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.
INTRODUCTION
The Hypenodinae, including the Micronoctuini Fibiger, in Fibiger & Lafontaine (2005) (Zahiri et al., 2012) comprise more than 500 species distributed worldwide, with the greatest species richness (more than 400 species) in the Old World and most belonging to the Micronoctuini (Fibiger, 2007, 2008, 2010, 2011; Holloway, 2008). The subfamily has been diagnosed on the basis of the loss of ocelli (Fibiger & Lafontaine, 2005) and by two larval synapomorphies articulated by Beck (1999, 2000). However, some included genera possess ocelli, and larvae are known only for two genera (Hypenodes Doubleday, 1850 and Schrankia Hübner, 1825) (Fibiger & Lafontaine, 2005: 25). The placement and status of the subfamily has changed several times in recent decades but is now well supported within the Erebidae (Fibiger & Lafontaine, 2005; Zahiri et al., 2012). Although no New World species of Hypenodinae were included among the taxa sampled by Zahiri et al. (2012), Lafontaine & Schmidt (2010) and Schmidt et al. (2018) built on this framework to include New World genera historically associated with Hypenodinae (e.g., Franclemont & Todd, 1983).
The described Western Hemisphere fauna of Hypenodinae comprises 18 species in five genera: Dasyblemma Dyar, 1923 (1 sp.), Dyspyralis Warren, 1891 (4 spp.), Hypenodes (8 spp.), Parahypenodes Barnes & McDunnough, 1918 (1 sp.), Quandara Nye, 1975 (1 sp.), and Schrankia (3 spp.) (Poole, 1989; Lafontaine & Schmidt, 2010; Savela, 2023). Of these, the Neotropical fauna accounts for three species described in Hypenodes, one in Quandara, and three in Schrankia.
This paper stemmed from a research program initiated 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 correctly 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 subsequently assigned to Schrankia, namely H. flualis Schaus, 1916 and H. musalis Schaus, 1916 (see for example Poole, 1989), both also described from Panama, are illustrated and shown to belong to Hypenodinae, but remain unassigned to genus.
MATERIAL AND METHODS
The material studied is deposited in the following institutions:
Canadian National Collection of Insects, Arachnids, and Nematodes, Ottawa, Ontario, Canada (CNC)
Charles Darwin Research Station, Santa Cruz Island, Galápagos, Ecuador (CDRS)
Cornell University Insect Collection, Ithaca, New York, U.S.A. (CUIC)
McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, Gainesville, Florida, U.S.A. (MGCL)
Muséum d'histoire naturelle de Genève, Geneva, Switzerland (MHNG)
Museum für Naturkunde Berlin, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Berlin, Germany (ZMB)
National Museum of Natural History, Smithsonian Institution, Washington, D.C., U.S.A. (USNM)
Natural History Museum, London, U.K. (NHMUK)
Other abbreviations used are BL for Bernard Landry, JFL for Jean-François Landry, SVII and SVIII for 7th and 8th sternites (S7 and S8 in figures), and TVII and TVIII for 7th and 8th tergites (T7 and T8 in figures).
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 hair-brush; 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).
Slide-mounted genitalia were photographed on a Nikon Eclipse 800 microscope equipped with a Nikon DSFi1 camera at 40-400× magnifications; NIS Elements software was used for generating deep-focused images from multiple stacked photos. Specimens were photographed with a Canon EOS 60D camera and MP-E 65-mm 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 specimens identified as Schrankia macula. Specimen data and sequences are available in a Barcoding of Life Datasystems (BOLD) (Ratnasingham & Hebert, 2013) dataset DS-HYHYSC19 ‘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 next-generation sequencing method described in Prosser et al. (2016) for century-old type specimens. Sequencing was performed at CCDB. After lysis, the remaining abdominal cuticle and genitalia were removed from the lysis liquid, briefly macerated in KOH, dissected and slide mounted following standard methods (e.g., Landry, 2007). Remaining lysate fractions were stored at -80 °C in the CNC pending final deposition in ZMB and USNM. All species formerly in Hypenopsis (macula, flualis, musalis) were selected for analysis as well as representatives of eight genera of Hypenodinae available in BOLD, including type species wherever possible (denoted by an asterisk): Dasyblemma straminea Dyar, 1923*, Dyspyralis illocata Warren, 1891*, Hypenodes humidalis Doubleday, 1850* (European), H. fractilinea (Smith, 1908) (North American), Luceria oculalis (Moore, 1877), Micronoctua karsholti Fibiger, 1997*, Parahypenodes quadralis Barnes & McDunnough, 1918*, Quandara hypozonata (Hampson, 1910)*, Schrankia taenialis (Hübner, 1809)* and S. costaestrigalis (Stephens, 1834) (both European). Scolecocampa liburna (Geyer, 1837) (Erebidae: Scolecocampinae) was selected as outgroup for the ML analysis, although it was also included in the distance analysis for comparability. The Scolecocampinae are the sister-group to the Hypenodinae (Zahiri et al., 2012) and S. liburna is the type species of Scolecocampa.
Barcode sequences were initially explored using the Taxon-ID tree tool as implemented in BOLD in order to visualize their grouping and quantify their BIN representation. Sequences were aligned with Kalign (Lassmann & Sonnhammer, 2005) implemented in BOLD, and downloaded as a fasta file. A neighbor-joining (NJ) tree was generated with MEGA X under the Kimura-2-Parameter distance model of nucleotide substitution. Distance analysis was performed with MEGA X (Kumar et al., 2018). Maximum likelihood (ML) analysis was carried out on the IQ-TREE 1.6 (Nguyen et al., 2015) on unpartitioned data. The best substitution model was GTR+F+G4 as determined by ModelFinder (Kalyaanamoorthy et al., 2017) using the Bayesian Information Criterion (BIC), as implemented in IQ-TREE. Clade support was assessed with 1000 iterations of ultrafast bootstrap (UFBoot2) and single-branch SH-aLRT and approximate Bayes tests; 50 independent tree searches using the --runs command were performed from which the best tree was selected. We employed posterior probability values (PP) ≥ 90, UFB ≥ 95 or SH-aLRT ≥ 80 (Nguyen et al., 2015) to indicate support. The resultant tree file was edited using FigTree v.1.4.4 (Rambaut, 2018) and Adobe Illustrator.
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 Y-shaped apex on male abdominal tergite VII; vestigial uncus; wholly membranous tuba analis (without scaphium or subscaphium); very thin and narrow teguminal walls; broad shield-like 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, hook-like 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 subsumed 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.
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. This is the species with which we initially compared the Galápagos material. Moreover, southwestern specimens of ‘macula’ from California, Texas, and Mexico were found to represent a third species, H. sonora sp. nov., also with different genitalia and barcode (BOLD:AAH4894). Both species are superficially similar to H. macula and are described as new below.
Distance analysis visualized in a neighbor-joining tree (Fig. 1) is congruent with the recognition of four species of Hypenopsis that are well differentiated morphologically, three of which are new, H. macula, H. calusa sp. nov., H. galapagensis sp. nov., and H. sonora sp. nov. Average interspecific distances ranged from 4.11% (between H. sonora and H. galapagensis) to 6.47% (between H. calusa and H. sonora), and intraspecific distances from 0.00-1.77%. Intraspecific haplotype divergence in both H. macula and H. galapagensis n. sp. was sufficient for each to be split into two BINs. Intra-BIN distances in H. galapagensis ranged from 0.00-0.99% (mean 0.44%) (BOLD:ADX5739) and 0.00-0.46% (mean 0.27%) (BOLD:ADX5740), and the two BINs were 2.58% distant. In H. macula intra-BIN distances ranged from 0.00-0.00% (BOLD:AAL2056) and 0.00-2.24% (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 p-distance 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 Hypenopsis group (11.89-15.76%), congruent with their distinct genitalia morphology.
Maximum-likelihood analysis showed three Hypenopsis clusters with high branch support, and a polytomy for H. macula (Fig. 2). Support values were also high for the clade uniting Luceria and the Hypenopsis group whereas branch support was generally low among other Hypenodinae. These results are congruent with observed morphological characters in showing that 1) these four species are not closely related to Schrankia, and 2) they are more closely related to Luceria, albeit distinct.
BIN separation in H. macula was not supported, and only moderately so for H. galapagensis. Close morphological scrutiny did not reveal any differences in genitalia of either sex that corresponded to the BIN splits in either species; nor did the haplotype divergence in H. galapagensis correspond to the inter-island distribution of the samples, as might have been expected for a species distributed over an archipelago. BIN splits uncorroborated by morphological or other differentiating evidence are not uncommon in other Noctuoidea, for example they are recorded in about 9% (140/1541) of species of Canadian Noctuoidea (Zahiri et al., 2014).
Regarding ‘Schrankia’ flualis and ‘S.’ musalis, originally described in Hypenopsis, both morphology and DNA barcodes clearly indicate that they do not belong there, nor in Schrankia or Hypenodes. We further discuss and illustrate them at the end of this work.
Hypenopsis Dyar, 1913, status revalidated
Hypenopsis Dyar, 1913: 296. Type species: Hypenodes macula Druce, 1891, by original designation.
Diagnosis: In habitus, Hypenopsis is not unlike species assigned to the Holarctic Hypenodes (type species Hypenodes humidalis Doubleday, 1850) or Palearctic Schrankia (type species Pyralis taenialis Hübner, 1809). They are small (9-18 mm in wingspan), narrow-winged, grey-brown moths with more or less contrasting postmedial transverse lines or fasciae on the forewing, sometimes with a discal spot, and without markings on the hindwing. The labial palpus of Hypenopsis (Figs 17-20) differs from that of Hypenodes in its long, straight second palpomere and short third palpomere directed upward at a nearly right angle, versus a broadly upturned second palpomere (and thus the whole palpus) in Hypenodes, and a straight and porrect or slightly erect second palpomere with a much shorter third palpomere that is also porrect or slightly directed upward in Schrankia. In the terminal segments of the male abdomen Hypenopsis differs from Schrankia and Hypenodes in possessing internal paired lateral apodemes projecting anteriorly from the base of sternite VIII and an internal Y-shaped, free apodeme stemming from the base of tergite VIII with the fork directed anteriorly (Figs 23-29, 35). In male genitalia Hypenopsis (Figs 30-34, 36-38) differs from Hypenodes and Schrankia in the markedly reduced uncus, the valva with a long, stout, downcurved ampulla with a forked apex extended to the apex of the cucullus, and one short, bulbous, densely setose lobe mesially on the ventral margin; the valva of Hypenodes is simple and narrow with two short digitiform projections near the base (costal lobes of Ferguson, 1954); the valva of Schrankia has an ampulla that is straight with an oblique apex and is much shorter than the cucullus, and the densely setose lobe from the ventral margin is an elongate projection extending to the dorsal margin or beyond (cf. Fibiger et al., 2010); both Hypenodes and Schrankia have a prominent, slender, downcurved uncus about as long as the tegumen. In Hypenopsis the juxta is a broad plate with a medial crease or fold line; in Schrankia the juxta is a thin, X-shaped sclerite. In habitus, shape of labial palpi, presence of apodemes on male abdominal tergite and sternite VIII, lack of or reduced uncus, and complex valva, Hypenopsis is similar to Luceria Walker, 1859 from the Old World tropics (Holloway, 2008), but Luceria has a more strongly developed set of apodemes 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 17-20) 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 R2-4 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 (quadrifid venation); anal vein strong, ending at anal angle. Hindwing venation with Sc+R ending just below start of terminal curve of costa, Rs ending on termen at apex (longest part of wing), M1 stemmed with RS for short distance after upper angle of cell, M2 feeble, arising from below middle of cell and ending in depression of termen, M3 and CuA1 short stemmed from lower angle of cell (trifine venation), crossvein at right angles between Rs+M1 stem and M3+Cubital stem, CuA2 from beyond middle of cell, 1A+2A to anal angle, 3A about half as long as preceding. Wing coupling mechanism in males with frenulum of one acantha held in place by costal frenulum hook; females with frenulum of 2-3 acanthae held in place by row of thin, curved scales directed dorsally above stem of M+Cu.
Abdomen (both sexes): Phragma lobes (Fig. 22) well developed, suborbicular, situated on pleural area near base of abdominal tergite I. First sternal sclerite elongate-subrectangular with short venulae and stubby anterior apodemes.
Male pre-genital abdomen (Figs 23-29, 35): Tergite VIII weakly sclerotized, half as wide as sternite VIII, anterior margin with a prominent internal mesial Y-shaped apodeme directed anteriorly with forked apex extending to or beyond middle of sternite VII. Sternite VIII weakly sclerotized, anterior margin mesially concave, laterally with pair of triangular apodemes pointed anteriorly, ending in thin point and extending to or beyond posterior margin of sternite VII.
Male genitalia (Figs 30-34, 36-38, 50, 51): Tegumen arms very thin, forming a slender arch with very narrow apical connection, penicular arms very thin, filament like, ventro-anteriorly fused along anterior margin of valva-vinculum. Tuba analis entirely membranous. Uncus reduced or vestigial, consisting of lightly sclerotized, ill-defined, short plate(s). Juxta a relatively large plate with medial unsclerotized, flexible crease. Valva (cf. Fig. 30 for details of parts) slender, apex not or barely exceeding uncus. Costa basally thick, ventral edge at mid-length with triangular protrusion at or near level of distal end of sacculus. Ampulla large, heavily sclerotized, downcurved, horn-like, with apex simply pointed or bifid, dorsal surface rugulose (galapagensis), gibbose (calusa, sonora), or smooth and darkly melanized (macula), and with small editum near base. Clasper (absent in macula) shaped as recurved median process with wider base and slender distal half, thickly sclerotized, apically variously spatulate, with dorsal surface smooth (galapagensis, sonora) or denticulate (calusa); ventral flange of clasper either small (macula), or large and club-shaped, sclerotized, densely setose with setae oriented dorsally. Cucullus almost membranous except for narrowly sclerotized costal and ventral margins, apically rounded; ventral margin bearing densely setose lobe with setae oriented oblique-dorsally. Sacculus thick, elongate, about half length of valva. Phallus slender, straight or curved, with base slightly dilated; vesica with single cornutus; bulbus ejaculatorius crescentic, its inception situated on dorsal side of phallus near base.
Female pre-genital abdomen: Segment VI nearly membranous or thinly sclerotized, contrasting with sclerotized preceding and following segments; sternite and tergite VII more markedly sclerotized and differently shaped than all preceding segments, symmetrical or asymmetrical.
Female genitalia (Figs 39-49, 52, 53): Sternite VIII symmetrical (calusa, galapagensis, sonora) with ostium bursae situated in a mesial notch, or strongly asymmetrical with ostium bursae skewed to right (macula), and about as wide as high (lateral aspect when not flattened). Anterior apophyses shorter than sternite VIII (galapagensis), or reduced to stubs (calusa, macula, sonora). Antrum wide, forming deep chamber almost as wide as sternite VIII. Ductus bursae slightly longer than sternite VIII, lightly sclerotized or membranous. Ductus seminalis from corpus bursae next to connection with ductus bursae (galapagensis, macula, sonora), or from middle of ductus bursae (calusa). Corpus bursae globular to ovoid; signum a small scobinate patch (macula, galapagensis), or absent (calusa, sonora). Ovipositor short, about equal in length to abdominal sternite VIII when fully extended. Posterior apophyses thin, longer than ovipositor or extending to posterior margin of sternite VIII. Papillae anales short and broad, separated from each other, with setation of variable length and extensive coverage of microtrichia.
Remarks: The free, Y-shaped apodeme of the male abdominal tergite VIII is moveable and can swing on its base with the Y-fork 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 was inadvertently flipped during mounting and thus is projected towards the caudal end of the abdomen; whereas in slides CNC 17611, CNC 17613 (both 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 Y-shaped 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, X-shaped 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, Y-shaped 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 barcode data available in BOLD (cf dataset DS-HYHYSC19) indicate that there may be additional putative species from Costa Rica represented by five BINs.
Host and biology: Hypenodinae biology and larval hosts are poorly known. Their larvae in general are said to feed on algae, lichens, and fungi, with pupation taking place in a dense silk cocoon covered with plant or wood fragments (Powell & Opler, 2009). The same authors as well as Covell (1984) mentioned that ‘Schrankia macula’ larvae feed on bracket fungi (Polyporaceae) without indicating the source for this observation. The ‘macula’ mentioned by them likely pertains to one of the North American species treated below; it remains to be determined whether this would apply to all of them. Skinner & Wilson (2009) stated that the larval foods of British species of Hypenodes and Schrankia are unknown in the wild but in captivity S. taenialis (Hübner, 1809) has been reared on lettuce (Lactuca, Asteraceae) and flowers of thyme (Thymus, Lamiaceae) and heath Calluna (Ericaceae) as well as sliced runner beans (Phaseolus, Fabaceae); S. costaestrigalis (Stephens, 1834) on lettuce, flowers of thyme and wild mint (Mentha, Lamiaceae) as well as damp and withered sallow (Salix, Salicaceae) leaves; and Hypenodes humidalis Doubleday, 1850 on Erica (Ericaceae). Fibiger et al. (2010) added that H. humidalis larvae also feed on Comarum palustre (L.) Scop. (Rosaceae) and sphagnum (Sphagnaceae) from September to spring. Leraut (2019) mentioned Heracleum spondylium (Apiaceae) as host for S. taenialis. Hellman & Parenzan (2010) added Ligustrum (Oleaceae), Melampyrum (Orobanchaceae) and Symphoricarpos (Caprifoliaceae) for this species and Clematis (Ranunculaceae), Fuchsia (Onagraceae), Galeopsis (Lamiaceae) and Melampyrum for S. costaestrigalis.
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 Kimura-2-Parameter 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.
Fig. 2.
Maximum-likelihood phylogram from an unpartitioned analysis of the barcode region of COI. Lower phylogram is continuation of Hypenopsis branch represented by triangle in upper phylogram. Specimen IDs to the right of branches. SH-aLRT values / posterior probabilities / Ultrafast bootstrap values to the left of the corresponding node. Node colours: green, PP ≥ 90 and either SH-aLRT ≥ 80 or UFB ≥ 95; yellow, one of PP ≥ 90, SH-aLRT ≥ 80 or UFB ≥ 95; red, PP < 0.90, SH-aLRT < 80 and UFB < 95. Scolecocampa liburna (Scolecocampinae) represented an outgroup.
KEY TO SPECIES OF HYPENOPSIS
1a Male
1b Female 5
2a Valva with ampulla darkly melanized, appearing almost black, clasper without median flange and small, inconspicuous ventral flange; phallus slender and distally curved with serrated apex (Fig. 31) macula
2b Valva with ampulla not melanized, appearing as other sclerotized parts of genitalia, clasper with prominent, downcurved median flange and with large, club-shaped ventral flange; phallus sinuate with smooth apex 3
3a Ampulla with apex simple (Fig. 36) galapagensis
3b Ampulla with apex Y-forked 4
4a Median flange of clasper denticulate on dorsal edge (Fig. 33) calusa
4b Median flange of clasper smooth (Fig. 37) sonora
5a Antrum and ostium asymmetrical, surface of sinus vaginalis covered with microtrichia; posterior margin of abdominal sternite VII with asymmetrical lobes (Fig. 39) macula
5b Antrum and ostium symmetrical, surface of sinus vaginalis smooth, without microtrichia; posterior margin of abdominal sternite VII without lobes, symmetrical, entire sclerite conical or trapezoid 6
6a Antrum globular, as wide as abdominal sternite VIII, with bowl-like posterior margin; ductus bursae membranous, inception of ductus seminalis in distal section of ductus bursae (Fig. 45) calusa
6b Antrum elongate, funnel-shaped, narrower than abdominal sternite VIII; ductus bursae at least partly sclerotized, inception of ductus seminalis near base of corpus bursae 7
7a Antrum narrowly and smoothly funnel-shaped, with U-shaped ostium; ductus bursae lightly sclerotized in distal section, straight (Fig. 48) galapagensis
7b Antrum irregular in outline, with irregular, almost slit-like ostium; ductus bursae entirely sclerotized up to inception of ductus seminalis, with a sharp bend before corpus bursae (Fig. 49) sonora
Hypenopsis macula (Druce, 1891), comb. rev.
Figs 8-10, 20, 26-27, 31-32, 39-41, 50, 52
Hypenodes macula Druce, 1891: 441, pl. 36 fig. 1. – Dyar, 1913: 296. – McDunnough, 1938: 129.
Schrankia macula (Druce): Franclemont & Todd, 1983: 122. – Poole, 1989: 899. – Savela, 2023.
Material examined: 15♂, 8♀, 2 of undetermined sex.
Described from two specimens (Druce 1891), a male and a female.
Lectotype (here designated): ♂ MfN-940363 (labels all white and in black ink), ‘Chiriqui | Ribbe’ [typed]; ‘Coll. | Staudinger’ [typed]; ‘Hypenodes | sp. n. m. 12.2.91’ [hand written]; ‘533.’ [hand written]; ‘Hypenodes | macula | Type. Druce’ [hand written]; ‘Lectotype ♂ | Hypenodes | macula Druce | Select. by J.-F. Landry, 2020’ [orange]. Genitalia slide JFL 1770. Deposited in ZMB.
Paralectotype (here designated): ♀ MfN-940364, with same first two labels as lectotype but without the third and fourth (i.e., no date and number); ‘Paralectotype ♀| Hypenodes | macula Druce | Select. by J.-F. Landry, 2020’. Genitalia slide JFL 1771. Deposited in ZMB.
Other specimens examined: – Costa Rica: 2 ♂ (09-SRNP-107251, 09-SRNP-107297, slide CNC 17629), Alajuela, Area de Conservacion Guanacaste, Sector Rincon Rain Forest, Protrero Chaves, 10.939°N, 85.322°W, 433 m, 19.viii.2009, light trap (F. Quesada & H. Cambronero) (CNC); 2 ♀, Alajuela, Area de Conservacion Guanacaste, Sector Rincon Rain Forest, Rio Negro, 10.904°N, 85.3029°W, 329 m, light trap, 23.i.2009 (09-SRNP-100187, slide CNC 17627), 24.i.2009 (09-SRNP-100900, slide CNC 17628) (H. Cambronero & F. Quesada) (CNC); 1 undetermined sex (11-SRNP-105137), Guanacaste, Area de Conservacion Guanacaste, 10.7314°N, 85.6008°W, 141 m, 25.x.2011, light trap (H. Cambronero & S. Rios) (CNC); 1 undetermined sex (17-SRNP-106225), Guanacaste, Area de Conservacion Guanacaste, 10.8287°N, 85.4030°W, 860 m, 18.x.2017, light trap (H. Cambronero & R. Franco) (CNC). – USA, Florida: 2 ♂, 3 ♀, Brevard Co[unty]., Melbourne, Weston Park, 28.1585°N, 80.6522°W, U[ltra]V[iolet]L[ight], 29.x.2017 (UF-FLMNH-MGCL-1112692, slide JFL 1788), 8.xi.2017 (UF-FLMNH-MGCL-1112693), 21.xii.2017 (UF-FLMNH-MGCL-1112697), 8.xii.2017 (UF-FLMNH-MGCL-1112695), 12.i.2018 (UF-FLMNH-MGCL-1112698) (F.M. Blaine) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112760, dissection in microvial), Broward Co., Dania Beach, 704 N Federal H[igh]w[a]y, 26.0613°N, 80.1365°W, 7.x.2015, UVL E15-5837 (E. Dougherty, J. Garcia) (MGCL); 2 ♂ (UF-FLMNH-MGCL-1112758, slide MGCL 1443; UF-FLMNH-MGCL-1112759, dissection in microvial), Broward Co., Fort Lauderdale, CAPS IMS., 25.0653°N, 80.1936°W, 3.x.2013, UVL E13-7395 (M. Dacosta, J. Garcia) (MGCL); 1 ♂(UF-FLMNH-MGCL-1112761, dissection in microvial), Broward Co., Fort Lauderdale, Hugh Taylor Birch S[tate].P[ark]., 26.1388°N, 80.1078°W, 6.xi.2015, UVL E15-6286 (J. Garcia, E. Dougherty) (MGCL); 1 ♀ (UF-FLMNH-MGCL-1112756, slide MGCL 5768), Miami-Dade Co., Coral Gables, USDA ARS SHRS, 25.6421°N, 80.2948°W, 23.ii.2020, UVL trap (J. Farnum) (MGCL); 1 ♀ (UF-FLMNH-MGCL-1112757, slide JFL 1776), Miami-Dade Co., Coral Gables, USDA ARS SHRS, 25.6421°N, 80.2945°W, 23.ii.2020, UVL trap E20-733 (J.E. Hayden) (CNCL); 1 ♂ (UF-FLMNH-MGCL-1112755, slide MGCL 4978), Miami-Dade Co., Florida City, Home 2 Suites, 25.4504°N, 80.4761°W, 30.xi.2018, dead on breakfast table in lobby, (J.E. Hayden) (MGCL); 6 ♂, 1 ♀, Miami-Dade Co., Miami Cutler USDA ARS SHRS cacao, 25.6424°N, 80.2946°W, UVL trap, 14-15.x.2015 (UF-FLMNH-MGCL-1112748, dissection in microvial; UF-FLMNH-MGCL-1112750; UF-FLMNH-MGCL-1112749, slide JFL 1784; UF-FLMNH-MGCL-1112751, slide JFL 1775), 15-16.x.2015 (UF-FLMNH-MGCL-1112752, slide JFL 1783, UF-FLMNH-MGCL-1112753), 24.x.2015 (UF-FLMNH-MGCL-1112754, slide MGCL 3541 (J.E. Hayden) (CNC, MGCL).
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, spine-like 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) with second palpomere straight, porrect, with scales slightly erect ventrally and dorsally, with outer side dark brown and inner side pale beige; third palpomere slightly < half length of second, dark brown with pale beige base and apex. Thorax dorsally pale beige with sparse greyish brown suffusion. Foreleg and midleg grey-brown. Hindleg pale beige, lighter than other legs. Wingspan (n=4): 13.0-16.0 mm; forewing length (n=6): 6.0-7.5 mm. Wing vestiture as shown (Figs 8-10). Forewing predominantly dark brown with indistinct, blackish-brown chevron-shaped antemedial line; a contrastingly pale beige transversely oblique subterminal fascia extending from apex of costa to near middle of dorsal margin and lined with a thin black postmedial line on inner side, in some specimens with sparse brown over pale fascia; a black discal spot at end of cell and six black terminal dashes. In some females (e.g., Fig. 10) forewing surface almost uniformly grey-brown with faint or concolorous subterminal fascia and faint anteromedial line; more uniform coloration of forewing seen only in females. Hindwing pale grey to pale brownish grey, anterodistal area slightly infuscated, with thin, dark brown terminal line. Abdomen pale beige with sparse brown irroration. Abdominal sternite VIII (Figs 26, 27) lateral projections of anterior margin broadly triangular with short apices, interspace broadly U-shaped. Abdominal tergite VIII with Y-shaped projection thin, about 1.3× length of tergite VIII and with short apical fork.
Male genitalia (n=12) (Figs 31-32). Tegumen weakly sclerotized, elongate V-shaped, slightly widened in distal half, with penicular arms thinly attenuate to base of valva. Vinculum with broadly rounded saccus, with small mesial protrusion. Uncus thinly sclerotized, with indistinct margins, elongate-trapezoid, about 0.2× length of tegumen (measured from base of valva to apex). Juxta consisting of two wide, elongate-trapezoid, divergent plates with a narrow mesial junction, dorsal edge transverse and concave forming a gutter at the manica. Valva slender, proportionally short, with apex of cucullus extending to, but not beyond, apex of uncus and base covered with microtrichiae. Costa narrowly sclerotized contrasting with membranous mesial wall, extending nearly to apex of cucullus. Ampulla strongly sclerotized with darkly melanized distal half, downcurved, apex bifurcate with dorsal fork smaller than ventral one and extending to apex of cucullus. Editum narrow, inconspicuous, with few setae. Ventral flange of clasper small, anvil-shaped, with 20-25 densely set setae. Cucullus narrowly elongate, with long ventral lobe, extending from ventral flange of clasper to near apex of cucullus, with dense cover of long setae oriented antero-dorsally. Sacculus about 0.5× length of valva, elongate-subrectangular with weakly convex ventral margin. Phallus thinly elongate, distally attenuate, 0.8× length of vinculum+valva, distal third dorsally upcurved, apically finely serrate and with caecum slightly narrowed; vesica with single, elongate cornutus about 0.15× length of phallus shaft; bulbus ejaculatorius about 0.5× length of phallus, wholly membranous, with crescent-shaped dilation.
Female pre-genital abdomen and genitalia (n=6) (Figs 39-41). Abdominal segment VI (Fig. 40) shorter than preceding segments, unevenly sclerotized/melanized, tightly associated or fused with segment VII, without distinguishable intersegmental membrane. Sternite VI 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. Tergite VIII broadly conical with sides asymmetrically extended around on ventral side and surrounding narrow sternite VIII. Sternite VIII < 0.5 width of tergite VIII, forming broad, funnel-shaped sinus vaginalis with surface densely spinulose. Anterior apophyses very short, reduce to stubs on anterior end of sternite VIII, left apodeme slightly longer than right one. Antrum funnel-shaped, asymmetrically skewed to left side, ∼half as wide as sinus vaginalis and ∼2× as wide as ductus bursae. Ostium bursae as wide as sinus vaginalis, opening asymmetrically off to left side. Ductus bursae weakly sclerotized and transversely wrinkled in posterior half, ∼3× as long as antrum. Corpus bursae ovoid, surface with very fine fishscale-like microsculpture, with one tiny, dentate signum situated in posterior third of corpus; inception of ductus seminalis at junction with ductus bursae. Posterior apophyses about 1.25-1.3× as long as papillae anales. Papillae anales melanized, together with ovipositor short, ∼0.5–0.6× as long as tergite VII+ tergite VIII.
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: In North America, confirmed records of H. macula are known only from Florida, mostly in the south from Miami-Dade, Brevard, and Broward Counties, with a single record from Gainesville, Alachua Co. All records that we examined and confirmed as true H. macula were collected from 2009 onward. It remains to be verified whether this is indicative that occurrence of the species in Florida and North America is recent or an artefact of our examined material. Other records of the species are from Panama (the type locality) and Costa Rica.
Remarks: Following his description of Hypenopsis, Dyar (1913) designated the type species as Hypenodes macula Druce, 1891, immediately followed by the following statement: “Described from two specimens from Chiriqui, Panama.” In the following sentence, Dyar added “I have before me thirteen [sic] from Jalapa and Orizaba, Mexico, one from Sao Paula, E.E. 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. Whatever specimens Dyar referred to in his second sentence were clearly not part of Druce's original type material of H. macula. In fact, Dyar did not explicitly state that these other thirteen specimens belonged to the same species nor did he describe them. We have not seen the other specimens mentioned by Dyar and do not know what they represent but this does not constitute a case of misidentification of the type species as explained and exemplified in Article 70.3 of the fourth edition of the International Code of Zoological Nomenclature.
DNA barcodes: Two BINs are represented, BOLD:AAE5679 and BOLD:AAL2056 separated by a distance of 3.20%. COI sequences of the two syntypes split into each BIN, the male lectotype in BOLD:AAL2056 and the female paralectotype in BOLD:AAE5679. Each BIN comprises specimens of both sexes from each area sampled in Costa Rica, Panama and Florida. Some specimens from both BINs were even collected together in Costa Rica and Florida. Careful scrutiny of genitalia did not reveal any morphological differences from either sex between 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 3-6, 17-18, 22-25, 33-34, 42-47, 51, 53
Schrankia macula (Druce): McDunnough, 1938: 129. – Forbes, 1954: 383. – Franclemont & Todd, 1983: 122. – Covell, 1984: 321, pl. 41 fig. 8. – Poole, 1996: 772. – Lafontaine & Schmidt, 2010: 28. – North American Moth Photographers Group, 2023.
Material examined: 89 ♂, 55 ♀.
Holotype: ♂, ‘USA, FL, Highlands Co. | Lake Placid | Archbold Bio. | Stn, Red Hill, 54 m | 27.185 N 81.3407 W | 20.VI.2006 @ BL & MVL | JF Landry & PDN Hebert | CNCLEP00025834’ [printed in black on card stock]; ‘Barcode of Life Project | DNA extracted’ [printed in black on blue card stock]; ‘HOLOTYPE | Hypenopsis | calusa | Landry & Landry’ [orange card stock, handwritten]. Abdomen undissected. BOLD Sequence ID MNAB243-07. Deposited in the CNC.
Paratypes: 66 ♂, 42 ♀. – USA, Alabama: 1 ♂ (UF-FLMNH-MGCL-1112725, slide JFL 1785), Baldwin Co[unty]., Weeks Bay Preserve, 30.416°N, 87.819°W, 2.viii.2009, pitcher plant bog, U[ltra]V[iolet] trap (J. Bolling Sullivan) (MGCL). – Florida: 4 ♂, 2 ♀, Alachua Co., Gainesville, 2004 SE 41st Avenue, 29.615°N, 82.298°W, 8.v.2005 (MGCL-FLMNH-57987), 17.v.2005 (MGCL-FLMNH-57997), 10.vi.2005 (MGCL-FLMNH-57990), 8.xi.2005 (MGCL-FLMNH-57988), 5.xii.2005 (MGCL-FLMNH-57978), 28.xii.2005 (MGCL-FLMNH-57982) (G.T. Austin) (MGCL); 2 ♂, Alachua Co., Gainesville, 2832 NW 41st, 29.692°N, 82.364°W, UVL[ight] trap, 29-30.ix.2019 (UF-FLMNH-MGCL-1112703), 6-7.x.2019 (UF-FLMNH-MGCL-1112704, slide FLMNH-MGCL 5683) (J.E. Hayden) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112706), Alachua Co., Gainesville, 6400 SW 20th Ave[nue], 29.641°N, 82.41°W, 25-26.vi.2016, UV light (J.E. Hayden) (MGCL); 1 ♀ (UF-FLMNH-MGCL-1112705), Alachua Co., Gainesville, Hull Road, FLMNH McGuire Center, 28.635°N, 82.37°W, 5.xi.2013, UV light (J.E. Hayden) (MGCL); 1 ♂, 1 ♀, Alachua Co., Paynes Prairie Preserve, overflow parking lot, Puggy Road, 29.527°N, 82.291°W, 6.x.2008 (MGCL-FLMNH-67480), 4.viii.2008 (MGCL-FLMNH-67446) (G.T. Austin) (MGCL); 1 ♂, 1 ♀, Brevard Co., Melbourne, Weston Park, 28.158°N, 80.652°W, UV, 5.ii.2017 (UF-FLMNH-MGCL-1112691), 13.xii.2017 (UF-FLMNH-MGCL-1112696, slide JFL 1789) (F.M. Blaine) (MGCL); 1 ♀ (UF-FLMNH-MGCL-1112710), Collier Co., Fakahatchee Strand, 25.98°N, 81.41°W, 12.ii.2013 (J. Troubridge) (MGCL); 1 ♂ (CNCLEP00210183), Collier Co., Fakahatchee Strand, 25.98°N, 81.41°W, 21.ii.2014 (J. Troubridge) (CNC); 1 ♂, 2 ♀ (CNCLEP00210184, slide CNC 17613; CNCLEP00210182, slide CNC 17612; CNCLEP 00210185), Collier Co., Fakahatchee Strand, 25.98°N, 81.41°W, 21.ii.2014 (J. Troubridge) (CNC); 1 ♂, 1 ♀(CNCLEP00210186, CNCLEP00210187, right wings on slide CNC 17615), Collier Co., Fakahatchee Strand, 25.98°N, 81.41°W, 4.ii.2014 (J. Troubridge) (MGCL); 1 ♀ (UF-FLMNH-MGCL-1112711), Collier Co., Fakahatchee Strand, 25.98°N, 81.41°W, 23.iii.2015 (J. Troubridge) (MGCL); 2 ♂, 1 ♀, Dade Co., Florida City, 25.4148°N, 80.44°W, 27.i.2014 (CNCLEP00281091), 2.iii.2014 (CNCLEP00281093, slide CNC 17626; CNCLEP00281092, slide CNC 17625) (J. Troubridge) (CNC); 1 ♂ (CNCLEP00281095), DeSoto Co., Nocatee, 27.16°N, 81.91°W, 1.iv.2012 (J. Troubridge) (CNC); 1 ♂(CNCLEP00281094), DeSoto Co., Peace River, 27.09°N, 82.08°W, 9.iii.2010 (J. Troubridge) (CNC); 1 ♀ (UF-FLMNH-MGCL-1112682), Dixie Co., near Gulf, 29.29°N, 83.17°W, 9.ii.1996 (J.G. Filiatrault) (MGCL); 1 ♂ (CNCLEP00281110, slide CNC 18287), Fort Ogden, 9.iv.1952 (J.R. Vockeroth) (CNC); 1 ♂ (UF-FLMNH-MGCL-1112678), Hernando Co., Withlacooche St[ate] Forest, Croom W.M.A, vic[inity]. Kirk Tower, 8.ix.1990 (W. Lee Adair Jr.) (MGCL); 1 ♂ (CUCI-Hypen-08), Highlands Co., Lake Placid, Bio[logical] St[atio]n, 2.iv.1959 (J.G. Franclemont) (CUIC); 1 ♂, 1 ♀(CNCLEP00025965, slide CNC 17606; CNCLEP 00025994, slide CNC 17614), Highlands Co., Lake Placid, Archbold Bio Stn, 125 m W Jay Cottage, 27.1716°N, 81.3493°W, 21.vi.2006, at B[lack]L[ight] and M[ercury]V[apour]L[ight] (J.-F. Landry) (CNC); 1 § (MHNG-ENTO-00011937, slide MHNG-ENTO-00011937), Highlands Co., Lake Placid, Archbold Bio Stn, behind labs & near Old State Road 8, 27.184°N, 81.34°W, 21.vi.2006, at uvl (B. Landry) (MHNG); 1 ♂(UF-FLMNH-MGCL-1112688), Leon Co., Trail Timbers Res[earch]. Station, Lake Iamonia, 22-24.ix.1986 (J.B. Heppner) (MGCL); 1 ♂ (CNCLEP00281096), Levy Co., Goethe State Forest, 29.15°N, 82.69°W, 12.iv.2014 (J. Troubridge) (CNC); 1 ♂, 2 ♀, Manatee Co., Oneco, 25.iii.1955 (CUCI-Hypen-07), 28.iii.1954 (CUCI-Hypen-05), 30.iii.1954 (CUCI-Hypen-06) (J.G. Franclemont) (CUIC); 3 ♂, Manatee Co., Oneco [near Bradenton], –.v.1954 (CNCLEP00210198, slide CNC 17618; CNCLEP00210199), 2.v.1953 (CNCLEP 00210197) (Paula Dillman) (CNC); 1 ♀(CNCNoctuoidea12869), Marion Co., Ocala National Forest Road 88, 3.9 mi SE of SR 316, 29.43°N, 81.8°W, 4.xii.2005, MV/UV light (Terhune S. Dickel) (CNC); 1 ♂, 1 ♀, Marion Co., West Anthony Road, 1.4 mi. WSW of Anthony, 29.283°N, 82.132°W, 4.i.2006 (CNCNoctuoidea12870), 10.i.2006 (CNCLEP 00038062) (Terhune S. Dickel) (CNC); 6 ♂, 6 ♀, Sarasota Co., North Port, 27.04°N, 82.08°W, 22.xi.2009 (CNCLEP00281079), 10.xii.2009 (CNCLEP00281080, CNCLEP00281081), 7.ii.2010 (CNCLEP00281082), 9.iii.2010 (CNCLEP00281083, CNCLEP00281084, CNCLEP00281085), 6.xi.2013 (CNCLEP00281086), 18.xi.2013 (CNCLEP00281087), 24.xi.2014 (CNCLEP 00281088, CNCLEP00281089, CNCLEP00281090) (J. Troubridge) (CNC); 4 ♂, Sarasota Co., Siesta Key, 17.iv.1953 (CNCLEP00210200), 29.i.1954 (CNCLEP 00281074), 26.ii.1954 (CNCLEP00030442), 5.i.1960 (CNCLEP00281075) (C.P. Kimball) (CNC); 1 ♂ (UF-FLMNH-MGCL-1112679, slide FLMNH-MGCL 1441), Sumter Co., Richloam W.M.A., 20.ii.1991 (W. Lee Adair Jr.) (MGCL); 1 ♂, 1 ♀ (UF-FLMNH-MGCL-1112684, UF-FLMNH-MGCL-1112685), Suwannee Co., Suwannee River St[ate] P[ar]k, 13 mi NE Live Oak, 24.x.1975, at blacklight (J.B. Heppner) (MGCL); 1 ♀(MHNG-ENTO-0113636, slide MHNG-ENTO-0113636), Volusia Co., 2 mi E of DeLand, 3.i.1993 (Y.F Hsu) (MHNG). – Louisiana: 4 ♂, 1 ♀, E Baton Rouge Parish, Baton Rouge, 27.v.1969 (UF-FLMNH-MGCL-1112739), 1.x.1970 (UF-FLMNH-MGCL-1112737), 4.x.1970 (UF-FLMNH-MGCL-1112738), 3.xii.1970 (UF-FLMNH-MGCL-1112735) home, 11.v.1972 (UF-FLMNH-MGCL-1112736) (G. Strickland) (MGCL); 1 ♂, 2 ♀, New Orleans, 8.xi.1974 (UF-FLMNH-MGCL-1112731), 18.viii.1974 (UF-FLMNH-MGCL-1112733), 7.viii.1974 (UF-FLMNH-MGCL-1112732) (V.A. Brou) (MGCL); 2 ♂, 1 ♀, S[ain]t. John Parish, Edgard, UV light trap, 30.vi.1973 (UF-FLMNH-MGCL-1112729), 18.x.1981 (UF-FLMNH-MGCL-1112727), 22.vi.1981 (UF-FLMNH-MGCL-1112728) (V.A. Brou) (MGCL); 1 ♂, 2 ♀ (CNCLEP00210179, CNCLEP00210180, CNCLEP00210181), Tammany Co., Fontainebleau State Park, 23.iv.1993, at MVL, southern mixed forest (J.-F. & B. Landry) (CNC). – Maryland: 1 ♂ (CNCLEP00090566), Calvert Co., Scientists Cliffs, 38.5195°N, 76.5135°W, 9.viii.2011, at MVL + BL (J.-F. Landry) (CNC). – Mississippi: 1 ♂(CNCLEP00210178, slide CNC 17611), Lowndes Co., near Crawford, T17N R16E sec 34, 26.iv.1993, at MVL, black belt prairie at edge of oak-hickory forest (B. & J.-F. Landry) (CNC). – New York: 1 ♂ (CNCNoctuoidea15043), Suffolk Co., Napeague, Promised Land, 12.viii.2007 (Hugh D. McGuiness) (CNC). – North Carolina: 1 ♀(UF-FLMNH-MGCL-1112712), Brunswick Co., 1 mi NE Pretty Pond, 20.ix.1994, 15W UV trap, Leiophyllum/slash pine woodland (J. Bolling Sullivan, Richard Broadwell, Brad Smith) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112716, slide MGCL 1447), Carteret Co., J[un] ct[ion]. State H[igh]w[a]y 101 & 181, 13.viii.1978 (J. Bolling Sullivan) (CNC); 1 ♂, 1 ♀, Craven Co., Croatan Forest Road 167, 24.vi.1993 (UF-FLMNH-MGCL-1112715), 21.vii.1993 (UF-FLMNH-MGCL-1112714) (J. Bolling Sullivan) (MGCL); 1 ♂, 1 ♀, Craven Co., North Harlowe, 7.xi.1990 (UF-FLMNH-MGCL-1112720), 12.ix.1990 (UF-FLMNH-MGCL-1112719, slide MGCL 4466) (J. Bolling Sullivan) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112713), New Hanover Co., Fort Fisher Maritime Forest, 27.ix.1994, 15W UV trap, coastal fringe evergreen forest (J. Bolling Sullivan, Richard Broadwell, Brad Smith) (MGCL). –South Carolina: 1 ♀ (CUCI-Hypen-04), Charleston Co., McClellanville, Wedge Plantation, 21.iii.1968 (J.G. Franclemont) (CUIC). – Tennessee: 1 ♀ (CNCLEP 00016512, slide CNC 17608), Tennessee, Blount Co., Great Smokies National Park, West Foothills Parkway at Little River, 37.716°N, 83.819°W, 22.v.2005, at MVL (J.-F. Landry) (CNC). – Texas: 4 ♂, Hidalgo Co., Santa Ana Refuge, 27.xi.1973 (USNMENT01769436, slide USNM 144180, USNMENT01769470), 4.xi.1974 (USNMENT01769369), 15.ix.1974 (USNMENT 01769410) (A&ME Blanchard) (USNM); 1 ♂, 1 ♀, Hidalgo Co., Santa Ana Wildlife Refuge, 27.x.1979 (UF-FLMNH-MGCL-1112743, slide JFL 1778), 6.ix.1992 (UF-FLMNH-MGCL-1112742, slide JFL 1777) (E.C. Knudson) (MGCL); 1 ♀ (UF-FLMNH-MGCL-1112744, slide JFL 1781), Jefferson Co., Beaumont, 2.vii.1992 (C. Bordelon) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112747, slide JFL 1774), Kerr Co., 10 mi W Aunt, 6.ix.1981 (E.C. Knudson) (MGCL); 2 ♀, Montgomery Co., Conroe, 13.x.1971 (USNMENT01769427), 30.vii.1975 (USNMENT01769469, slide USNM 144181) (A&ME Blanchard) (USNM); 1 ♂ (UF-FLMNH-MGCL-1112745, slide JFL 1779), Orange Co., Sabine River Orange THWMA, 5.vi.2004 (Bordelon & Knudson) (MGCL); 1 ♂ (USNMENT01769458), San Patricio Co., Sinton Welder Wildlife Foundation, 16.xi.1966 (A&ME Blanchard) (USNM); 1 ♂ (UF-FLMNH-MGCL-1112746, slide JFL 1780), Tyler Co., BITH Ranch House, Turkey Creek Ut., 21-22.vi.1996 (E.C. Knudson) (MGCL); 1 ♂(USNMENT01769445), Tyler Co., Town Bluff (Dam B), 24.xi.1967 (A&ME Blanchard) (USNM). – Virginia: 1 ♂, 2 ♀, Arlington, 4.xi.1947 (CUCI-Hypen-01, slide JFL 1786), 11.x.1949 (CUCI-Hypen-03, slide JFL 1787), 17.ix.1949 (CUCI-Hypen-02) (J.G. Franclemont) (CUIC).
Other specimens examined: 22 ♂, 13 ♀. USA, Alabama: 1 ♀ (UF-FLMNH-MGCL-1112740), Mobile Co., Mobile, 1.viii.1974 (R.H. Leuschner) (MGCL). – Florida: 1 ♀ (CNCLEP00210196), [no locality], –.vii.1914 (Wooley-Dod) (CNC); 3 ♂(CNCLEP00281113, CNCLEP00281114, CNCLEP 00281115), 16 mi W Fort Lauderdale, 23.iii.1953 (W.R.M. Mason) (CNC); 3 ♂, Alachua Co., Gainesville, 2832 NW 41st, 29.692°N, 82.364°W, UVL trap, 18-19.vi.2019 (UF-FLMNH-MGCL-1112700), 8-9. vii.2019 (UF-FLMNH-MGCL-1112701), 29-30. ix.2019 (UF-FLMNH-MGCL-1112702) (J.E. Hayden) (MGCL); 1 ♀ (UF-FLMNH-MGCL-1112694), Brevard Co., Melbourne, Weston Park, 28.158°N, 80.652°W, 14.xi.2017, UV (F.M. Blaine) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112675), Cassadaga, 23.ix.1965, at blacklight, (S.V. Fuller) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112699), Collier Co., Collier-Seminole St Pk, 14.ii.1988, (V.P. Lucas) (MGCL); 1 ♀ (CNCLEP00281111), Elfers, 19.iv.1952 (J.R. Vockeroth) (CNC); 1 ♀ (UF-FLMNH-MGCL-1112683), Highlands Co., Highlands Hammock St Pk, 3.v.1974, at UV blacklight (J.B. Heppner) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112681), Highlands Co., Highlands Hammock St Pk, 10.xi.1983 (H.D. Baggett) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112676), Homestead, 6.xi.1959 (D.O. Wolfenbarger) (MGCL); 1 ♂ (CNCLEP00309353), Lake Co., Lake Dorr Camp near Altoona, 9.iv.1979 (D. & V. Hardwick) (CNC); 1 ♀ (UF-FLMNH-MGCL-1112680), Lee Co., Sanibel Island, 17-18.iv.1965, at light (E. & I. Munroe) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112689), Leon Co., Trail Timbers Res. Station, Lake Iamonia, 22-24. ix.1986 (J.B. Heppner) (MGCL); 2 ♂, 1 ♀, Levy Co., Goethe St Forest, Gasline & Beehive R[oa] ds, 29.16°N, 82.598°W, pine flatwoods, 4.v.2012 (UF-FLMNH-MGCL-1112708) UV (J.E. Hayden & A. Jansen), 4.v.2012 (UF-FLMNH-MGCL-1112707) beer-fruit bait (J.E. Hayden), 28.vi.2014 (UF-FLMNH-MGCL-1112709) MVL/UVL (J.E. Hayden) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112690), Liberty Co., Torreya St Pk, 25–27.ix.1986 (J.B. Heppner) (MGCL); 1 ♀ (UF-FLMNH-MGCL-1112677), Manatee Co., Oneco, –.v.1954 (Paula Dillman) (MGCL); 1 ♀(CNCLEP00281112), Punta Gorda, 12.iv.1952 (G.S. Walley) (CNC); 1 ♂, 1 ♀, Putnam Co., Welaka Forest Cons. Station, 9-10.vi.1986, Welaka site 4 live oak xeric hammock (UF-FLMNH-MGCL-1112686), Welaka site 5 slash pine palmetto flatwoods (UF-FLMNH-MGCL-1112687) (J. Heppner & J. Powell) (MGCL). – Louisiana: 1 ♂ (UF-FLMNH-MGCL-1112734), Prairieville, 28.iv.1974, UV light (Vernon A. Brou) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112730), “Cut Off” [sic], 24.vi.1975 (V.A. Brou) (MGCL). – North Carolina: 1 ♂ (UF-FLMNH-MGCL-1112718), Carteret Co., Beaufort, 15.xi.1991 (J. Bolling Sullivan) (MGCL); 1 ♂ (UF-FLMNH-MGCL-1112717), Carteret Co., Jct. State Hwy 101 & 181, 10–11.xi.1972 (J. Bolling Sullivan) (MGCL); 2 ♀, Dare Co., Buxton Hatteras Island, 1.viii.1975 (UF-FLMNH-MGCL-1112721), 7.x.1975 (UF-FLMNH-MGCL-1112722) (Samuel M. Gifford) (MGCL). – South Carolina: 1 ♂, 1 ♀(UF-FLMNH-MGCL-1112723, UF-FLMNH-MGCL-1112724), The Wedge Plantation, McClellanville, 12-25. ix.1979 (E.G. & I.M. Munroe) (MGCL). – Texas: 1 ♂(USNMENT01769430), St. Augustine Co., 2 mi SW Broaddus, Lake Sam Rayburn, 20.ix.1972 (E.L. Todd) (USNM).
Etymology: The species is named after the Calusa people, a Native American group, now extinct, who inhabited the southwestern coast of Florida and whose influence is thought to have extended to much of the southern half of that state (Wikipedia: https://en.wikipedia.org/wiki/Calusa). Part of the type material was collected in that region. It is a noun in apposition.
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 spur-like 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 bowl-like 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. 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. Differences in genitalia are pronounced between the species pair H. calusa-H. sonora versus H. macula and, in some dry specimens, can be viewed without dissection if those parts are partly extruded by brushing off the scale cover (Figs 50-53). This must be done very carefully, however, particularly in males, because the structures are delicate and brittle and can be easily damaged or broken. In males, the darkly melanized, black ampullae with indistinct apical forks of H. macula contrast markedly with the paler fuscous brown and strongly forked ampullae of H. calusa-H. sonora. In females, the asymmetrically skewed and recessed ostial 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 17-18) pale beige mixed with brown, darker greyish brown between antennae. Antenna with scales greyish brown; ventral sensilla about 1.5× flagellomere length. Labial palpus with second palpomere straight, porrect, with scales slightly erect ventrally and dorsally, outer side dark brown, inner side pale beige; third palpomere ∼ 0.4× length of second, dark brown with pale beige base and apex. Thorax dorsally pale beige mixed with greyish brown suffusion. Foreleg and midleg grey-brown. Hindleg pale beige, lighter than other legs. Wingspan (n=25): 12.0-17.5 mm; forewing length (n=25): 6.0-8.0 mm. Wing vestiture as shown (Figs 3-6). Forewing predominantly dark brown with blackish-brown chevron-shaped antemedial line, though interrupted, indistinct or reduced to a dot in many specimens; with contrastingly pale beige transversely oblique subterminal fascia extending from apex of costa to near middle of dorsal margin, lined with a thin black postmedial line on inner side, in many specimens with sparse brown suffused over pale fascia; with black discal spot at end of cell and six black terminal dashes of variable intensity. In several females (Figs 5-6) forewing surface mostly grey-brown or brown, most markings indistinct except discal spot, with nearly concolorous subterminal fascia and faint anteromedial line; black discal spot with small pale centre in some specimens (Fig. 6). Hindwing pale grey to pale brownish grey, slightly infuscated, with dark brown terminal line well marked in most specimens. Abdomen pale beige with sparse brown irroration.
Male pre-genital abdomen (Figs 23-25). Sternite VIII lateral projections of anterior margin narrowly triangular, with slender, subacuminate apices, interspace broadly U-shaped. Tergite VIII projection with variably widened shaft which is about 1.3× length of tergite VIII and apical fork with proportionally longer ‘Y’-branches.
Male genitalia (n=18) (Figs 33-34). Tegumen weakly sclerotized, elongate V-shaped, with penicular arms thinly attenuate to base of valva. Vinculum with narrowly rounded saccus, with pronounced mesial protrusion. Uncus surface weakly and unevenly sclerotized, outline indistinct, subelongate or subquadrate with apical margin lobate, about 0.2× length of tegumen (measured from base of valva to apex), with lateral margins concave. Juxta consisting of two elongate-subcrescentic plates with a narrow mesial junction/gap, an oblique dorsal edge, and recurved ventral edge. Valva wider in distal half, proportionally short with apex of cucullus extending to, but not beyond, apex of uncus. Costa thickly sclerotized, contrasting with membranous mesial wall, extending to apex of cucullus. Ampulla strongly sclerotized, slightly downcurved, with series of small protuberances along upper side of shaft (variable in number and distribution), apically bifurcate with dorsal fork variously rounded 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, with many very densely set, thick setae. Cucullus oval, ventral edge sclerotized, ventral lobe small, with long setae oriented antero-dorsally. Sacculus ∼0.6× length of valva, elongate-subrectangular with slightly concave ventral margin. Phallus elongate, straight in dorsal aspect, medially with slight sinuation in lateral aspect, 0.8× length of vinculum+valva, with apex oblique and pointed with smooth surface; caecum slightly wider than shaft; vesica with single, very short tooth-like cornutus; bulbus ejaculatorius about 0.5× length of phallus shaft, wholly membranous, with crescent-shaped dilation.
Female pre-genital abdomen and genitalia (n=10) (Figs 42-47). Abdominal segment VII (Figs 42-44) sclerotized, about 2× as long as segment VI. Tergite VII broadly transverse, in undissected preparation extending ventrally and surrounding segment VIII. Sternite VII trapezoid, extended as a lip over and covering basal half of sternite VIII, laterally sinuated. Sternite VIII 0.75× length of tergite VII, forming a large, bulbous, deep, symmetrical sinus vaginalis/antrum, its posterior edge with deep emargination, surface of sinus vaginalis smooth. Anterior apophyses short but variable in length, from short stubs to ∼0.25× as long as posterior ones. Ostium bursae opening mesially, about 0.25× as wide as sinus vaginalis. Ductus bursae membranous, as long as or shorter than sternite VIII (length variable depending on amount of stretching). Corpus bursae elongate-ovoid (shape variable depending on whether female mated or not), surface with faint isodiametric microsculpture, signum absent; inception of ductus seminalis situated half-way between antrum and ductus bursae. Posterior apophyses about 1.5× as long as papillae anales. Papillae anales melanized, together with ovipositor slightly longer than tergite VIII.
Variation: Differences between male and female external aspect are slight overall, except in forewing coloration, some females having reduced postmedian fascia and being nearly unicolorous. More individual variation was observed for this species because of the abundant material available.
Biology: Reported to feed on bracket fungus (Polyporaceae) (Forbes, 1954: 383, as macula, quoting Franclemont). We have not seen the specimen(s) on which this observation was based so our presumption that this pertains to H. calusa and or to one of the other two North American species is unverified. Adults have been collected at light in all months of the year depending on locations.
Distribution: Confirmed records verified by the authors are from Florida, Alabama, Mississippi, 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, Maryland, New Jersey, 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.
Remarks: DNA barcodes in BIN BOLD:AAB2278. The nearest neighbour is H. macula BOLD:AAL2056 at an average pairwise distance of 4.96%.
Figs 3-10.
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 UF-FLMNH-MGCL-1112705 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 UF-FLMNH-MGCL-1112748 from Miami-Dade Co., Florida. (10) H. macula female, specimen UF-FLMNH-MGCL-1112754 from Miami-Dade Co., Florida.
Hypenopsis galapagensis B. Landry & J.-F. Landry, sp. nov.
Figs 11-16, 19, 21, 35-36, 48
Material examined: 35 ♂, 24 ♀.
Holotype: ♂, ‘ECUADOR | GALÁPAGOS | San Cristóbal | pampa zone | 15.II.1989, M[ercury]V[apour] L[ight] | B. Landry’ [printed in black on white card stock, with ‘ECUADOR’ sideways on left]; ‘BL 1817 ♂’ [green paper, hand written]; ‘GALAP | Hypen-CO1- | 01’ [orange paper, typed]; ‘HOLOTYPE | Hypenopsis | galapagensis | Landry & Landry’ [red card stock, hand written]. Deposited in the CNC.
Paratypes: 2 ♂, 4 ♀ from the Galápagos Islands, Ecuador. – San Cristóbal: 1 ♀ (dissected, slide BL 1825), 4 km SE P[uer]to Baquarizo [sic], 12.ii.1989, M[ercury]V[apour] L[ight] (B. Landry); 1 ♂ (dissected, slide BL 1158), 1 km S El Progreso, 14.ii.1989, MVL (B. Landry); 1 ♀, pampa zone, 18.ii.1989, MVL (B. Landry); 1 ♀, base of Cerro Pelado, 22.ii.1989, MVL (B. Landry); 1 ♀ (dissected, slide MHNG-ENTO-3613; sequenced for DNA barcode, GALAP-Hypen-CO1-02), La Toma, ca. 5.6 km E EL Progreso, G[lobal]P[ositioning]S[system]: 299 m elev[ation]., S 00°55.356', W 089°31.089', 23.ii.2005, u[ltra]v[iolet]l[ight] (B. Landry); 1 ♂ (dissected, slide MHNG-ENTO-11577; sequenced for DNA barcode, GALAP-Hypen-CO1-03), shore of El Junco, GPS: elev. 655 m, S 00°53.714', W 089°28.869', 17.iii.2004, net (B. Landry). Deposited in CDRS, CNC, and MHNG.
Other specimens examined: 32 ♂, 20 ♀, from the Galápagos Islands: – Floreana: 1 ♂ (dissected, slide MHNG-ENTO-11940, sequenced for DNA barcode, GALAP-Hypen-CO1-011), Scalesias near Cerro Pajas, G[lobal]P[ositioning]S[system]: elev[ation]. 329 m, S01°17.743', W90°27.111', 12.iv.2004, u[ltra] v[iolet]l[ight] (P. Schmitz); 1 ♂ (dissected, slide MHNG-ENTO-3612, sequenced for DNA barcode, GALAP-Hypen-CO1-012), Cerro del Asilo, GPS: elev. 366 m, S01°18.931', W90°27.232', 14.iv.2004, uvl (P. Schmitz). – Isabela: 3 ♂ (one dissected, slide BL 1826, sequenced for DNA barcode, GALAP-Hypen-CO1-09), V[olcan]. Alcedo, Los Guayabillos, 5.iii.2007, UVL (L. Roque); 1 ♂, 8.5 km N P[uer]to Villamil, 11.iii.1989, M[ercury]V[apour]L[ight] (B. Landry); 1 ♂ (dissected, slide BL 1828, sequenced for DNA barcode, GALAP-Hypen-CO1-010), Sierra Negra, pampa zone, 1000 m [elevation], 12.iii.1989, MVL (B. Landry); 1 ♂, 1 ♀, 11 km N Pto Villamil, 13.iii.1989, MVL (B. Landry); 1 ♂, V. Alcedo, cumbre, 1200 m[e] t[er]s elev., La caseta, 9.iv.1999, UVL-F[luorescent] L[ight] (L. Roque); 1 ♀, Tagus Cove, 13.v.1992, MVL (B. Landry); 2 ♂ (dissected, slides BL 1157 & 1820, one sequenced for DNA barcode, GALAP-HypenCO1-05), 1 ♀ (dissected, slide BL 1833, sequenced for DNA barcode, GALAP-Hypen-CO1-06), V. Darwin, 300 m elev., 15.v.1992, MVL (B. Landry); 1 ♂, V. Darwin, 630 m elev., 16.v.1992, MVL (B. Landry); 1 ♂ (sequenced for DNA barcode, GALAP-HypenCO1-07), V. Darwin, 1240 m elev., 19.v.1992, MVL (B. Landry); 1 ♀ (dissected, slide BL 1824, sequenced for DNA barcode, GALAP-Hypen-CO1-08), ± 15 km N Pto Villamil, 25.v.1992, MVL (B. Landry); 1 ♂, Sierra Negra, Alemania, xi.1974 (T. J. deVries). – Pinta: 1 ♂(dissected, slide BL 1819, sequenced for DNA barcode, GALAP-Hypen-CO1-014), 400 m elev., 17.iii.1992, MVL (B. Landry); 2 ♂ (one dissected, slide BL 1827), 400-650 m elev., 18.iii.1992, day (B. Landry); 1 ♀(dissected, slide BL 1823, sequenced for DNA barcode, GALAP-Hypen-CO1-013), 400 m elev., 18.iii.1992, MVL (B. Landry). – Santa Fé: 1 ♂ (dissected, slide BL 1818, sequenced for DNA barcode, GALAP-Hypen-CO1-04), Tourist trail, 28.v.1992, MVL (B. Landry). – Santa Cruz: 1 ♀, Los Gemelos, 21.i.2004, UVL (L. Roque); 1 ♂, Los Gemelos, 31.i.1989, MVL (B. Landry); 1 ♂, 1 ♀, Los Gemelos, Scalesia Forest, 4.ii.2002, UVL (L. Roque); 3 ♂ (one sequenced for DNA barcode, GALAP-Hypen-CO1-016), 6 ♀ (2 dissected, slides BL 1821, 1822, 1 sequenced for DNA barcode, GALAP-Hypen-CO1-015), Media Luna, pampa zone, 8.ii.1989, MVL (B. Landry); 1 ♂(dissected, slide MHNG-ENTO-3611, sequenced for DNA barcode, GALAP-Hypen-CO1-017), low agriculture zone, GPS: S00°42.132' W 90°19.156', 13.iii.2004, uvl (B. Landry, P. Schmitz); 1 ♀, Finca S[teve]. Devine, 17.iii.1989, MVL (B. Landry); 1 ♂, 1 ♀ (dissected, slide LR151, sequenced for DNA barcode, GALAP-Hypen-CO1-018), Gemelos, iii. [20]04 [L. Roque.]; 1 ♀ (dissected, slide L[azaro] R[oque] 150), Los Gemelos, Scalesia forest, 4.v.2002, UVL (L. Roque); 3 ♂ (2 dissected, genitalia slide LR 149 & wing slide MHNG-ENTO-11969, 1 sequenced for DNA barcode, GALAP-Hypen-CO1-019), 2 ♀(1 sequenced for DNA barcode, GALAP-HypenCO1-020) Los Gemelos, 27.v.1992, MVL (B. Landry); 1 ♂ (dissected, slide BL 1829), 1 ♀, Indefatigable, vi.1970, Ref. No. L.157 (R. Perry, Tj. De Vries); 2 ♂, 1 ♀, Los Gemelos, Scalesia forest, 580 m[etros]s[obre el]n[ivel del]m[ar], 00[°]37'29.4”S, 090[°]23'05.3[”] W, 14.x.1996, in fluorescent light trap (L. Roque); 1 ♂, Barranco, C[harles]D[arwin]R[esearch]S[tation], 23.x.2001 (L. Roque). Deposited in CDRS, CNC, MHNG, NHMUK.
Etymology: The species epithet refers to the collecting locality of all known specimens, the Galápagos Islands.
Diagnosis: In forewing (Figs 11-16) H. galapagensis is more greyish brown or blackish brown compared to the warmer brown of H. macula and H. calusa. 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 tooth-like 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, funnel-shaped with a U-shaped 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.
Description: Male (n=3): Head greyish brown, slightly paler on frontoclypeus. Antenna with scales greyish brown; ventral sensilla about 1.5x their own flagellomere length. Labial palpus (Fig. 19) with second palpomere straight, porrect, about twice as long as eye diameter, with short scales slightly projecting ventrally and dorsally, dark brown laterally, paler medially; with third palpomere about 4/5 second, dark brown with paler tip. Thorax dorsally greyish brown, paler distally. Foreleg and midleg dark brown, with single row of apical scales of tibia and tarsomeres paler. Hindleg lighter greyish brown than other legs. Wingspan (n=2): 9.5-12.0 (holotype) mm; forewing length: 4.5-5.5 (holotype) mm. Wing vestiture as shown (Figs 11, 15, 16). Abdomen greyish brown.
Male pre-genital abdomen and genitalia (n=14) (Figs 35-36). Abdominal sternite VIII lateral projections of basal margin about 3/7 to 4/7th length of sternite. Abdominal tergite VIII median projection highly variable in length and width of shaft, about 1.5× to 2/3 length of tergite VIII. Tegumen arms slightly widened in distal half, then constricted apically. Vinculum slightly shorter (ca. 15%) than valva, with base about as long as narrow arms, with narrowly rounded apex. Uncus thinly sclerotized, about 1/3 as long as tegumen, composed of a median, narrow plate and pair of narrow plates laterally, without sclerotized connection to central plate, margins indistinct. Juxta short, consisting of two narrow, diverging plates more thickly sclerotized in basal half. Valva of medium width and length, about 30% longer than tegumen; basal section, to midlength, with margins more thickly sclerotized, apicoventrally with bunch of medium-length setae directed apically; ventral margin slightly constricted at midlength, followed by short, bulbous ventral flange of clasper tightly set with thick setae of medium length on dorsomedial surface; dorsal of ventral flange of clasper with narrow, S-shaped median flange of clasper well separated from median wall of valva in situ, directed apically, ending at level of ventral margin and apically thickly sclerotized and rounded; with larger, more thickly sclerotized and wider ampulla only slightly curved, pointed, reaching ventral margin, with small patch of setae directed upward at base dorsally; ventral margin following bulbous process more tightly sclerotized for short distance and set with 4-5 long setae directed medially; median wall of valva at about 9/10th of length and next to ventral margin with ventral lobe of cucullus harbouring patch of setae projecting dorsally on median side of dorsal process, reaching dorsal margin. Phallus about two-thirds length of vinculum+valva, straight, with apex finely serrate; vesica with single, slender, narrowly triangular cornutus about 1/10th length of phallus shaft. Female (n=4): Antenna with ventral sensilla minute. Vestiture (Figs 12-14) as in males. Wingspan (n=4): 10.5-13.0 mm; forewing length: 5.0-6.0 mm.
Female pre-genital abdomen and genitalia (n=9) (Fig. 48). Abdominal sternite VII a broad triangle with truncated apex, about as long as preceding sternite. 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 11-12). 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 36c-f) 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.
Distribution: Galápagos islands of Floreana, Isabela, Pinta, San Cristóbal, Santa Cruz, and Santa Fé.
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. 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.
Figs 11-16.
Specimens of Hypenopsis galapagensis. (11) Male holotype from San Cristóbal Island, pampa zone. (12) Female from Isabela Island, 11 km N of Puerto Villamil. (13) Female paratype from San Cristóbal Island, pampa zone. (14) Female paratype from San Cristóbal Island, base of Cerro Pelado. (15) Male paratype from San Cristóbal Island, shore of El Junco, 655 m elev. (16) Male from Santa Cruz Island, Los Gemelos.
Figs 17-20.
Heads of Hypenopsis specimens in lateral and dorsal view. (17) Holotype of H. calusa, lateral view. (18) Same, dorsal view. (19) Holotype of H. galapagensis. (20) H. macula, specimen UF-FLMNH-MGCL-1112752 from Florida.
Fig. 21.
Wing venation of Hypenopsis galapagensis, specimen from Santa Cruz Island, slide MHNG-ENTO-11969.
Figs 22-29.
Male abdominal segments of Hypenopsis species. (22) SI–II (on left) and TI–II with phragma (on right), H. calusa from Florida, slide CNC 17613. (23-29) SVIII (left) and TVIII (right), posterior end of abdomen oriented down. (23) H. calusa from Texas, slide JFL 1774. (24) H. calusa from Texas, slide JFL 1777. (25) H. calusa from Alabama, slide JFL 1785. (26) H. macula from Florida, slide JFL 1783. (27) H. macula lectotype from Panama, slide JFL 1770. (28) H. sonora from Mexico, slide CNC 17609. (29) H. sonora from Texas, slide CNC 17623.
Hypenopsis sonora J.-F. Landry & B. Landry, sp. nov.
Figs 7, 28-29, 37-38, 49
Schrankia macula (Druce): North American Moth Photographers Group, 2023.
Material examined:
Holotype ♂: ‘CA: San Diego Co | SanDiegoTierrasanta | VII 5 2013 | BL N.Bloomfield’; ‘Barcode of Life | DNA voucher specimen | SmpleID BIOUG23753-H09 | BOLD Proc ID: LOCBF4844-15 [yellow label]’; ‘CNC | genitalia slide # | 17607 ♂’ [green label]; ‘CNCLEP | 00281099’ [printed in black on card stock]; ‘HOLOTYPE ♂ | Hypenopsis | sonora | Landry & Landry’ [orange card stock, handwritten]. Deposited in the CNC.
Paratypes: 5 ♂, 1 ♀. – USA: Arizona: 1 ♂(CNCLEP00210174), Yuma Co[unty]., Yuma, on Colorado River, 32.692°N, 114.49°W, 22.iii.2001, at blacklight (Ian A. Watkinson) (CNC). – Texas: 2 ♂, 1 ♀ (CNCLEP00281076, ♀ slide CNC 17622; CNCLEP00281077, ♂ slide CNC 17623; ♂CNCLEP00281078), Brownsville, 9.ii.1937 (T.N. Freeman) (CNC); 1 ♂ (UF-FLMNH-MGCL-1112741, slide JFL 1773), Cameron Co., Laguna Atascosa Nat[ional] Wild[life]. Ref[uge]., 11.xi.1979 (E. Knudson) (MGCL). – Mexico: 1 ♂ (CNCLEP00210158, slide CNC 17609), Oaxaca, 17-18.viii.1969 (D. Kritsch) (CNC).
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, spur-like. 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 funnel-shaped, half the width of the posterior margin of abdominal tergite VIII with a narrow and irregularly V-shaped 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).
Description: Head pale brownish grey or pale beige. Antenna pale beige; ventral sensilla about 1.5× flagellomere length. Labial palpus similar to calusa except slightly paler brown. Thorax dorsally pale beige. Foreleg and midleg grey-brown. Hindleg pale beige, lighter than other legs. Wingspan (n=2): 18.0-18.5 mm; forewing length (n=6): 7.0-8.5 mm. Wing 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. Abdomen grey 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 U-shaped. Tergite VIII Y-shaped 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 V-shaped with small mesial protrusion and rounded saccus. Uncus surface weakly and unevenly sclerotized, outline resembling a double-headed battle axe, ∼ 0.25-0.3× length of tegumen (measured from base of valva to apex). Juxta consisting of two elongate-subcrescentic 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 pointed depending on orientation). Ventral flange of clasper large, club-shaped, with very dense set of thick setae. Cucullus oval, with sclerotized ventral edge, its ventral lobe larger than in calusa and with long, dense setae oriented antero-dorsally. Sacculus ∼0.5× length of valva (proportionally shorter than in calusa), elongate-subrectangular with straight ventral margin. Phallus elongate, distally slightly tapered, straight in dorsal aspect, medially with slight sinuation in lateral aspect, ∼0.8× length of vinculum+valva, apically oblique and tapered with ventral surface finely spinulate, caecum dilated, wider than shaft; vesica with single, spine-like 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 crescent-shaped dilation.
Femalepre-genitalabdomenandgenitalia(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, funnel-shaped, slightly asymmetrical, ∼0.5× as long as sternite VIII, with narrowly U-shaped 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 elongate-ovoid, 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.
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 separate H. sonora from other Hypenopsis with an average pairwise distance of 4.01% to nearest neighbour H. galapagensis BOLD:ADX5739. Barcoding was attempted for the Texas and Mexico specimens, which included the single female available, but failed. The association of the female with the males remains
unverified by barcode matching. It was collected together with two males in Brownsville, Texas and these males have genitalia that match those of barcoded males from southern California and Arizona. No other putative species with similar male genitalia were found among all the Hypenopsis specimens examined, so we consider the female associated with the H. sonora males likely to be conspecific.
Figs 31-32.
Male genitalia of H. macula, (a) genitalia, ventral view, phallus removed; (b) phallus of same, lateral view with dorsal (upper) side facing left; (c) close-up of phallus tip showing cornutus and apical serration. (31) slide JFL 1770, lectotype from Panama, setae of ventral lobe of cucullus missing. (32) slide CNC 17629, from Costa Rica.
Figs 33-34.
Male genitalia of H. calusa. (33a) genitalia, ventral view, phallus removed, slide CNC 17611, paratype from Mississippi. (33b) phallus of same, dorsal view. (33c) close-up of phallus tip, lateral view, with vesica slightly everted and cornutus protruding, slide CNC 17626 from Florida. (34a) genitalia, ventral view, phallus removed, slide JFL 1789, paratype from Florida. (34b) phallus of same, dorso-lateral view with bulbus ejaculatorius bent to the left (upper side facing left). (34c) median flange of clasper slide CNC 17626 paratype from Florida.
Figs 35-36.
Abdomen and male genitalia of H. galapagensis. (35) Abdominal segment VII; (a) holotype slide BL 1817; (b) paratype, slide BL 1158. (36a) Genitalia without phallus, paratype, slide BL 1158; (b) phallus of same; (c) distal portion of valva, from Isabela Island, slide BL 1157; (d) idem, from San Cristóbal Island, slide BL 1158; (e) Idem, from Floreana, slide BL 3612; (f) idem, from Santa Cruz Island, Los Gemelos, slide LR 021; the scale bar of 36a is the same for 36b-f.
Figs 37-38.
Male genitalia of H. sonora. (37a) genitalia, ventral view, phallus removed, slide CNC 17607, holotype from California, setae of ventral lobe of cucullus missing; (b) phallus of same, dorsal view with bulbus ejaculatorius bent to the left; (c) closeup of median flange of clasper of same. (38a) genitalia, ventral view, phallus removed, slide CNC 17623, paratype from Texas; (b) phallus of same, lateral view (upper/dorsal side facing left) with bulbus ejaculatorius in natural orientation; (c) close-up of phallus tip of same, lateral view, differential-interference contrast illumination showing fine denticulation.
Figs 39-41.
Female genitalia of H. macula, ventral view. Segment 6 removed in Figs 39 and 41. (39) slide JFL 1771, paralectotype, specimen MfN-940364 from Panama. (40) slide JFL 1776, specimen UF-FLMNH-MGCL-1112757 from Florida; dashed lines indicate position of S6 separated on its left side and along its posterior margin, and unrolled. (41) slide CNC 17628, specimen 09-SRNP-100900 from Costa Rica; (a) close-up of signum.
Figs 42-44.
Female genitalia and abdominal segments of H. calusa. (42) dissection CNC 17612, specimen CNCLEP00210182 from Florida, cleaned and immersed in lactic acid, ventral view. (43) same preparation, lateral view. (44) slide CNC 17608, specimen CNCLEP00016512 from Tennessee, abdominal segments VI-VII, unrolled.
Figs 45-47.
Female genitalia of H. calusa, ventral view. Segment 7 removed in Figs 46 and 47. (45) slide JFL 1787, specimen CUCI-Hypen-03 from Virginia. (46) slide CNC 17608, specimen CNCLEP00016512 from Tennessee. (47) slide CNC 17612, specimen CNCLEP00210182 from Florida.
Figs 48-49.
Female genitalia of H. galapagensis and H. sonora, ventral view. (48) H. galapagensis paratype slide MHNG-ENTO-3613, from La Toma, San Cristóbal Island. (49) H. sonora paratype slide CNC 17622, specimen CNCLEP00281076 from Texas.
Figs 50-53.
Partially denuded apex of undissected Hypenopsis abdomens showing exposed parts of genitalia. (50) H. macula male, dorsal view, specimen UF-FLMNH-MGCL-1112693. (51) H. calusa male, dorsal view, specimen UF-FLMNH-MGCL-1112688. (52) H. macula female, ventral view, specimen UF-FLMNH-MGCL-1112695. (53) H. calusa female, ventral view, specimen UF-FLMNH-MGCL-1112685.
Figs 54-57.
Hypenopsis flualis. (54) female holotype from Panama. (55) same, head in lateral view. (56a) male genitalia, MHNG-ENTO-11939, specimen HYPENODINE-MHNG-CO1-03 from French Guiana. (56b) same, male abdominal segment VIII. (56c) same, phallus shown at same scale as rest of genitalia. (57) holotype female genitalia; note hair-like pheromone scales near anterior apophyses.
Hypenopsis flualis Schaus, 1916, Hypenodinae incertae sedis
Figs 54-57
Material examined: Holotype ♀: ‘Trinidad Riv[er] | Pan[ama] May .11 | August Busck’ [printed]; ‘Hypenopsis | flualis | type Schs’ [handwritten]; ‘Type No. | 19743 | U.S.N.M.’ [red, printed]; ‘USNMENT | 01440684’ [printed with QR code]. Dissected, slide BL 1832. In the USNM.
Remarks: The barcode sequence (658bp[89n]) obtained from the old female holotype collected in 1911 matched BOLD:ADR3240. This BIN contained another record (HYPENODINE-MHNG-CO1-03), originally identified only as Hypenodinae, a male from French Guiana collected by BL in 2008 (MHNG-ENTO-11939). Both NJ and ML analyses place H. flualis in the Hypenodinae but far removed from the Hypenopsis group as well as from Schrankia and all other Hypenodinae. Most branches are weakly supported among the Hypenodinae analyzed, thus providing no clear pattern of relationship, except for the Hypenopsis-Luceria pair. In BOLD the nearest neighbour to flualis is BOLD:AEK5416 at 4.11% distance with two members representing an unidentified taxon labeled as ‘Schrankia Malaise5416’ (unavailable private records). Morphologically, the presence of hair-like pheromone scales near the anterior apophyses and the multiple spiniform signa arranged in a transverse row in the female genitalia (Fig. 57) appear to be unique. The male genitalia (Fig. 56) also differ conspicuously from those of Hypenopsis. Three females and a male collected in the Dominican Republic in 2022 by BL (MHNG) resemble this species although in the female corpus bursae the spiniform signa are 2-3 times more numerous, smaller, and form a complete ring. A male collected in Venezuela by BL in 2009 (MHNG) also matches this species closely. In view of the incomplete knowledge of New World Hypenodinae, we treat H. flualis as Hypenodinae incertae sedis.
Figs 58-60.
Holotype male of Hypenopsis musalis. (58) habitus. (59) head in lateral view. (60a) genitalia without phallus. (60b) phallus shown at same scale as rest of genitalia.
Hypenopsis musalis Schaus, 1916, Hypenodinae incertae sedis
Figs 58-60
Material examined: Holotype ♂: ‘Trinidad Riv[er] | Pan[ama] March .11 | August Busck’ [printed]; ‘Hypenopsis? | musalis | type Schs’ [handwritten]; ‘Type No. | 19744 | U.S.N.M.’ [red, printed]; ‘USNMENT | 00973066 [printed with QR code]. Dissected, slide BL 1831. In the USNM.
Remarks: The incomplete barcode sequence (463 bp [200n]) 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 well-developed, elongate hook-like 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.
ACKNOWLEDGEMENTS
For sending photos of the syntypes of H. macula we are thankful to T. Léger and ZMB volunteer Marie Rehse. For his help in extracting the DNA of the holotypes of S. flualis and S. musalis we thank Lionel Monod (MHNG). For his support of the DNA sequencing part of the project we thank Evgeny V. Zakharov (Director, Canadian Centre for DNA Barcoding, Centre for Biodiversity Genomics, Biodiversity Institute of Ontario, University of Guelph). We thank the authorities of the Galápagos National Park for the collecting and export permits, the authorities and staff, especially Lazaro Roque Albelo (between 2002-2005) of the Charles Darwin Research Station for facilitating field work, logistical support, friendship, and companionship in the field, and Stewart B. Peck (Carleton University, Ottawa) for inviting BL to take part in his inventory of the Galápagos insects, funded by an operating grant of the Natural Sciences and Engineering Research Council of Canada for field research on arthropod evolution, and for the organization of our 1989 and 1992 expeditions. Furthermore, we wish to thank N. Castillo, C. Causton, J. Cook, M. Inca, J. Loaiza, R. Palma, P. Schmitz, B. Sinclair, and E. Vilema for companionship and help in the field, the Galápagos Conservation Trust (London, U.K.) for financing BL's visit to the NHMUK in 2000, and the City of Geneva for funding BL's 2002, 2004 and 2005 field trips to the Galápagos archipelago. For the loan of material in their care we are very grateful to M. Shaffer and K. Tuck (NHMUK), T. Léger (ZMB), and Ian Watkinson (Yuma, Arizona). We also thank Valentin Nidergas (France) for a preliminary morphological description and microphotographs (Figs 36c-f) of H. galapagensis; Alberto Zilli (NHMUK) for checking the type of Luceria novatusalis and for his detailed review which greatly improved the manuscript; Laszlo Ronkay for his comments on morphological terms; Jim Vargo (Indiana) for checking his collection for possible specimens of sonora; Adam Brunke (CNC) for assistance and advice on molecular analysis; Don Lafontaine (CNC, retired), Chris Schmidt (CNC), and Reza Zahiri (Canadian Food Inspection Agency) for references, discussion of morphology, phylogeny, and molecular analysis; Jim Hayden (Florida State Department of Agriculture) for loaning specimens, for sharing information on Hypenopsis collected by him in Florida, especially critical material of macula, and for supplying us with two GenBank sequences obtained from his specimens. Jim alerted us to the occurrence of two species in Florida (calusa, macula), based on his own dissections and DNA sequences when at the time we were aware of only one. Finally, we thank Christina Lehmann-Graber (MHNG) for the drawing of the wing venation of H. galapagensis (Fig. 21) and for cleaning habitus photos; Lisa Bartels (CNC) for data capture, barcoding preparation, specimen and genitalia photography, image editing, and plate preparation; and Alberto Zilli and Emmanuel Toussaint for their critical reviews of the manuscript. Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the USDA; USDA is an equal opportunity provider and employer.
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