in reference to the complete, regular row of discal setae on T4, a character state unique to this species.
Female: not known at this time.

Diagnosis
Calolydella renemalaisei can be distinguished from all other other species of Calolydella by the following combination of characters: frontal setae extending to base of postpedicel, fronto-orbital plate sparsely setose througout, with four thoracic vittae, the outermost two twice as wide as the innermost two, anatergite with three or more hair-like setae arranged in a small tuft, and up to two pairs of discal setae on T4.

Etymology
The specific epithet is in honor of René Edmond Malaise  of Stockholm, Sweden, in recognition of his invention of the Malaise trap, which has captured so many ACG Tachinidae and other insects, as explained by Fredrik Sjöberg in his wonderful book "The Fly Trap" (Sjöberg 2014, translation by Thomas Teal), a book that explains much about Malaise traps.

Diagnosis
Calolydella susanaroibasae can be distinguished from all other species of Calolydella by the following combination of characters: fronto-orbital plate with a single row of fine setulae outside of frontal setae, thoracic pollinosity gold dorsally and on 50% of lateral surface, with outer two thoracic vittae twice as wide as inner two, anatergite with three or more hair-like setae arranged in a small tuft, and base of ST1+2 black lateroventrally.

Calolydella tanyadapkeyae
Male: not known at this time.

Diagnosis
Calolydella tanyadapkeyae can be distinguished from all other other species of Calolydella by the following combination of traits: parafacial all silver, frontal setae extending to base of postpedicel, thoracic pollinosity gold dorsally and over 50% of lateral surfaces, postpronotum with only two setae (inner basal seta absent), anatergite bare, and scutellar discal setae situated as wide apart as subapical scutellar setae.

Etymology
The specific epithet is in honor of Tanya Dapkey James of Levittown, Pennsylvania, in recognition of her efforts curating and preparing ACG parasitoid flies for DNA barcoding.
Female (Fig. 34d, e, f). Length: 6mm. As male, except for the following characters: fronto-orbital plate 1.41X as wide as in male; palps dark grayish-orange.

Diagnosis
Calolydella tenebrosa can be distinguished from all other species of Calolydella by the following combination of characters: frontal setae extending beyond base of postpedicel, palps black in males and dark grayish-orange in females, abdominal ground color dark brown-orange, anatergite bare, and abdominal T5 black apically in both sexes.

Etymology
The specific epithet Calolydella tenebrosa is derived from the Latin adjective " tenebrosum", meaning black or dark, in reference to the dark-colored palpi and darkcolored legs, character states found in both sexes of this species.

Diagnosis
Calolydella timjamesi can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate sparsely setulose to bare, frontal setae extending beyond base of postpedicel, abdominal ground color black, anatergite with three or more setae often in a small tuft, and scutellar discal setae slightly closer togetherthan subapical scutellar setae.

Etymology
The specific epithet is in honor of Tim James of Levittown, Pennsylvania, in recognition of the moral and family support of Tanya Dapkey in her efforts curating and preparing ACG parasitoid flies for DNA barcoding.

Diagnosis
Calolydella virginiajamesae can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate with a single row of small black setulae outside of frontal setae, parafacial silver-pollinose, anatergite with three or more setae arranged in a small tuft, and T3-T4 each with a single pair of discal setae.

Etymology
The specific epithet is in honor of Virginia James of Levittown, Pennsylvania, in recognition of the moral and family support of her mother, Tanya Dapkey, in her efforts curating and preparing ACG parasitoid flies for DNA barcoding.

Ecology
Calolydella virginiajamesae has been reared 34 times from various species of Lepidoptera from several families (Nymphalidae; Hesperiidae; Lycaenidae;

Key to the species of Calolydella of the Mesoamerican region
This key is confined to species of Calolydella occurring in the Mesoamerican biogeographical region. It is applicable to both males and females.

23
Fronto-orbital plate with a single row of setulae outside of frontal setae; postpronotum with three setae; wing vein R with 5-7 setulae dorsally, reaching from base almost to crossvein R-M; abdominal T4 with two pairs of discal setae (Fig. 32a, b,
-Fronto-orbital plate with setulae interspersed among frontal setae; postpronotum with two setae, inner basal seta absent; wing vein R with at most three small setulae dorsally at base; T4 with one pair of discal setae (Fig. 3a, b) Calolydella alexanderjamesi sp. n.

DNA Barcoding
A phylogenetic tree based on DNA barcodes was used to visually demonstrate the variation within and between species, and is presented in Fig. 39. Interested readers can consult the Barcode of Life Data System (BOLD) (Ratnasingham and Hebert 2007) for all information associated with each sequence (including GenBank accession numbers), derived from each individual specimen using the persistent DOI: dx.doi.org/10.5883/DS-ASCALOL.

Maximum Likelihood tree showing DNA barcode variation within ACG species of
Calolydella. The inter-and intra-specific variation within the DNA barcode region for the 23 species of Calolydella from ACG is illustrated using a Maximum Likelihood tree based on the General Time Reversible model (GTR) (Nei and Kumar 2000). GTR was selected due to the lowest BIC scores (Bayesian Information Criterion) of the Maximum Likelihood fits of 23 different nucleotide substitution models run in MEGA6 (Tamura et al. 2013). Tip labels include species name|specimen code|host family of each species. A habitus image in lateral view of each holotype is presented with the tree.