On the species formerly assigned to Schrankia Hübner (Erebidae: Hypenodinae) in the Western Hemisphere, with the revalidation of Hypenopsis Dyar and descriptions of three new species

Abstract: The genus Hypenopsis Dyar, 1913 (type species: Hypenodes macula Druce, 1891) is removed from synonymy with Schrankia Hübner, 1825 (Lepidoptera, Erebidae, Hypenodinae) and reinstated as valid. Hypenopsis and its type species H. macula (Druce, 1891) are both redescribed based on the first morphological study of the genitalia of its two syntypes, and a lectotype is designated. Three new species are described based on morphology and supported by COI sequence data: H. calusa J.-F. Landry & B. Landry, sp. nov., H. sonora J.-F. Landry & B. Landry, sp. nov., two North American species long misidentified as ‘Schrankia’ macula, and H. galapagensis B. Landry & J.-F. Landry, sp. nov. from the Galápagos Islands. DNA barcodes support the separation of Hypenopsis from Schrankia in congruence with morphology. Two other species from Panama described in Hypenopsis (H. flualis Schaus, 1916 and H. musalis Schaus, 1916), and later included in Schrankia are illustrated for the first time: the genitalia of their holotypes show that they do not share diagnostic characters of either Schrankia or Hypenopsis, and their COI barcode sequences are distinct from either genus as well as other Hypenodinae genera; they are here regarded as Hypenodinae incertae sedis.


Morphological analysis
Morphological observations were made either on a Nikon SMZ18 or Nikon SMZ25 stereomicroscope.Wing venation was examined from a few slide preparations of cleared wings following the method presented in Landry (1991); forewings were checked on several additional dry specimens by wetting the wings with 95% ethanol on a camel hairbrush; however, the delicate hindwings tended to fold up when wetted so were more difficult to assess.Genitalia dissections and slides were prepared following Landry (2007).Slidemounted genitalia were photographed on a Nikon Eclipse 800 microscope equipped with a Nikon DS Fi1 camera at 40-400× magnifications; NIS Elements software was used for generating deepfocused images from multiple stacked photos.Specimens were photographed with a Canon EOS 60D camera and MPE 65mm lens attached to a Stackshot™ system for image capture; Zerene Stacker™ software was used to generate focused images from multiple stacked photos.The collecting methods used by BL and JFL were presented in Landry (2006) and the methods developed for collecting and preparing small moths can be found in Landry & Landry (1994).
Descriptions follow the terminology of Kristensen (2003) and Holloway (2008).Terms for male valva structures specific to Erebidae follow Goater et al. (2003).Primary type label data are cited verbatim.Citations of the label data of the paratypes are standardized as mentioned in Landry (2006).

DNA sequence data and analysis
DNA was extracted from a single leg of 20 dry museum specimens from the Galápagos Islands from all of the islands on which Hypenopsis specimens were found, i.e., Floreana, Isabela, Pinta, San Cristóbal, Santa Cruz, and Santa Fe, and sequenced for the barcode region of the mitochondrial cytochrome oxydase I gene.Extraction and sequencing were performed at the Canadian Centre for DNA Barcoding (CCDB) at the University of Guelph following standard protocols (deWaard et al., 2008).The same procedure was applied to some North American in 1989 by BL to inventory the Lepidoptera of the Galápagos Islands and to produce identification tools for them.Although the Noctuoidea of the archipelago had already been reviewed in detail by Hayes (1975), a species of Hypenodinae encountered by BL in 1989 and in subsequent collecting expeditions in 1992 and 2004 had not been treated in Hayes' review.
In order to place the Galápagos species to genus cor rectly and confirm that it was undescribed, we examined the known Hypenodinae of the Western Hemisphere.Externally (in shape, size, and forewing pattern) it resembled North American material identified as Schrankia macula, but differed in certain details.Genitalia examination showed that they shared many similarities that could be construed as congeneric.
The widespread distribution of S. macula in North America with an abundance of records (cf.North American Moth Photographers Group, 2023) and the detection of three distinct barcode clusters subsumed under that name, including one that is geographically allopatric with the other two, suggested the possibility of an unresolved species complex.Moreover, the fact that the type series of S. macula was from Panama, coupled with the fact that their genitalia had never been examined, highlighted the need to dissect and sequence them to clarify its identity and determine how closely related it is to the Galápagos taxon.This led us to the actions presented below, i.e., to resurrect the genus Hypenopsis Dyar, 1913 (type species Hypenodes macula Druce, 1891) from synonymy with Schrankia, to redescribe Hypenopsis and its type species based on the first morphological study of the genitalia of its two syntypes, and to describe, as new within Hypenopsis, two North American species formerly confused with H. macula, and a new species from the Galápagos Islands.Two other Western Hemisphere species originally described in Hypenopsis but subse quently assigned to Schrankia, namely H. flualis Schaus, specimens identified as Schrankia macula.Specimen data and sequences are available in a Barcoding of Life Datasystems (BOLD) (Ratnasingham & Hebert, 2013) dataset DSHYHYSC19 'Hypenopsis Hypenodes Schrankia'.Additional records from BOLD used for exploratory and distance analyses, either publicly available or used with their owner's permission, are also included in the dataset.The two syntypes of H. macula and the holotypes of 'Schrankia' flualis and 'Schrankia' musalis were processed differently in Ottawa and Geneva, respectively, as follows: each abdomen was removed, placed in lysis buffer and incubated in proteinase K solution (following the recommended Qiagen DNeasy® kit instructions); the lysate was carefully flushed out of the abdomen and used for DNA amplification and sequencing using the nextgeneration sequencing method described in

RESULTS
Morphological examination revealed that the Galápagos species has male and female genitalia that share several characters with North American specimens traditionally regarded as 'Schrankia macula': porrect labial palpi with an elongate second palpomere; presence of a free and anteriorly directed apodeme with a Yshaped apex on male abdominal tergite VII; vestigial uncus; wholly membranous tuba analis (without scaphium or subscaphium); very thin and narrow teguminal walls; broad shieldlike juxta with a midline; long and stout ampulla with a forked apex; slender phallus; strongly sclerotized and modified female abdominal segment VII partially encapsulating segment VIII/ostium bursae.This combination of characters contrasts markedly with characters of the type species of Schrankia, S. taenialis, and of other Hypenodinae genera, which have a large, hooklike uncus almost as long as the tegumen and bent down over it, and an unmodified male abdominal segment VII.This indicated that the Galápagos species and specimens from North America historically sub sumed under the name S. macula are more closely related to each other than to Old World Schrankia and other Hypenodinae.They are also distinct from each other in DNA barcodes.The type series of H. macula consists of two syntypes, a male and a female, both from Chiriqui, Panama.Their DNA barcodes, although partial, each match one of two BINs, namely BOLD:AAL2056 for the male lectotype and BOLD:AAE5679 for the female paralectotype.Each BIN also comprises specimens from Florida and Costa Rica, with both sexes from both places represented in each BIN.The male and female genitalia from either BIN match those of the syntypes from Panama.Morphological scrutiny did not reveal differences in genitalia associated with each BIN and we consider these two BINs to represent one morphological species, H. macula.This confirms the occurrence of 'true' macula in North America.

Remarks:
The free, Yshaped apodeme of the male abdominal tergite VIII is moveable and can swing on its base with the Yfork oriented anteriorly or posteriorly, although the natural orientation appears to be with the fork pointed anteriorly.This is noticeable when dissecting and the structure can be oriented either way without causing any distortion when preparing a permanent slide mount.For example, in the slide of H. calusa holotype (CNC 17606) the fork on the male sternite VIII and the phallus is much shorter and stouter than in Hypenopsis.
Redescription: Head with short, appressed scales; frons gently rounded; ocellus absent; maxillary palpus inconspicuous; labial palpus (Figs 1720) with porrect, second palpomere about twice as long as head, third palpomere shorter than second and projecting upward; haustellum well developed; antennal flagellomeres in males with cilia slightly longer than flagellomeres bearing them, in females much shorter.Forewing (Fig. 21) with Sc ending on costa at about 2/3, veins R24 on one stem arising before angle of cell, R5 from upper angle of cell and ending on outer margin below apex, M1 also from upper angle of cell, M2 and M3 from lower angle of cell, CuA1 from slightly before lower angle of cell, CuA2 from cell postmedially  H. calusa) and JFL 1770 (lectotype of H. macula) it is pointed towards the anterior end of the abdomen, which seems to be its natural position prior to dissection.The position of this apodeme, its Yshaped apex, and mobility suggest that it could function as a proxy for the reduced uncus in guiding the phallus.It would be interesting to ascertain whether there are muscles attached to the apodeme.
The dense setae of the ventral flange of the clasper and ventral lobe of the cucullus are easily removed during dissection so extra care must be exercised in order to preserve them.They also easily break off during spreading of the valvae and after dehydration prior to slide mounting.
Regarding the phragma, it is difficult to keep them intact when removing the abdomen from a dry specimen, even when doing so with great care.Most of the time, the abdomen will fracture irregularly somewhere in the first or second segment leaving the phragma attached to the metathorax; or part of one or both phragma will remain with the abdomen but in a damaged condition.To obtain intact preparations (e.g., Fig. 22), we severed the metathorax+abdomen together after removing the wings and legs.Separation was carefully done after maceration and cleaning.The juxta is a pair of large, mesially hinged plates, or a single plate with an unsclerotized medial flexible line; in its unspread position it forms a kind of gutter under the phallus.Species of Palearctic Schrankia have a proportionally smaller, Xshaped juxta with thin lower arms (Fibiger et al., 2010, figs 15-18).Species of the genus Luceria share some characters of the male genitalia with Hypenopsis, notably the free, Yshaped apodeme of abdominal tergite VIII and a vestigial uncus (Holloway, 2008).It must be noted, however, that the type species of Luceria, L. novatusalis Walker, described from Sri Lanka, was not represented in our analysis.The type specimen of L. novatusalis in the UKNHM has not been dissected (Alberto Zilli, pers.comm.2022), its genitalia remain unstudied, and the identification of the species is based solely on external aspect.Thus, it remains unknown whether the type species of Luceria shares genital characters with other species currently included in the genus, such as L. oculalis, which was part of our analysis.Notwithstanding the external features presented above in the diagnosis, the only sure way to recognize members of Hypenopsis is to dissect and examine the genitalia.The likelihood of significant undescribed diversity in the Neotropics, as suggested by the many unidentified barcode BINs in BOLD, coupled with our unexpected discovery that most North American material was previously misidentified, makes this a necessity.Differentiation of the four species treated below, which show significant variation in coloration and external aspect, requires likewise.

Diversity and distribution:
The four species treated below are all New World and distributed from the southern half of North America to Central America down to Panama, with the exception of H. galapagensis which is restricted to the Galápagos as far as known.DNA

Diagnosis:
In the male, the lack of median flange and the small ventral flange of clasper, darkly melanized and smooth ampulla, large subtrapezoid juxta plates, base of valva with microtrichia, curved phallus with serrate apex, and slender, spinelike cornutus easily distinguish H. macula from the other three species.These latter three all possess a large, recurved median flange and club-shaped ventral flange of clasper, an ampulla with rugulose or gibbose surface, narrower and proportionally smaller juxta plates, no microtrichia on valva base, a straight or slightly sinuate phallus with a smooth and pointed apex, and a short, tooth like cornutus.In the female, the sclerotized abdominal sternite VI, strongly asymmetrical abdominal sternite VII fused with segment VIII and with asymmetrically lobate posterior margin, asymmetrical and microtrichia covered antrum, presence of a signum with diffuse edges and fish-scale-like microsculpture, are uniquely distinctive.
Redescription: Head pale beige to light brown, darker greyish brown between antennae.Antenna with scales greyish brown; ventral sensilla about 1.5× antennomere length in male, shorter than antennomere width in female.Labial palpus (Fig. 20

Diagnosis:
In the male, the ampulla has a forked apex and a gibbose dorsal surface, the medial flange of the clasper is strongly denticulate, the apex of the phallus is acutely pointed and smooth, and the cornutus is a short, triangular tooth.It is similar to H. sonora in most features, but in the latter the median flange of clasper is smooth, the apex of phallus is less pointed and with microtrichia, and the cornutus is a spurlike tooth.In the female, abdominal sternite VII is trapezoid, about 1.6 wider than long, with a lateral sinuation and a straight and wide posterior margin; the antrum is bulbous, deep, and as wide as the distal margin of abdominal tergite VIII, with a bowllike ostium indentation, and the ductus bursae is membranous over its entire length.
Both galapagensis and sonora differ markedly in the shape and proportions of these structures, as detailed in each species diagnosis.

Etymology:
The species epithet refers to the collecting locality of all known specimens, the Galápagos Islands.H. galapagensis is more greyish brown or blackish brown compared to the warmer brown of H. macula and H. calusa.

Diagnosis: In forewing (Figs 1116)
In forewing pattern H. galapagensis has a paler postmedian fascia that is often broken and that never reaches the costa whereas in the other three species the postmedian fascia, when distinct, is usually complete and extending to the apex of the wing.In male genitalia (Fig. 36), the ampulla with a simple, pointed apex and smooth or slightly rugulose surface is distinctive; the median flange of clasper has a smooth surface (similar to H. sonora, denticulate in H. calusa), the phallus is straight (slightly sinuate in both calusa and sonora) with a slender toothlike cornutus which is intermediate in length between that of H. calusa and H. sonora.In female genitalia (Fig. 48), abdominal sternite VII is narrowly conical, about 4x wider than long, smaller and more weakly sclerotized than in H. calusa and H. sonora; the antrum is elongate, funnelshaped with a Ushaped posterior margin, and about 1/3 the width of sternite VIII whereas it is as wide as sternite VIII in H. calusa; the ductus bursae is slightly sclerotized over its distal half, the inception of the ductus seminalis is near the base of the corpus bursae whereas it is in the distal section of the ductus bursae in H. calusa.
Abdominal tergite VII a broad plate with parallel margins dorsally, about 1/3 longer than preceding tergite.Anterior apophyses about half as long as posterior, but stronger.Ostium bursae opening in middle of broad plate subbasally.Antrum funnel shaped, slightly more than twice as wide as ductus bursae at widest cross section, with ventral margin broadly emarginated.Ductus bursae lightly sclerotized, about 2.5× as long as antrum funnel.
Ductus seminalis from corpus bursae next to connection with ductus bursae.Corpus bursae elongate, about 2.5× as wide as antrum and about as long as antrum+ductus bursae, with tiny signum near base.Posterior apophyses about twice as long as papillae anales.Papillae anales short, quadrangular.
Variation: Based on specimens from all islands.In forewing vestiture some specimens are more uniformly paler greyish brown, with poorly contrasting markings (Figs 1112).Some males barely reach 9.0 mm in wingspan while some females reach a wingspan of 17 mm.In the male genitalia the main variation (Figs 36cf) occurs in the length and curvature of the ampulla of the valva, that of the type specimens, from San Cristóbal being the shortest and least strongly curved.
In addition, the median flange of the clasper also varies in curvature.In females the length and shape of sternite VII varies slightly and the position of the ductus seminalis sometimes appears to come from the ductus bursae before it merges into the corpus bursae.
Biology: Unknown except that moths are attracted to light but also fly in the daytime.Also, specimens have been found mostly at higher elevations (up to 1240 m), but two were collected a few meters above sea level on Isabela and Santa Cruz.

Remarks:
The COI barcode sequences obtained from 16 specimens revealed two clusters of Galápagos Hypenopsis separated by a distance of 2.58%, a gap sufficient for BOLD to assign them to different BINs, BOLD:ADX5739 and BOLD:ADX5740.The latter is represented by specimens from Isabela Island whereas the former is represented by specimens from Floreana, Pinta, San Cristóbal, Santa Cruz, and Sante Fé.This divergence in COI sequences is not reflected in morphological differences observed in genitalia, with the San Cristóbal specimens being more distinct from all others by virtue of their shorter and less curved ampulla of the male valva (Fig. 36d).This is the reason why the type series of H. galapagensis is restricted to specimens from one island and one BIN, BOLD:ADX5739

Etymology:
The species epithet refers to the Sonoran biogeographic region where the type locality is situated.

Diagnosis:
In male (Fig. 37), the ampulla has a forked apex and gibbose surface, the medial flange of clasper is smooth, the apex of phallus is moderately pointed and subterminally bears a set of microtrichia, and the cornutus is sharply pointed, spurlike.It resembles H. calusa in most features except that in the latter, the medial flange of clasper is denticulate, the apex of phallus is acute and lacks microtrichia, and the single cornutus is dentiform and short.In female (Fig. 49), abdominal sternite VII is broadly conical with a narrow and slightly indented posterior margin, the antrum is asymmetrically funnelshaped, half the width of the posterior margin of abdominal tergite VIII with a narrow and irregularly Vshaped ostium, and the ductus bursae is sclerotized over its entire length up to the ductus seminalis with a sharp bend between the latter and the corpus bursae.Females of both H. calusa and H. galapagensis differ significantly in the shape and proportions of these structures, differences given under each species diagnosis (vide supra).vestiture as shown (Fig. 7).Forewing similar to calusa but with paler overall appearance and with more contrast, oblique subterminal fascia uniformly pale in at least one specimen (Laguna Atascosa, Texas).In the single female, pale areas, especially subterminal fascia, predominantly light brown with little contrast.Hindwing pale grey suffused with pale brown to pale brownish grey, darker near brown terminal line.Biology: Unknown.Adults were collected at blacklight in different months of the year suggesting that it could be found throughout the year like H. calusa.
Distribution: Known from southern California, southern Arizona, southern Texas and Oaxaca, Mexico.Additional unrecognized records likely exist in collections.

Remarks:
The few specimens studied were variously rubbed or damaged.Coloration did not appear substantially different from that of H. calusa or H. macula as far as could be assessed.The slightly larger size was marginally different and possibly not significant considering that few specimens were seen, and would not be easily observed without comparing examplars of both species side by side.The bend in the anterior section of the ductus bursae of the female appears to be real, not an artifact of the single slide preparation available.Barcode sequences obtained formed one BIN, BOLD:AAH4894.COI sequences (MHNG) also matches this species closely.In view of the incomplete knowledge of New World Hypenodinae, we treat H. flualis as Hypenodinae incertae sedis.) obtained from the old male holotype did not match anything in BOLD, but both NJ and ML analyses place the species in the Hypenodinae, separated by long branches with weakly supported nodes.Morphologically the welldeveloped, elongate hooklike uncus is reminiscent of the state seen in many Hypenodinae such as Hypenodes and Schrankia, and more generally in Erebidae.The labial palpi (Fig. 59) are upcurved with the third palpomere longer than the second one, unlike in Hypenopsis.In contrast to Schrankia the valva is slender, devoid of ornamentation or accessory lobes like ampulla and clasper, and the sacculus is reduced and indistinct.The phallus is stocky and massive relative to the genitalia (both shown at the same scale in Fig. 60).In view of the incomplete knowledge of New World Hypenodinae, we also treat H. musalis as Hypenodinae incertae sedis.

Fig. 1 .
Fig. 1.NJ tree of Hypenopsis species and representatives of other Hypenodinae genera based on the barcode region of COI, generated by MEGA X using the Kimura2Parameter distance model of nucleotide substitution.Scale bar = divergence of 2%.Specimen identifiers to the right of branches.Asterisks denote generic type species.Scolecocampa liburna (Scolecocampinae) represented an outgroup.

Hypenopsis Dyar, 1913, status revalidated
This is the species with which we initially compared the Galápagos material.
However, contrary to expectations, the bulk of the North American specimens were found to represent a different species, H. calusa sp.nov., distinct in both genitalia and barcode (BOLD:AAB2278), and more widely distributed in southeastern USA than true H. macula but sympatric with it in Florida.77%.Intraspecific haplotype divergence in both H. macula and H. galapagensis n. sp. was sufficient for each to be split into two BINs.IntraBIN distances in H. galapagensis ranged from 0.000.99%(mean 0.44%) (BOLD:ADX5739) and 0.000.46%(mean 0.27%) (BOLD:ADX5740), and the two BINs were 2.58% distant.In H. macula intraBIN distances ranged from 0.000.00%(BOLD:AAL2056) and 0.002.24%(mean 0.47%) (BOLD:AAE5679), whereas the two BINs were 3.19% distant.Based on available sampling, the nearest neighbour to the Hypenopsis clusters was Luceria oculalis, a species widespread in Asia, Australia, and Africa, separated from H. macula by a minimum pdistance of 7.23%.The other genera were all markedly more distant from the Hypenopsis cluster with mean distances ranging from 8.30% (S. taenialis) to 17.05% (M.karsholti).Both 'Schrankia' flualis and 'S.' musalis were also markedly distant from each other at 13.31% as well as from both species of Palearctic Schrankia (>13%) and from the
short and wide.Tergite VI anteriorly margined.Tergite VII indistinguishably fused with tergite VIII (or reduced).Sternite VII also tightly fused to, and shorter than sternite VIII, posterior margin forming asymmetrical lip skewed to right side of segment, with two tongue-like flanges around ostium bursae.Variation: Differences between male and female in external aspect and coloration are slight overall, except in forewing coloration.In some females the forewing maculation is subdued, being almost unicolorous grey brown with faint markings.Most specimens examined were unspread or had damaged wings, consequently wing measures were based on the few that were in adequate condition.Biology: Powell & Opler (2009) mentioned that 'Schrankia macula' larvae feed on bracket fungi (Polyporaceae).Distribution: Brazil, one from Castro, Parana, Brazil, all from Mr. Schaus' collection, and four from La Puerta Valley, near San Diego, California, from Mr. G. H. Field.".Thus, Dyar first referred to the original type specimens (i.e.syntypes) which were also indicated by Druce (1891) in the original description of H. macula.the BINs and the significance of this barcode split is unclear.We consider both barcode clusters to represent the same species.The closest species is H. calusa BOLD:AAB2278 at 4.96% distance.Hypenopsis calusa J.-F.Landry & B. Landry, sp.nov.Figs 310.Specimens of Hypenopsis.(3) H. calusa male holotype, specimen CNCLEP00025834 from Archbold Bio.Stn, Florida.(4) H. calusa male paratype, specimen USNMENT01769369 from Hidalgo Co., Texas.(5) H. calusa female paratype, specimen UFFLMNHMGCL1112705 from Gainesville, Florida.(6) H. calusa female paratype, specimen USNMENT01769427 from Conroe, Texas.(7) H. sonora male holotype, specimen CNCLEP00281099 from Tierrasanta, San Diego Co., California.(8) H. macula male lectotype from Panama.(9) H. macula male, specimen UFFLMNHMGCL1112748 from MiamiDade Co., Florida.(10) H. macula female, specimen UFFLMNHMGCL1112754 from MiamiDade Co., Florida.
Ferguson et al. (1991)and Massachusetts, as well as from Cuba and Puerto Rico, and disjunct records from southern California.It is probable that most, if not all, of the records from the eastern part of the American mainland refer to H. calusa, although records from Florida may also comprise H. macula.The records from Cuba and Puerto Rico remain unverified and could easily represent something else that is superficially similar to H. calusa and may not even be Hypenopsis.Records from southern California are likely the superficially similar H. sonora described below, which occurs in the American Southwest and Mexico.We have not seen any H. calusa specimens from western United States.Ferguson et al. (1991)report of H. macula from Connecticut is likely H. calusa.The same authors also recorded it from Bermuda, adding that the island specimens "have a tendency to be a paler brown color than mainland ones"; the specific identity of Bermuda specimens also remains unverified by genitalia examination.
The habitus of H. calusa is very similar to that of H. macula and H. sonora and genitalia must be examined to distinguish them with certainty.region of H. macula contrasts with the symmetrical and exposed antrum and ostium of H. calusa.Undissected females of H. sonora were not available for comparison.Description: Head (Figs 1718) pale beige mixed with brown, darker greyish brown between antennae.Antenna with scales greyish brown; ventral sensilla or blunt, ventral one sharply pointed.Editum a small lobe with few setae.Mesial flange of clasper markedly developed and sclerotized, extending to about half length of ampulla, medially bent at 90˚ towards ventral margin of cucullus, dorsal surface densely spinulate, apex spatulate.Ventral flange of clasper large, club-shaped, Louisiana, Texas, South Carolina, North Carolina, Virginia, Maryland and New York (Long Island).Under "Schrankia macula" the North American Moth Photographers Group website (2023) shows records from numerous places in the American Southeast from Florida to Texas and Oklahoma, north to southern Illinois, Remarks: DNA barcodes in BIN BOLD:AAB2278.The nearest neighbour is H. macula BOLD:AAL2056 at an average pairwise distance of 4.96%.Hypenopsis galapagensis B. Landry & J.-F.Landry, sp.nov.
. In COI sequence data H. galapagensis is closest to H. sonora, separated by an average distance of 4.01% from BOLD:ADX5739.The collecting locality of Puerto Baquarizo is properly spelled Puerto Baquerizo.
Abdomen g29) suffused with pale brown.Male pre-genital abdomen(Figs 28,29).Sternite VIII lateral projections of anterior margin narrowly triangular, with slender, pointed apices, interspace broadly and shallowly Ushaped.Tergite VIII Yshaped projection with widened shaft about 2× length of tergite VIII and proportionally short apical 'Y'branches.Male genitalia (n=4)(Figs 37, 38).Tegumen elongate arched, broader and with thicker penicular arms than in calusa.Vinculum narrowly Vshaped with small mesial protrusion and rounded saccus.Uncus surface weakly and unevenly sclerotized, outline resembling a doubleheaded battle axe, ~ 0.250.3×length of tegumen (measured from base of valva to apex).Juxta consisting two elongatesubcrescentic plates with a narrow mesial junction/gap, dorsal edge oblique, ventral edge recurved, similar in shape to that of calusa but proportionally shorter.Valva, proportionally short with apex of cucullus extending to, but not beyond, apex of uncus, wider in distal half.Costa thickly sclerotized and wider than in calusa.Ampulla strongly sclerotized, downcurved, with series of small protuberances along upper side of shaft (variable in number and distribution but fewer than in calusa, apically bifurcate with dorsal fork variously rounded, blunt or explanate, ventral one a rounded point.Editum a small lobe with a few setae similar to calusa.Mesial flange of clasper well developed and sclerotized, extending to about half length of ampulla, smoothly curved towards ventral margin of cucullus, with smooth dorsal surface, apically weakly spatulate (may appear ~0.8× length of vinculum+valva, apically oblique and tapered with ventral surface finely spinulate, caecum dilated, wider than shaft; vesica with single, spinelike cornutus which appears apically rounded or acuminate depending on orientation (cf.Figs 37b vs 38b); bulbus ejaculatorius about 0.5× length of phallus, wholly membranous, with crescentshaped dilation.Female pre-genital abdomen and genitalia (n=1) (Fig.49).Abdominal segment VII sclerotized, about 2× as long as segment VI. Tergite VII broadly transverse.Sternite VII conical, extended as a lip over and covering basal half of sternite VIII.Sternite VIII ~0.5× length of tergite VII.Anterior apophyses ~0.5× as long as posterior ones.Antrum elongate, funnelshaped, slightly asymmetrical, ~0.5× as long as sternite VIII, with narrowly Ushaped posterior emargination, and surface of sinus vaginalis smooth.Ostium bursae opening mesially.Ductus bursae sclerotized to inception of ductus seminalis, ~3× as long as SVIII, with sharp bend in anterior section above ductus seminalis.Corpus bursae elongateovoid, its surface with isodiametric microsculpture, signum absent; inception of ductus seminalis close to corpus bursae just posterior to bend of ductus bursae.Posterior apophyses about 1.5× as long as papillae anales.Papillae anales melanized, together with ovipositor slightly longer than tergite VIII.