Systematics of the Scythrididae (Lepidoptera) of the Galápagos Islands, Ecuador

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Abstract

The Scythrididae moths of the Galápagos Archipelago are taxonomically revised. Ten species are recognised, all of them new: Scythris ancystra J.-F. Landry & Bucheli, sp. nov., S. cristobalensis J.-F. Landry & B. Landry, sp. nov., S. falcata J.-F. Landry & Bucheli, sp. nov., S. forcipata J.-F. Landry & Bucheli, sp. nov., S. furculata J.-F. Landry & Bucheli, sp. nov., S. galapagensis J.-F. Landry & Bucheli, sp. nov., S. isaferna J.-F. Landry & Bucheli, sp. nov., S. lisae J.-F. Landry & Bucheli, sp. nov., S. pistillata J.-F. Landry & Bucheli, sp. nov., and S. roquealbeloi J.-F. Landry & Bucheli, sp. nov. Based on a putatively unique apomorphic character of the male genitalia, they appear to form a monophyletic group restricted to the Galápagos. Their relationships to other known Scythrididae are undetermined, at best they share some generalised features with the type species and other species with similar genitalia configurations which may represent ground plan states or homoplasies. DNA barcodes showed weak differentiation among several of the species. Several species occur on more than one island, even sympatrically, and species distributions seem unrelated to the archipelago geography. Larval host plants are recorded for three species: S. isaferna on Darwiniothamnus lancifolius (Hooker f.) Harling ssp. lancifolius (Asteraceae); S. galapagensis as a leaf miner on Sesuvium portulacastrum (L.) L. (Aizoaceae) and on Portulaca sp. (Portulacaceae); and S. falcata as a leaf miner on Alternanthera halimifolia Standl. ex Pittier (Amaranthaceae).

INTRODUCTION

The family Scythrididae comprises nearly 900 described species worldwide, however, the actual diversity is estimated to be much higher based on undescribed material already in collections (Landry, 1991; Passerin d'Entrèves & Roggero, 2007; Heppner, 2010b; Bengtsson, 2014; Nupponen & Sihvonen, 2022). The majority of genera other than the type genus Scythris (16/22) apply to Old World taxa (Bengtsson, 2014), but contrastingly the vast majority of species globally are placed in Scythris.

Scythridid species diversity is higher in warmer and dryer regions. Detailed knowledge of any fauna is available only for Europe (Bengtsson, 1997) and, to a lesser extent, the Afrotropical region (Bengtsson, 2014). Many scythridids have diurnal habits and are not or are poorly attracted to lights, which is the most common method used to sample moths. In many situations it can be equally or even more productive to use sweep nets during daytime to obtain adults (Powell, 1976; Landry, 1991). Knowledge of larval host plants and life histories is also generally scant, even for the better known fauna such as that of Europe (Bengtsson, 1997; Baran, 2005). The Neotropical fauna is comparatively poor with only 35 described species (Heppner 2010b; Nupponen & Sihvonen, 2022) despite a recent revision prompted by new material from significant collecting efforts in several South American countries (Nupponen & Sihvonen, 2022). But again, this paucity is suspected to reflect a combination of inadequate collecting and dearth of taxonomic study.

The present work is the first to document species of Scythrididae from the Galápagos Islands. Ten species are represented, all of them new and described below.

Figs 1-8.

Adults of Galápagos Scythris. (1) S. isaferna ♂ holotype, specimen CNCLEP00014088 from Fernandina. (2) S. isaferna ♂paratype, specimen CNCLEP00004070 from Isabela. (3) S. isaferna ♂ paratype, specimen CNCLEP00004085 from Isabela. (4) S. isaferna ♂ paratype, specimen CNCLEP00014085 from Fernandina. (5) S. isaferna ♀ paratype, specimen CNCLEP00004162 from Isabela. (6) S. pistillata ♂ holotype, specimen CNCLEP00004166 from Pinta. (7) S. pistillata ♂ paratype, specimen CNCLEP00004165 from Pinta. (8) S. pistillata ♀ paratype, specimen CNCLEP00004167 from Pinta.

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MATERIAL AND METHODS

Specimens

This study is based on 277 specimens mostly collected by B. Landry during expeditions to the Galápagos first with S. B. Peck and others in 1989 and 1992, and later with Lazaro Roque Albelo and Patrick Schmitz starting in 2002 and in 2004-2006. They aimed at conducting a comprehensive survey of the Lepidoptera of this archipelago. The material collected by B. Landry in 1989 is deposited in the Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada (CNC) and that collected by Landry and Schmitz in 1992, 2002, 2004, 2005 and 2006 is deposited in the Muséum d'histoire naturelle in Geneva, Switzerland (MHNG). The material collected by Roque Albelo and some others is deposited at the Charles Darwin Research Station, Santa Cruz Island, Galápagos (CDRS). We examined three additional, older specimens deposited in the California Academy of Sciences, San Francisco, California, USA (CAS) and one in the Natural History Museum, London, UK (NHMUK). Additional abbreviations used are GPS for Global Positioning System, BL for B. Landry, JFL for Jean-François Landry, MIC for Microlepidoptera slides in the CNC, and SRB for Sibyl Ray Bucheli.

Morphological analysis

Morphological observations were made either on a Leica Wild-M10 or Nikon SMZ25 stereomicroscope; the former was used for observing and describing external aspect, the latter for genitalia dissection and examination. Forewing length was measured through a Nikon SMZ25 stereomicroscope at 20x magnification as viewed on a monitor with a Nikon DS-Ri2 camera and using the length measuring tool in NIS Elements software. Many specimens had their wing tips drooped which would result in length measures to be slightly underestimated. Genitalia dissections and slides were prepared following Landry (1991) with some modifications regarding neutralising and staining described in Landry (2007). For female genitalia, extraction of the inflated corpus bursae using the technique described in Landry (1991) was attempted on several specimens but inexplicably unsuccessful. A semi-inflated bursa was observed in a few dissections but deflated when the abdominal cuticle was slit open along a pleural margin, despite proceeding carefully. In preparation for slide mounting, the entire and intact female abdomen was carefully cleaned of cuticular scales and internal debris, and the five basal segments were gently flattened; the remaining portion, that is segment VI to the papillae anales, was slightly pressure-inflated by pushing the dissecting medium (30% ethanol followed by lactic acid) with a brush from the base toward the apex, taking care that the brush strokes did not go beyond the fifth segment. This was to preserve the tridimensional aspect of segments VI-VII as well as the delicate texture of the membranous wall and sinus vaginalis of sternum VIII. Without this precaution, the sternal and tergal walls would become appressed together, making it difficult to distinguish the fine structures of the sinus vaginalis, ostium bursae, and antrum/colliculum. For the permanent slide mounts in Euparal, small vinyl props of the appropriate thickness were applied around the edges of the cover slips to prevent compression and maintain tridimensionality. The slide mounts were prepared in two stages separated by one or more days; the first stage was to apply a small amount of Euparal and to position the genitalia and props, after which the preparation was covered in a dust-free plastic box for hardening; the orientation of the genitalia was checked several times in the following hour or two and adjusted if they had shifted or rolled; the second stage took place the next day or later after sufficient hardening had taken place by wetting the surface with Euparal essence, adding more Euparal, and applying the coverslip.

For male genitalia, the abdomen was slit lengthwise from base to segment VIII. The genitalia were then separated while taking care to leave the modified segment VIII attached to the abdomen. Some male genitalia were separated into their main components whereas others were left intact. The same two-stage slide mounting process was used except that the abdomen and genitalia were mounted under separate coverslips because they required different thicknesses of cover props.

Illustrations

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. Some genitalia were photographed unmounted and immersed in lactic acid (Figs 119-125, 142-143) in order to illustrate structures in orientations that cannot be easily maintained in permanent slide mounts. Slide-mounted genitalia were photographed on a Nikon Eclipse 800 microscope equipped with a Nikon DS-Fi1 camera at 40-400× magnifications. For imaging finely textured but transparent membranous structures, such as the sinus vaginalis of sternum VIII of females (Figs 136-147) differential interference contrast (DIC) illumination was used with the polariser rotated in the pale blue range which provided the best enhancement for membranes and transparent details. Nikon NIS Elements-BR© software or Zerene Stacker were both used for generating deep-focused images from multiple stacked photos. All photographs were edited with Adobe Photoshop© CC (2025). DIC images were converted to grayscale after stacking using the desaturate filter.

Line drawings (Figs 44-50, 69-77) were prepared by SRB, first drawn as pencil sketches with the aid of a drawing tube mounted on a compound microscope and magnified at 100x, digitised by scanning, and then traced over using Adobe Illustrator© to create scalable vector graphics.

Figs 9-16.

Adults of Galápagos Scythris. (9) S. galapagensis ♂ holotype, specimen CNCLEP00004160 from Santa Fé. (10) S. galapagensis ♀ paratype, specimen CNCLEP00005969 from Seymour Norte. (11) S. galapagensis ♀ paratype, specimen CNCLEP00004101 from Seymour Norte. (12) S. galapagensis ♂ paratype, specimen CNCLEP00004130 from Española. (13) S. galapagensis ♀ paratype, specimen CNCLEP00014092 from San Cristóbal. (14) S. falcata ♂ holotype, specimen CNCLEP00004047 from Santa Cruz. (15) S. falcata ♂ paratype, specimen CNCLEP00014100 from San Cristóbal. (16) S. falcata ♀ paratype, specimen CNCLEP00004048 from Santa Cruz.

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Figs 17-22.

Adults of Galápagos Scythris. (17) S. ancystra ♂ holotype, specimen CNCLEP00014091 from Santa Cruz. (18) S. ancystra ♂ paratype, specimen CNCLEP00014090 from Santa Cruz. (19) S. ancystra ♀ paratype, specimen CNCLEP00005949 from Isabela. (20) S. roquealbeloi ♂ holotype, specimen CNCLEP00005951 from Isabela. (21) S. roquealbeloi ♀ paratype, specimen CNCLEP00005953 from Isabela. (22) S. roquealbeloi ♀ paratype, specimen CNCLEP00005950 from Isabela.

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DNA sequences and analysis

DNA was extracted from single legs taken from pinned, dry specimens. Specimen data, photographs, and sequences are available in Barcoding of Life Datasystems (BOLD) (Ratnasingham & Hebert, 2013) dataset DS-SCYGAL ‘Scythrididae of the Galapagos’. 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). Sequences equal or greater than 400bp were selected for analysis, with the exclusion of those that contained either a large number of ambiguous positions (>50bp), or highly divergent positions near trim sites which likely represented noise or reading errors. A total of 245 sequences representing all the morphological species from the Galápagos were analysed.

Barcode distance analysis was performed using the Taxon-ID tree tool as implemented in BOLD in order to visualise groupings, with the BOLD aligner (Amino Acid based HMM) and Kimura 2-parameter distance model of nucleotide substitution. Distances were calculated with MEGA X (Kumar et al., 2018).

For maximum likelihood (ML) analysis, sequences were downloaded unaligned from BOLD and viewed and realigned with Geneious Prime 2025. For outgroup selection, we began by examining the ML tree in the supplementary file provided by Nupponen & Sihvonen (2022). Their tree was based on all public sequences of Scythrididae available on BOLD from North, Central, and South America, covering a wide taxon sampling; it also included sequences from the Galápagos material developed by JFL and treated in the present work, but at the time of their publication these sequences were labelled with provisional designations only. In their tree, S. limbella, S. mixaula, S. tibicina, and S. sanfranciscoensis were the described Scythrididae that clustered the closest to the Galápagos group. However, all of them were highly divergent from the Galápagos Scythris and from each other with high support values, with the Galápagos sequences forming a distant monophyletic clade. We retained only S. limbella for outgroup, which is the type species of Scythris and was also the species closest to the Galápagos cluster. The ML analysis was carried out on 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.

Figs 23-28.

Adults of Galápagos Scythris. (23) S. furculata ♂ holotype, specimen CNCLEP00005957 from Santa Cruz. (24) S. furculata ♀ paratype, specimen CNCLEP00005961 from San Cristóbal. (25) S. furculata ♀ paratype, specimen CNCLEP00005956 from Santa Cruz. (26) S. furculata ♂ paratype, specimen CNCLEP00014078 from Fernandina. (27) S. cristobalensis ♀ holotype, specimen CNCLEP00005966 from San Cristóbal. (28) S. cristobalensis ♀ paratype, specimen CNCLEP00005964 from San Cristóbal.

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Formats for descriptions and citation of specimen data

The format of the descriptions, that of the citations of label data, as well as collecting and preparation methods for specimens are given in Landry (2006). The descriptions are based on all specimens of the type series. The number of scales mentioned as forming the pecten does not represent the maximal number because these scales are easily lost. The metathorax always has small white scales laterally and light greyish brown, usually longer scales medially. The legs are described in lateral view and hence the paler scaling medially on the darker tibia and tarsus is not mentioned here. The terminology follows Landry (1991).

Accurate species identification necessitates dissection and examination of genitalia. Although the species diagnoses mention some external characters, these are secondary in all cases and should not to be relied upon for correct identification. Note also that DNA-based identification from COI sequence data by itself does not permit to distinguish several species reliably and, if available, must be used in conjunction with genitalia.

Provisional manuscript species names used by Bucheli (2005) in her Ph.D. thesis dissertation have been found on the Internet (for example,  https://ftp.funet.fi/pub/ sci/bio/life/insecta/lepidoptera/ditrysia/gelechioidea/scythrididae/scythris). These names are not available according to the rules of the International Code of Zoological Nomenclature (ICZN, 1999, and 2012 amendments), despite the fact that descriptions were provided and name-bearing types designated in Bucheli's (2005) thesis. Under Articles 9.11 and 9.12, linking in turn to Article 8, taxon names introduced in a thesis are not available because a thesis does not constitute a published work. A thesis essentially never complies with Articles 8.1 and 8.5 in that there are not enough physical printed copies AND those copies are not obtainable when first issued.

Table 1.

Unavailable Scythris species names used in Bucheli (2005) thesis dissertation and their available equivalents newly described in the present work.

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Bucheli's dissertation encompassed nine species at the time. One additional species was discovered after her work was completed. Some of the unavailable names used in Bucheli (2005) were changed in the present work, whereas others were maintained and are made available as per the present work with its publication date and the species authorship “J.-F. Landry & Bucheli” (Table 1).

RESULTS

Species recognition

Initial recognition of species was conducted by sorting specimens based on external appearance, localities and collecting dates, followed by selective dissections of genitalia from various series. This resulted in the detection of nine morphospecies well delineated by moderate to pronounced male genital differences. These morphospecies were attributed provisional designations and specimens so identified. Female genitalia, however, showed much more subtle or little obvious differences compared to males, and associating them with corresponding males was found to be highly problematic at first.

Barcode analysis was used to corroborate species delimitation based on male genitalia and to assist in associating males and females. ML analysis (Figs 150-152) showed that many clusters of the Galápagos samples had low branch support or were unsupported. Taken as a group, however, the Galápagos Scythris form a compact, strongly supported clade which is markedly distant from all other New World Scythrididae, including the type species S. limbella. The latter is 11.52% distant from the nearest of the Galápagos Scythris, which is the pair S. isafernaS. pistillata. Despite low divergences, all the morphologically defined species were recovered as shallow subclusters.

Two BINs are represented among the samples, BOLD:AAB1085 with two of the morphospecies (S. isaferna, S. pistillata), and BOLD:AAB0053 comprising the remaining eight species. These BINs are separated by 4.6% as nearest neighbours. Sequence divergences were very low to moderate within each BIN: 0.74% between S. isaferna and S. pistillata; and ranging from 0.86% between S. falcata and S. ancystra to 3.01% between S. furculata and S. lisae. The results initially were perplexing when contrasted with the pronounced morphological differences among most of the species. However, the BINs each coincide with putatively apomorphic morphological traits. In BOLD:AAB1085 S. isaferna and S. pistillata share possession of a club-shaped process on the male valva, and tergum VI of the female is reduced to a heavily sclerotized, girdle-like band extended ventrally to the spiracles. All species in BOLD:AAB0053 have the posterior part of the male valva armed with spiniform or peg-like setae, a character that may be unique in the family.

Figs 29-34.

Adults of Galápagos Scythris. (29) S. forcipata ♂ holotype, specimen CNCLEP00014079 from Fernandina. (30) S. forcipata ♂ paratype, specimen CNCLEP00004145 from Pinta. (31) S. forcipata ♀ paratype, specimen CNCLEP00004051 from Santa Cruz. (32) S. forcipata ♂ paratype, specimen CNCLEP00005976 from Isabela. (33) S. forcipata ♀ paratype, specimen CNCLEP00004157 from Pinta. (34) S. lisae ♀ holotype, specimen CNCLEP00005952 from Isabela.

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Low support for the two subclusters of S. isaferna and S. pistillata is in contrast to the striking morphological differences that distinguish these two species, especially in their male genitalia. Conversely, the trio of species S. galapagensis - S. ancystra - S. roquealbeloi are the closest morphologically but have distinct albeit close DNA barcodes. The pair S. falcata and S. ancystra were the closest in barcodes (0.88%) with poor support but are very distinct from each other in genitalia.

The cluster of S. cristobalensis is nested within S. falcata although on a long (1.35%) and strongly supported branch. Initially these specimens, all females, had been tentatively identified prior to dissection as ‘S. furculata’ due to their similar forewing pattern as well as being from the same collecting event as some true S. furculata. Dissections revealed that this divergent subcluster represented an initially undetected species, S. cristobalensis, with female genitalia strikingly different from those of all other species.

Male-female association

Correctly associating males and females is problematic in such a group where external differences between the species are variable and unreliably confusing, and the distribution of two or more species overlap. Moreover, the initial examination of a limited number of female genitalia showed small differences that did not lead to straightforward association with male counterparts. DNA barcodes through distance analysis helped to solve this problem. Despite generally low divergence, barcodes correctly associated females with their respective males, although many additional dissections were required to confirm that congruence and to assess the consistency and specific value of the small morphological differences. Dissections were performed for 53% of males and 36% of females, or an overall 45% of all specimens available for study.

Geographical distribution (Table 2)

Several species are sympatric on some islands, notably the main island of Isabela, and have been found at the same localities, even on the same date. For example, seven species (S. ancystra, S. falcata, S. forcipata, S. furculata, S. isaferna, S. lisae, S. roquealbeloi) were collected around Volcan Alcedo on Isabela, albeit some at different elevations. Three species have been found each on a single island, S. cristobalensis on San Cristóbal, S. lisae on Isabela, and S. pistillata on Pinta. Each of these three species is known from few specimens; S. cristobalensis and S. pistillata each from a single collecting event. The morphologically and genetically most distinct S. isaferna was found on Fernandina and Isabela, which are geologically the two youngest islands of the archipelago.

Table 2.

Distribution of Scythris species by island in the Galápagos archipelago. isa= S. isaferna, pis= S. pistillata, gal= S. galapagensis, anc= S. ancystra, roq= S. roquealbeloi, fal= S. falcata, for= S. forcipata, fur= S. furculata, lis= S. lisae, cri= S. cristobalensis.

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TAXONOMY

Scythris Hübner, 1825

  • Globally, the great majority of Scythrididae are assigned to the type genus which remains non-monophyletic and highly heterogeneous. Taxa are often differentiated by pronounced, even extreme morphological differences in male and female genitalia which often obscure homologies (Landry, 1991; Baran, 2005; Passerin d'Entrèves & Roggero, 2009).

  • Landry (1991) tentatively proposed a restricted definition for a small group of Nearctic species that shared putative synapomorphies with the type species, S. limbella (Fabricius) of Europe, the latter being an established introduction in North America. This was done primarily to avoid placing in Scythris the numerous Nearctic taxa unrelated to Old World groups but this hypothesis still left the genus paraphyletic or perhaps polyphyletic and all the character states were homoplasious (Baran, 2005). In addition to defining a few new genera to account for endemic Nearctic groups, Landry (1991) also used species groups within Scythris for a number of distinctive species that could not be accommodated in other genera. In absence of global phylogenetic analysis, the species-group approach has been used by several authors for other regional faunas to organise clusters of species that shared genitalia similarities without being necessarily monophyletic, yet at the same time recognising genera for some groups (Landry, 1991; Bengtsson, 1997; Passerin d'Entrèves & Roggero 2012a; Bengtsson, 2014). Recently, Nupponen & Sihvonen (2022) applied a similar hybrid approach in revising the relatively small Neotropical fauna (34 species).

  • The Scythrididae of the Galápagos are all placed here in genus Scythris, on account of the following traits:

    • • male sternum VIII expanded beneath genitalia, subtriangular in outline, convex with anterior edge emarginate, lateral portion dorsally expanded and surrounding basal part of genitalia, posterior margin with a medial projection or process (Figs 35-50);

    • • tegumen elongate with anterodorsal edge deeply V-emarginate, emargination extended to two-thirds or three quarters of length of tegumen and with a thick dorsomedial sulcus (Figs 78, 79, 82-90);

    • • uncus-socii bilobate and setose (Figs 79-90);

    • • valvae symmetrical.

  • These traits are shared by many Scythris in a broad sense and either are homoplasious within the family, or represent a grade or ground plan states. Nevertheless, they confer to the male genitalia a rather distinctive, if somewhat generalised aspect that differentiates them from the various extreme developments, reductions, or asymmetries observed in other Nearctic and Neotropical Scythrididae.

  • The Galápagos species as a group possess a unique trait – putative autapomorphy – in having the lateroposterior walls of the tegumen distally expanded under the gnathos, each side being apically joined to form a bulging hump at the apex (Figs 78-90). The uncus-socii-gnathos complex proper is small and fused dorsally; in lateral aspect it is on top of the tegumen extension (Figs 78-81). There is also a pair of internal ducts extended internally into the tegumen cavity and each with a terminal opening dorsally just distal of the uncus-socii lobes (Figs 79-81).

  • Given the many peculiar developments in abdominal segments and genitalia of the Galápagos species, we present a descriptive overview of these structures. Species-specific details are given in the formal descriptions.

  • Male abdomen

  • Terga and sterna I-VII are of typical construction as subquadrate or transversely rectangular plates which are moderately sclerotized, the sterna more so than the terga. Segment VIII differs markedly from preceding segments. Tergum VIII is much smaller than the preceding terga, either linguiform or trapezoid with a rounded posterior margin, or transverse with a sinuate posterior margin which gives it a bilobate aspect (Figs 51-59). Sternum VIII (Figs 35-50) is markedly enlarged, heavily sclerotized, subquadrate or transverse with a deep medioanterior emargination, sides recurved dorsally around the genitalia, and with heavily sclerotized, thickened margins; its posterior margin has a thick transverse sulcus where the abdominal membrane ends, and a long mediocaudal, prominent projection extended under the genitalia; this mediocaudal projection is upcurved to various degrees depending on the species (Figs 44-50) (except in S. lisae), with the apex dilated or club-shaped in some, or reduced to a small knob (S. furculata, Fig. 39); the surface of the anterior portion is smooth to the posterior transverse sulcus and devoid of scales. Pleura VIII are conically projected and bear tufts of long scales (Fig. 36) (which are easily lost in dissection).

  • Male genitalia

  • The tegumen in dorsal aspect is elongate V-shaped, or trapezoid with a deep inverted V-shaped anteromedial emargination, the median margin is a thickened sulcus; the lateral walls are posteriorly extended under the uncus-gnathos-anal cone, their apices caudally contiguous and projected dorsally to form a hump under the gnathos (Figs 78-79, 82-90); in two species (S. furculata, S. lisae) the tegumen extension is further developed ventrally into a spined or hooked process (Figs 89-90). The caudal extensions of the tegumen are less sclerotized than the main basal part and there is a membranous or unmelanized band below the uncus-socii-gnathos.

  • The uncus-socii-gnathos complex (Figs 78-90) is small, situated dorsally above the laterocaudally extended walls of the tegumen. The uncus-socii (Fig. 78) is a pair of small, setose lobes projecting caudally from the dorsocaudal edge, with long, stiff, closely set setae forming a dense dorsal tuft. Landry (1991) tentatively interpreted as socii paired, setose lobes situated in a similar position at the dorsal apex of the tegumen, but it remained uncertain or unclear whether these lobes, variously developed in many species, represent the uncus (tergum X) (see also Kristensen, 2003, p. 107). Given this uncertainty, we here refer to the setose lobes as “uncus-socii”. The lateral/ proximal arms of the gnathos form a sort of small medial ridge on each side posterior to the uncus-socii lobes, ending in a short, beak-like distal apex which protrudes just above and anterior to the tegumen lateral extensions (Figs 78-80, 82-90); the whole structure appears fused or syncleritous with the caudolateral wall of the tegumen extensions and its boundary is the unmelanized dorsal band of the tegumen extensions. There is a pair of internal ducts that are extended anteriorly into the tegumen cavity and open laterodorsally between the uncus-socii and the gnathos base; the internal terminal portion of these ducts is sclerotized and their openings are very small (shown enlarged in Figs 79-81 but also clearly visible in Figs 82-90). These ducts remind of the paired glandular openings present in some Nearctic Scythrididae, notably Rhamphura Landry (see Landry, 1991, figs 97-98, 226-227, 229), although in the latter the larger openings lead internally into a pair of blind sacs inside the tegumen. Whatever anatomical structures are connected to these ducts in the present group of species have not been observed and were likely destroyed by the KOH maceration process. The anal cone opens posterior to the uncus-socii lobes, at the base or middle of the gnathos (Fig. 80).

  • The vinculum is a transverse, subquadrate, or subconical plate with a deep anterior emargination and a thickened medial sulcus (Figs 60-68). It is articulated with the ventral pedunculi of the tegumen, the base of the valvae and the base of the phallus. In S. isaferna and S. pistillata, the vinculum is more extensive and forms a large, keel-like process projected below the valvae (Figs 60-61). The valvae are symmetrical, articulated, although with limited movability, the inner angles are extended upward to the articulation with the phallus base. In S. isaferna and S. pistillata, the basal half is wide and the distal half wedge-like or blade-like with a ventral, club-shaped projection with a setose apex (Figs 91, 92). In the other species, there is a zone of dense, short, peg-like or spiniform setae on the inner surface of the distal or apical half, in addition to sparse, normal setae scattered over the distal half (Figs 93-99).

  • The phallus is asymmetrically and variously bent, curved from right to left, articulated on the inner base of the valvae, its base has small lateral, winglet-like or lobe-like extensions articulated to the base of valvae (Figs 69-77, 100-108). In most species except S. lisae, the apex of the phallus is hooked, the subapex is humped, and the phallotreme is preapical and oriented dorsally (Figs 100-107). The bulbus ejaculatorius is small, ellipsoid or hemi-ellipsoid.

  • Female abdomen

  • Terga and sterna I-V are of typical construction as subquadrate or transversely rectangular plates which are moderately sclerotized, the sterna more sclerotized than the terga.

  • Segments VI-VII differ markedly from preceding segments (Figs 109-118).

  • Both tergal and sternal plates of segment VI are more heavily sclerotized and modified than preceding segments in all species, with tergum VI extended down laterally to wrap around the pleural region, the sclerotization surrounding the spiracles, which are produced into prominent lobes in one species (isaferna). The integument at the junction of tergum VI and sternum VI is folded under each side of the sternum, forming a pair of shallow pockets which are lined with tufts of firmly set scales. The scales of these tufts are resistant to the brushing/rubbing action while cleaning the abdomen and often can be removed only by pulling them with tweezers. When left, they obscure the sclerite boundaries. Tergum VI is narrowly constricted medially, almost divided into two separate plates in several species. Sternum VI is narrower than tergum VI.

  • Tergum VII and sternum VII are also heavily sclerotized like the plates of segment VI. Tergum VII is more or less subquadrate or subrectangular, without major development or modification. Sternum VII is elongate-rectangular or subconical, with a dense vestiture of scales except for a posteromedian zone which is devoid of scales and smooth (best visible in Figs 136-146). That smooth patch varies in extent among the species from semi-circular and extended to nearly the middle of the plate (S. galapagensis, S. ancystra), to a narrow, transverse band (S. furculata, S. cristobalensis), or reduced to a narrow line (S. lisae).

  • All species except S. cristobalensis possess a flap-like lobe protruding from the middle of the posterior margin of sternum VII, which is flat and recurved posteroventrally (Figs 126-133, 135-145, 147). In one species (S. furculata), the lobe (or the same area) is developed into a heavily sclerotized, concave, vertically oriented

  • plate which is nearly as wide as sternum VIII (Figs 130, 145). In S. cristobalensis, the lobe is reduced to a narrow band and indistinct (Figs 134, 146).

  • A posteromedial lobe of sternum VII is present in several Nearctic Scythrididae, for example several species of Arotrura Walsingham as well as Scythris trivinctella (Zeller) and S. ypsilon Braun (see Landry, 1991, figs 323-329, 343, 346), but in these groups the lobe is membranous.

  • Female genitalia

  • Tergum VIII has a horseshoe-shaped or U-shaped sclerotization (marginosclerite?) around the anterior and lateral margins, as is typical of many Scythrididae (Landry, 1991; Baran, 2005; Bengtsson, 2014). The posterior apices of the marginosclerite are variously widened (e.g., S. ancystra, S. roquealbeloi) and the posterior edge of tergum VIII is lined with fine, sparse setae.

  • The surface of sternum VIII is mostly membranous in all Galápagos species except S. lisae and S. cristobalensis, and nearly as long as sternum VII (Figs 126-133, 137-145). The sides have a swollen surface with oblique wrinkles and folds which are directed posteriorly, the middle of the segment is concave, forming a membranous longitudinal trough (sinus vaginalis) from the ostium bursae to the posterior margin of the segment. The posterior area is covered with a zone of microtrichia, a narrow strip of which is extended medially into the sinus vaginalis in some species (Figs 136, 137, 139-141, 145). In S. cristobalensis, sternum VIII has crescentic sclerotizations on each side of the sinus vaginalis (Fig. 146), whereas in S. lisae the entire ventral surface of sternum VIII forms a sclerotized plate with oblique wrinkles (Fig. 147).

  • The ostium bursae is located at the anterior end of sternum VIII immediately behind segment VII, at the anteriormost part of the sinus vaginalis. In S. furculata, the ostium bursae is surrounded anteriorly by a small, semicircular sclerotization (Fig. 145). Because sternum VIII is mostly membranous, it seems improper here to apply the term ‘sterigma’, a collective term for sclerotizations in front of, and behind the ostium bursae (Kristensen, 2003: 118) which has been widely employed in other Scythrididae (e.g. Landry, 1991).

  • The most posterior section of the ductus bursae forms a short colliculum which is barely widened and has walls with densely packed, darkly melanized cuticular folds (Figs 136-141). The length of the colliculum varies from about one-third the length of sternum VIII to a very short, stubby cone (antrum). The term colliculum is here applied as described by Kristensen (2003: 118) to a discrete sclerotized section of the ductus bursae just anterior of the antrum whereas the antrum is the most posterior, often widened part of the ductus. The boundary between these two zones is not always distinct, nor are two different zones always distinguishable. The ductus bursae is thin, membranous, flat, more or less ribbon-like, its inception point situated in distal third of the bursa copulatrix (preserved well in S. isaferna slide SRB 214). The corpus bursae is ovoid, single, delicately membranous, without signum, a state shared with all other Scythrididae.

  • The anterior apophyses are inserted in the anterior third or half of the lateral sclerotization and extended to the middle of segment VII, with a slight jog or band just beyond their insertion point. The extended ovipositor has a single section (segment IX), as is typical of other Scythrididae, about 2x as long as sternum VIII in all species. The papillae anales are elongate, two-thirds the length of segment X, and are longitudinally finely wrinkled. The membrane of segment IX is covered with extremely fine microspinules arranged in transverse rows. At low magnification with transmitted illumination this membrane appears to have a finely granular texture; under high magnification with differential interference contrast illumination, this microsculpturing appears as tiny ridges each terminating in a small point (Fig. 149).

  • Species arrangement

  • The species are arranged as follows, reflecting similarities in characters of male and female genitalia:

  • Scythris isaferna J.-F. Landry & Bucheli, sp. nov.

  • Scythris pistillata J.-F. Landry & Bucheli, sp. nov.

  • Scythris galapagensis J.-F. Landry & Bucheli, sp. nov.

  • Scythris ancystra J.-F. Landry & Bucheli, sp. nov.

  • Scythris roquealbeloi J.-F. Landry & Bucheli, sp. nov.

  • Scythris falcata J.-F. Landry & Bucheli, sp. nov.

  • Scythris forcipata J.-F. Landry & Bucheli, sp. nov.

  • Scythris furculata J.-F. Landry & Bucheli, sp. nov.

  • Scythris lisae J.-F. Landry & Bucheli, sp. nov.

  • Scythris cristobalensis J.-F. Landry & B. Landry, sp. nov.

  • The pair S. isaferna and S. pistillata have the male valva with a ventral club-shaped process and the female tergum VI reduced to a thickly sclerotized, girdle-like band which is extended and curved ventrally to join the spiracles. They lack spiniform or peg-like setae.

  • The species S. galapagensis, S. ancystra, S. roquealbeloi, S. falcata, S. forcipata, S. furculata, S. lisae have the posterior (distal) part of the male valva armed with spiniform or peg-like setae. Possession of such modified setae is unique among known New World Scythrididae, and possibly unique globally, thus likely autapomorphic. The species S. galapagensis, S. ancystra, S. roquealbeloi, S. falcata, S. forcipata have the distal (pre-apical) part of the phallus with a prominent, lobiform dorsal crest and the phallus markedly arched.

  • The trio S. galapagensis, S. ancystra, and S. roquealbeloi have the dorsal margin of the male valva crested and the apical portion wedge-shaped.

  • The pair S. falcata and S. forcipata possess a recurved spiniform process on the mediodorsal edge of the male valva and the apical portion is paddle-like with a very dense cover of peg-like setae.

  • Scythris furculata is unique in having a massively bifurcate phallus and heavy lateroposterior extensions of tegumen with a terminal upcurved hook.

  • Scythris lisae is unique in having the apex of the lateroposterior extensions of tegumen asymmetrical and heavily spined and the ostium-bearing sternum VIII of the female entirely sclerotized.

  • Scythris cristobalensis is known from females only and is uniquely distinct in having the ostium bursae surrounded by a semicircular sclerotization of sternum VIII and the medioposterior projection of sternum VII reduced to a small, indistinct sliver.

  • Figs 35-43.

    Male sternum VIII of Galápagos Scythris, ventral aspect; posterior apex oriented to the right. (35) S. isaferna, slide MIC 4970. (36) S. pistillata, slide JFL 1812. (37) S. galapagensis, slide SRB 218. (38) S. ancystra, slide JFL 1625. (39) S. furculata, slide SRB 221. (40) S. roquealbeloi, slide JFL 1630. (41) S. lisae, slide JFL 1626. (42) S. falcata; a) slide MIC 8712; b) apex only, slide BL 1156. (43) S. forcipata, slide JFL 1634.

    img-z10-1_147.jpg

    Figs 44-50.

    Male sternum VIII of Galápagos Scythris, lateral aspect; posterior apex oriented to the right. (44) S. isaferna, from dissection MIC 4971. (45) S. pistillata, from dissection SRB 227. (46) S. galapagensis, from dissection SRB 200 (holotype). (47) S. ancystra, from dissection SRB 219. (48) S. falcata, from dissection JFL 1631. (49) S. roquealbeloi, from dissection JFL 1627 (holotype). (50) S. forcipata, from dissection JFL 1634.

    img-z12-1_147.jpg

    Figs 51-59.

    Male tergum VIII of Galápagos Scythris, dorsal aspect; posterior apex oriented to the right. (51) S. isaferna, slide MIC 4968. (52) S. pistillata, slide JFL 1812. (53) S. galapagensis, slide SRB 209. (54) S. ancystra, slide JFL 1625. (55) S. roquealbeloi, slide JFL 1624. (56) S. falcata, slide MIC 8712. (57) S. forcipata, slide JFL 1634. (58) S. furculata, slide SRB 220. (59) S. lisae, slide JFL 1626.

    img-z13-1_147.jpg

    Figs 60-64.

    Male genitalia of Galápagos Scythris, lateral aspect. (60) S. isaferna, slide MIC 4970. (61) S. pistillata, slide JFL 1812. (62) S. galapagensis, slide JFL 742. (63) S. ancystra, slide MIC 4966. (64) S. roquealbeloi, slide JFL 1813.

    img-z14-1_147.jpg

    KEYS TO SPECIES OF GALAPAGOS SCYTHRIS BASED ON GENITALIA

    Males

    Note: the male of S. cristobalensis is unknown.

    1A Lower margin of valva with a pedunculate process extended ventrally with a club-like, setose apex; all setae in distal half of valva fine, hair-like (Figs 91,92) 2

    1B Lower margin of valva variously sinuate but without ventral process; distal portion of valva with densely set, spiniform or peg-like setae (Figs 93-99) 3

    2A Explanate distal half of valva horizontally transverse, wedge-like, its plane perpendicular to the vertical basal half; ventral pedunculate process caudally curved and with its apex much lower than the ventral margin of valva (Fig. 91) S. isaferna

    2B Explanate distal half of valva in the same vertical plane as the basal half; ventral pedunculate process straight with its apex only slightly below ventral margin of valva, and followed distally by a digitiform process (Fig. 92) S. pistillata

    3A Upper margin of valva with a curved, spine-like process at about two-thirds (Figs 96,97) . 4

    3B Upper margin of valva variously sinuate, but without spine-like process (Figs 93-95, 98, 99) 5

    4A Apex of spine-like process of valva sharp, upper margin humped (Fig. 96); uncus-socii low (lateral view), sloping gently to the same level as the gnathos; posterior projection of sternum VIII apically subtruncate or notched (Fig. 42); phallus evenly arched, with an angulate crest (Fig. 104) S. falcata

    4B Apex of spine-like process of valva dull, upper margin gently curved (Fig. 97); uncus-socii markedly protruded above the gnathos, which is also protruded; posterior projection of sternum VIII apically rounded, tongue-like (Fig. 43); phallus sharply bent, with a rounded crest (Fig. 106) S. forcipata

    5A Posterior projection of sternum VIII as long as or longer than basal plate, narrow, stem-like and distally narrowed before wider apex (Figs 37, 38, 40) 6

    5B Posterior projection of sternum VIII shorter than basal plate or absent (Figs 39, 41) 8

    6A Upper margin of valva evenly arched/rounded with a sharp sinuation just before apex (Fig. 95); phallus with a broad dorsal crest and a tiny, short, double-hooked apex (Fig. 105) S. roquealbeloi

    6B Upper margin of valva humped with only a slight sinuation before apex (Figs 93, 94); phallus crest with a constricted base and slender apex about as long as the crest (Figs 102, 103) 7

    7A Lower margin of valva evenly curved in basal half (Fig. 93); phallus crest recurved, with a basal hump (Fig. 102) S. galapagensis

    7B Lower margin sinuate in basal half (Fig. 94); phallus crest straight, bulb-like, with a concave base (Fig. 103) S. ancystra

    8A Distal half of valva markedly narrowed, digitiform (Fig. 99); phallus short, stout, beak-like (Fig. 108); posterior projection of sternum VIII conical, about as long as basal plate (Fig. 41) S. lisae

    8B Distal half of valva dilated, spatulate or paddle-like (Fig. 98); phallus with apex bifurcate (Fig. 107); posterior projection of sternum VIII absent (Fig. 39) S. furculata

    Females

    1A Tergum VI reduced to narrow, markedly thickened bar (hind margin) which is lateroventrally extended and curved, girdling around the base of abdominal segment VII (Figs 119, 120, 121, 123, 125) 2

    1B Tergum VI a transverse plate without thickening, albeit sides lateroventrally bent, plate may be narrowed or nearly divided in middle 3

    2A Margin of tergum VI extended all the way around the spiracles and ventrolaterally fused to sternum VI, forming a complete girdle around the base of segment VII (Figs 119, 120); spiracles prominently protruded from pleural wall (Fig. 122); sternum VII elongate-rectangular with straight posterior margin (Fig. 122) S. isaferna

    2B Margin of tergum VI tapering to spiracles in pleural wall, its apices separated from, albeit almost contiguous to, sternum VI (Figs 121, 125); spiracles not significantly protruded from pleural wall (Fig. 124); sternum VII elongate-linguiform with sinuate posterior margin (Fig. 124) S. pistillata

    3A Sternum VIII mostly or entirely sclerotized, either over entire surface, or if in part then sclerotization forming half-rings on each side around and posterior of ostium bursae (Figs 146, 147) 4

    3B Sternum VIII membranous, unmelanized, with wrinkled surface (Figs 136-141, 144), or at most with very small area of sclerotization anterior of ostium bursae (Fig. 145) 5

    4A Sternum VIII sclerotization forming half-rings around ostium bursae and interrupted medioposteriorly; posterior margin of sternum VII slightly sinuate but not emarginate and separated from sternum VIII, its medioposterior lobe reduced, indistinct; antrum with very dense, darkly melanized cuticular folds posterior to ostium (Fig. 146) S. cristobalensis

    4B Sternum VIII forming a complete subrectangular plate around ostium bursae's basal two-thirds with transverse wrinkles/creases laterally elongate-subquadrate; posterior margin of sternum VII widely excavated and encircling base of sternum VIII, its medioposterior lobe prominent, axe-like; antrum darkly melanized and extended into long colliculum (Fig. 147) S. lisae

    5A Sternum VII subcordiform in outline, anterolateral angles with short extensions recessed into pockets around spiracles, posterior margin broadly concave, thickly rimmed, medioposterior lobe a thickly sclerotized, subquadrate plate with concave surface and set under sternum VII margin (Fig. 145) S. furculata

    5B Sternum VII subconical or subtrapezoid in outline, anterolateral angles rounded, without extension (Figs 136-141, 144) 6

    6A Smooth medioposterior zone of sternum VII extended anteriorly to half or nearly half of sternal plate length (Figs 140, 141) 7

    6B Smooth medioposterior zone of sternum VII one-fifth or less of sternal plate length (Figs 128, 129, 133, 138, 139, 144) 8

    7A Elevated ridge of sinus vaginalis conical; section of colliculum with dense cuticular folds about as long or slightly shorter than smooth sinus ridge (Fig. 140); anterior emargination of sternum VI less than half length of plate (Fig. 113) S. galapagensis

    7B Elevated ridge of sinus vaginalis subrectangular; section of colliculum with dense cuticular folds longer than smooth sinus ridge (Fig. 141); anterior emargination of sternum VI deeper than half length of plate (Fig. 114) S. ancystra

    8A Sternum VI transversely semicircular, anterior margin straight (Figs 112, 129) S. forcipata

    8B Sternum VI trapezoid with sinuate/excavated anterior margin (Figs 111, 115, 128) 9

    9A Antrum a short, transverse, stubby cone; posteromedial lobe of sternum VII a thin, transverse band, almost indistinct (Fig. 144) S. roquealbeloi

    9B Antrum a narrow, barely sclerotized curved rim; posteromedial lobe of sternum VII wide, subrectangular, promiment (Fig. 138) S. falcata

    Figs 65-68.

    Male genitalia of Galápagos Scythris, lateral aspect. (65) S. falcata, slide MIC 8711. (66) S. forcipata, slide JFL 1636. (67) S. furculata, slide SRB 207. (68) S. lisae, slide JFL 1824.

    img-z17-1_147.jpg

    Figs 69-77.

    Male genitalia of Galápagos Scythris, caudal aspect. (69) S. isaferna, from dissections MIC 4968 and MIC 4971. (70) S. pistillata, from dissection SRB 227. (71) S. lisae, from dissection JFL 1626. (72) S. galapagensis, from dissection SRB 200 (holotype). (73) S. ancystra, from dissection SRB 219. (74) S. roquealbeloi, from dissections JFL 1627 (holotype) and JFL 1629. (75) S. falcata, from dissection JFL 1631. (76) S. forcipata, from dissection JFL 1634. (77) S. furculata, from dissection MIC 4965.

    img-z18-1_147.jpg

    Scythris isaferna J.-F. Landry & Bucheli, sp. nov.
    Figs 1-5, 35, 44, 51, 60, 69, 82, 91, 100, 109, 119, 120, 122, 123, 126, 136, 149

  • Material examined

  • Holotype: ♂. 1- “ECU[ADOR], Galápagos, Fernandina | SW side, GPS: 352 m elev[ation]. | S 00° 20.503', W 091° 36.969' | u[ltra]v[iolet]l[ight], 10.ii.2005 | B. Landry & P. Schmitz”. 2- “Database # | CNCLEP | 00014088”. 3- “Barcodes of Life Project | Leg removed | DNA extracted”. 4- “genitalia slide ♂ | JFL 1820”. 5-HOLOTYPE ♂ | Scythris | isaferna | Landry & Bucheli”. Deposited in MHNG.

  • Paratypes: 9 ♂, 13 ♀ from the Galápagos Islands. – Isabela: 1 ♀ (dissected, slide SRB 214), 1 km W Puerto Villamil, 3.iii.1989, M[ercury]V[apour]L[ight] (B. Landry) (CNCLEP00004060, BOLD DNA # jflandry1387); 1 ♂ (dissected, MIC 1999), 8.5 km N P[uer]to Villamil, 11.iii.1989, MVL (B. Landry) (CNCLEP00004070, BOLD DNA # jflandry1324); 1 ♂ (dissected, slide MIC 4971), same data except CNCLEP00004071); 1 ♂, same data except CNCLEP00004072, BOLD DNA # jflandry1325; 1 ♂, same data except CNCLEP00004073, BOLD DNA # jflandry1326; 1 ♂ (dissected, slide MIC 4968), same data except CNCLEP00004079; 1 ♀ (dissected, slide MIC 4961), same data except CNCLEP00004076, BOLD DNA # jflandry1327); 1 ♀ (dissected, slide MIC 8713), same data except CNCLEP00004078, BOLD DNA # jflandry1329); 1 ♂ (dissected, slide MIC 4970), Sierra Negra rim, NE side, arid zone, 15.iii.1989, swept [with net] from flowers and leaves of Darwiniothamnus sp. (B. Landry) (CNCLEP00004085, BOLD DNA # jflandry1372); 1 ♂, same data except CNCLEP00004086, BOLD DNA # jflandry1373); 1 ♀ (dissected, slide MIC 8762, same data except CNCLEP00004081, BOLD DNA # jflandry1374); 1 ♀ (dissected, slide MIC 8716, same data except CNCLEP00004082, BOLD DNA # jflandry1375); 1♀, same data except CNCLEP00004083, BOLD DNA # jflandry1376); 1 ♀ (dissected, slide SRB 213), same data except CNCLEP00004084, BOLD DNA # jflandry1377); 1 ♀, ± 15 km N Pto Villamil, 25.v.1992, MVL (B. Landry) (CNCLEP00004162, BOLD DNA # jflandry1330); 1 ♀, Volcán Darwin, 2.iii.2000, Malayse [sic] trap (L. Roque, LR # 2000-06) (CNCLEP00009425); 1 ♀, Alcedo lado [side] NE, 200 m, camp arida alta, 14.iv.2002, u[ltra]v[iolet]l[ight] (B. Landry, L. Roque) (CNCLEP00005947, BOLD DNA # jflandry1413). – Fernandina: 1 ♂, N side, 300 m, S 00° 20.541' W 091° 36.815', 12.i.2002, UVL (L. Roque, C. Causton) (CNCLEP00009426); 1 ♀, same data except CNCLEP00009427; 1 ♀, Cabo Douglas, S 00° 18.251' W 091° 39.047', 11.i.2002, Barrenador de flores Darwiniothamnus lancifolius (C. Causton, L. Roque) (CNCLEP00009467); 1 ♂, same data as holotype except CNCLEP00014085; 1 ♀, same data except CNCLEP00014087. Deposited in CDRS, CNC, and MHNG.

  • Etymology: The new name refers to the occurrence of the species on the islands of Isabela and Fernandina and is treated as a noun in apposition.

  • Diagnosis: The male genitalia are characterized by markedly arched tegumen (lateral aspect); the valva with the distal half wedge-shaped, horizontally transverse, and perpendicular to the basal half, and without pegged or spiniform setae; the ventral margin has a curved pedunculate process with a club-shaped apex; the phallus is downcurved with a slight preapical hump dorsally and a small upcurved apical hook; sternum VIII has a markedly upcurved medioposterior projection with an apical dilatatation and an anterior emargination which is broadly V-shaped. Scythris pistillata is similar in lacking pegged setae on the valva and having a pedunculate process with a club-shaped apex but differs in the broader distal half of valva which is vertical and has a granular surface microsculpture, the shorter pedunculate process followed by a digitiform ventral lobe, the straighter phallus, and thicker medioposterior projection of sternum VIII with a double apical dilatation.

  • In female genitalia, tergum VI is reduced to a narrow, markedly thickened bar which is extended all the way around the spiracles and ventrolaterally fused to sternum VI, forming a complete girdle around the base of segment VII; this bar is arched and laterally angular; the spiracles of segment VI are prominently protruded from a sclerotization of the pleural wall; tergum VII has a thickly sclerotized, arched anterior margin; sternum VII is elongate-rectangular with a straight posterior margin. The sterigma is a short, sclerotized band with a shallow medial notch. Scythris pistillata also has tergum VI reduced to a narrow bar but it is evenly arched, and its pleural extensions are not fused to sternum VI, only contiguous with it, and the spiracles are on a membranous pleural wall and not protruded; its sternum VII is linguiform with a notched posterior margin and the anterior margin of tergum VII is membranous.

  • In females, the abdomen ventrally has a striped pattern of greyish brown at base of tergites and white distally to cream distally on the two distal sternites, with the distal sternite sometimes all greyish brown; males as well as females of the other Galápagos species do not have the abdomen striped ventrally.

  • Description: Male (n=10) (Figs 1-4). Head: occiput mostly blackish brown with faint purple shine, with paler brown to white scales laterally along antennal bases; lateral tufts of scales projecting dorsomedially from behind antennae dark brown medially and paler brown to white laterally; frontoclypeus blackish brown with paler brown to white scales laterally and toward ventral margin, sometimes mostly white. Antenna blackish brown with white ventrally on scape and pedicel, with white patch at base above pecten, with row of pale greyish brown scales along flagellum ventrally; pecten with up to 18 brown scales paler at their bases. Labial palpus ivory white with greyish brown on 1st palpomere, mostly dark greyish brown on 2nd with few white scales at base, dorsally and apex, greyish brown on 3rd with white at base and dorsally. Haustellum scales white at base, greyish brown toward end of scaled section. Thorax blackish brown, with few scattered paler brown to white scales or with more abundant white scaling. Wings as shown (Figs 1-4); forewing upperside varying in amount of white scaling and contrast of pattern, frequently with black spot surrounded with white above tornus; forewing underside dark brown, with greyish brown patch from costa as distal section of retinaculum; hindwing underside pale greyish brown at base, darker brown beyond. Forewing length: 4.4-6.0 mm, mean 5.3 ± 0.5, n=10 (holotype: 5.4 mm). Foreleg coxa dark greyish brown with white at apex and scattered elsewhere; rest of leg dark greyish brown, with white at base and scattered elsewhere on femur sometimes, at apex of tibia, base and apex of tarsomere I, and sometimes apex of tarsomere II. Midleg coxa satiny white and pale greyish brown; femur dark greyish brown with white scales except toward distal 3rd; tibia dark greyish brown with white medially and apically; tarsomeres dark greyish brown with white at base and apex of 1st. Hindleg coxa satiny white and pale greyish brown; femur satiny white with dark greyish brown apically; tibia mostly white, with brown patches increasing in size at base, submedially, and subapically, with crest of hair-like scales ivory white; tarsomeres dark greyish brown, with white at base and apex of 1st, and pale greyish brown at apex of 2nd. Abdomen dorsally mostly greyish brown, warmer dark brown on ultimate scaled tergite; ventrally all ivory white.

  • Male abdomen and genitalia (5 preparations) (Figs 35, 44, 51, 60, 69, 82, 91, 100). Tergum VIII elongate-linguiform, anterior margin concave, posterior margin with straight sides and rounded corners. Sternum VIII with very wide base, anterior margin broadly emarginate, strengthening sulci very thick and extended along medioposterior projection. Sides and medioposterior projection markedly upcurved, apex dilated, club-shaped with tuft of firmly set scales on a subapical hump, posterolateral angles prominent.

  • Tegumen bent at right angle at midlength, with basal half vertical and distal half horizontal (at 90 degrees from basal half), ventrocaudal extension with blunt, straight, weakly sclerotized apex. Uncus-socii lobes mediobasally contiguous, dorsally protruded, stiff setae projected dorsally. Paired ducts very short, indistinct. Gnathos with very small, hook-like, downcurved apex. Vinculum large, extended to middle of valvae, base with heavy, V-shaped dorsal extensions articulated to tegumen and along base of valvae. Valva with smooth basal half, distal half wedge-shaped or securiform, projected transversely (in horizontal plane at right angle with basal half), with long, ventral, curved, stem-like projection with club-shaped, setose apex; clusters of fine setae on ventral margin near base of stem and of distal wedge. Phallus downcurved, with subterminal hump on right side and a small, recurved, sharp hook at apex.

  • Female (n=13) (Fig. 5). Head, thorax, and wings dorsally as in male; forewing underside with patch of scales projecting from costa as distal part of retinaculum often white, sometimes pale greyish brown, with paler scaling along and below costa usually continuing until middle of wing; forewing length: 4.4-5.4 mm, mean 4.8 ± 0.3, n=12; hindwing underside with costal 1/3 dark brown almost from base until apex, with dorsal 2/3 pale cream colored until before apex, or more homogeneously coloured different shades of brown; frenulum usually with 3 bristles, sometimes 2 (e.g., CNCLEP00004078, CNCLEP00004081; CNCLEP00004083 has 2 on the left hindwing and 3 on the right hindwing). Abdomen dorsally mostly greyish brown, with satiny white on 1st tergite, darker brown sometimes on distal two tergites; ventrally with striped pattern of greyish brown at base of tergites and white distally to cream distally on distal two sternites, with distal sternite sometimes all greyish brown.

  • Female abdomen and genitalia (7 preparations) (Figs 109, 119, 120, 122, 123, 126, 136, 149). Abdominal terga II-V with thickened anterior margins. Tergum VI a narrow, thickly sclerotized band forming a girdle-like ring which is laterally downcurved along the pleura, its distal ends tightly juxtaposed to the lateral margins of sternum VI and lobate around the spiracles which jut out posterolaterally; these spiracle lobes are densely covered with scales that are difficult to remove during the cleaning process. Sternum VI transverse, linguiform with a thickly sclerotized, broadly incurved (concave) anterior margin, posterior margin more or less straight with rounded lateral angles. Tergum VII elongate-subcylindrical with very thick, rim-like, curved anterior margin. Sternum VII elongate-subrectangular about 2x longer than sternum VI, with thick, concave anterior margin which is recessed under sternum VI, the latter forming like an overlapping lip; surface densely covered with scales except for small, semicircular bare patch posteromedially; posterior margin straight, medioposterior lobe transverse. Tergum VIII membranous, half the length of tergum VII, surface with very fine longitudinal wrinkles, posterior margin lined with row of fine setae; lateral and anterior margins sclerotized, together forming a horseshoe-like band or arc. Anterior apophyses extended from anterolateral corners of tergum VIII and about 1.5x length of tergal plate, extended to middle of tergum VII, marginosclerite lateroposteriorly widened. Sternum VIII membranous, with only a hint of sclerotization or melanization, surface obliquely wrinkled, medially with longitudinal trough-like depression (sinus vaginalis) with finely spinulate surface in posterior half. Ostium bursae situated at anterior end of sinus vaginalis and surrounded by a small, short, stubby, darkly melanized colliculum.

  • Biology: One moth of this species was reared from a caterpillar boring in flowers of Darwiniothamnus lancifolius near the shore on Cape Douglas. Six specimens were swept with a net from flowers of a Darwiniothamnus species on the rim of Sierra Negra volcano. Both sites are on Isabela Island. Hence, the species can be found at a range of elevations, providing that host plants of the genus Darwiniothamnus are present. Darwiniothamnus lancifolius (Hooker f.) Harling ssp. lancifolius (Asteraceae) is endemic to Isabela and Fernandina islands and is classified as vulnerable to extinction (Galápagos Species Database, 2024).

  • Distribution: The species is presumed to be endemic to the Galápagos and has been found on the islands of Isabela and Fernandina.

  • Figs 78-81.

    Uncus-socii-gnathos complex of Galápagos Scythris. Dashed lines delineate the hindgut and anal cone. (78) S. roquealbeloi, dorsal aspect, slide JFL 1629. (79) S. falcata, lateral aspect, slide MIC 8711. (80) S. falcata, lateral aspect, slide JFL 1631; extruded apex of anal cone delineated by thin solid line. (81) S. falcata, lateral aspect showing the paired glandular ducts, slide MIC 8711.

    img-z20-1_147.jpg

    Figs 82-90.

    Male uncus-socii-gnathos complex of Scythris, lateral aspect. (82) S. isaferna, slide MIC 1999. (83) S. pistillata, slide JFL 1812. (84) S. galapagensis, slide JFL 1815. (85) S. ancystra, slide JFL1833. (86) S. roquealbeloi, slide JFL 1624. (87) S. falcata, slide MIC 8711. (88) S. forcipata, slide JFL 1636. (89) S. furculata, slide MIC 4965. (90) S. lisae, slide JFL 1824.

    img-z21-1_147.jpg

    Figs 91-99.

    Male valva of Galápagos Scythris, lateral aspect of inner side. (91) S. isaferna, slide MIC 4968. (92) S. pistillata, slide JFL 1618. (93) S. galapagensis, slide JFL 1835. (94) S. ancystra, slide JFL 1833. (95) S. roquealbeloi, slide JFL 1627. (96) S. falcata, slide MIC 2001. (97) S. forcipata, slide JFL 1634. (98) S. furculata, slide MIC 4965. (99) S. lisae, slide JFL 1626.

    img-z22-1_147.jpg

    Scythris pistillata J.-F. Landry & Bucheli, sp. nov.
    Figs 6-8, 36, 45, 52, 61, 70, 83, 92, 101, 110, 121, 124, 125, 127, 137, 148

  • Material examined

  • Holotype: ♂. 1- “ECU[ADOR]., GALAPAGOS | Pinta, 16.iii.1992 | 200 m elev[ation]., M[ercury]V[apour] L[ight] | leg[it]. B. Landry”. 2- “Database # | CNCLEP | 00004166”. 3- “Barcodes of Life Project | Leg removed | DNA extracted”. 4- “genitalia slide ♂ | JFL 1812”. 5- “HOLOTYPE ♂ | Scythris | pistillata | Landry & Bucheli”. Deposited in MHNG.

  • Paratypes: 2 ♂, 3 ♀ from the Galápagos Islands. – Pinta: 1 ♂ (dissected, slide SRB 227, preparation lost), same data as holotype except CNCLEP00004165; 1 ♀ (dissected, slide JFL 1826), same data as holotype except CNCLEP00004146, BOLD DNA # jflandry1354; 1 ♂ (dissected, slide JFL 1618), 18.iii.1992, 400 m elev[ation]., M[ercury]V[apour]L[ight] (B. Landry) (CNCLEP00004168, BOLD DNA # jflandry1350); 1 ♀ (dissected, slide MIC 8718), same data except CNCLEP00004167, BOLD DNA # jflandry1351; 1 ♀ (dissected, slide JFL 1805), N 00°34.591', W 90°45.137', 421 m elev., u[ltra]v[iolet]l[ight], 17.iii.2006 (P. Schmitz, L. Roque) (CNCLEP00024704). Deposited in CNC and MHNG.

  • Etymology: The new name is derived from the Latin pistillum which is a club-shaped pounder, in reference to the club-shaped ventral process of the valva.

  • Diagnosis: In male genitalia, the valva has a broadly explanate, vertically oriented distal half in the same plane as the basal half, with an irregularly sinuate apical edge and the inner surface with very fine granular microsculpture (Fig. 148) and without peg-like setae, its ventral margin has both a club-shaped but short pedunculate process and a digitiform lobe; the phallus is slightly downcurved with the apex bearing a small, recurved tooth; sternum VIII has a thick and sharply upcurved medioposterior projection with a bilobate apex, the preapical lobe being inwardly directed, and the anterior emargination is broadly U-shaped. Both S. isaferna and S. pistillata are distinguished from the other Galápagos species by their pedunculate ventral process of the valva and the lack of spiniform or peg-like setae in the distal portion of the valva; the other species lack a pedunculate ventral process and have the terminal part of the valva with dense, spiniform or peg-like setae. In female genitalia, the sclerotized bar of tergum VI forms an evenly curved ark which is laterally curved down the pleural wall and tapers to a narrow strip subcontiguous with the sides of the sternum plate. Sternum VII is linguiform with a notched posterior margin and the anterior margin of tergum VII is membranous.

  • The extensive medial white stripe across the frontoclypeus may be diagnostic, but genitalia need to be examined for positive identification.

  • Description: Male (n=3) (Figs 6-7). Head: occiput dark greyish brown with purple shine, white laterally along antennal bases and scattered elsewhere; lateral tufts of scales projecting dorsomedially from behind antennae mostly white with some dark greyish brown scales medially; frontoclypeus striped longitudinally pale greyish brown dorsally, white medially, dark greyish brown ventrally, or with greyish brown showing in middle of white stripe. Antenna blackish brown, with paler greyish brown scape adorned with white patch at base dorsally; ventrally with satiny white on scape and pedicel and with row of cream-colored scales along flagellum; pecten made of up to 14 brown scales with their base usually paler. Labial palpus: 1st palpomere greyish brown with white along dorsal and ventral margins, 2nd greyish brown with white at base, dorsally, and apex, 3rd greyish brown with white at base and dorsally. Haustellum white at base, pale greyish brown further along scaled section. Thorax greyish brown, with more or less extensive white scaling, except on distal half of tegulae, also with darker, blackish brown laterally before apex and apically on mesothorax. Wings as shown (Figs 6, 7); forewing upperside slightly variable in contrast of pattern, i.e., with less white scaling in blackish brown areas and vice versa; forewing underside brown, with greyish brown patch from costa as distal section of retinaculum followed by white band along costa to about midlength of wing; forewing length: 4.8-5.0 mm, mean 4.9 ± 0.1, n=3 (holotype: 5.0 mm); hindwing underside pale whitish at base along costa and on dorsal 2/3 of wing until tornus, brown on costal 1/3 to apex. Foreleg coxa dark greyish brown with extensive white scaling; femur dark greyish brown with scattered white scales; rest of leg dark greyish brown, with white at 1/3 and apex of tibia, base and apex of tarsomere I, and sometimes apex of tarsomere II. Midleg coxa satiny white and pale greyish brown; femur satiny white with dark greyish brown at apex; tibia dark greyish brown with white submedially, postmedially, and apically; tarsomeres dark greyish brown with white at base and apex of 1st and apex of 2nd. Hindleg coxa pale greyish brown with satiny white along dorsal margin; femur satiny white with dark greyish brown apically; tibia mostly white, with brown patches increasing in size at base, submedially, and subapically, with crest of hair-like scales ivory white; tarsomeres dark greyish brown, with white at base and apex of 1st and apex of 2nd. Abdomen dorsally not checked before dissections; ventrally white, although not checked in detail.

  • Male abdomen and genitalia (3 preparations) (Figs 36, 45, 52, 61, 70, 83, 92, 101, 148). Generally similar/resembling S. isaferna except as follows: Tergum VIII transverse-conical with posterior margin evenly rounded, anterior margin broadly V-concave. Sternum VIII with very wide, slightly asymmetrical base, anterior margin more rounded, median posterior projection thicker, apex asymmetrically humped on left side and with another subterminal ventral process pointed anteriorly; right posterolateral angle markedly produced, much more so than left side.

  • Uncus-socii more dorsally protruded. Apex of tegumen extensions more square and more thickly sclerotized. Paired ducts short. Valva with basal half very wide, distal half also very wide but oriented vertically (in dorso-ventral plane) as basal half, outline irregular, anterodorsal surface sparsely setose, ventral margin with club-shaped projection setose, short-stemmed and with flat lobe distal of it, inner surface of distal half with very fine granular microsculpture (Fig. 148). Vinculum with distal half shorter, less projected, shallower. Phallus less downcurved, subterminal hump poorly developed and ventral, apex with a short, blunt hook.

  • Female (n=3) (Fig. 8). Head, thorax, and wings as in male; forewing length: 4.5-4.8 mm, mean 4.7 ± 0.2, n=3; frenulum with 3 bristles. Abdomen dorsally: tergite one pale greyish brown with satiny white, subsequent tergites greyish brown with paler apical margin; ventrally ivory white to cream white with some pale greyish brown distally.

  • Female abdomen and genitalia (3 preparations) (Figs 110, 121, 124, 125, 127, 137). Abdominal segment I-V of typical construction and sclerotization. Sternum V anterior margin thickened medially. Tergum VI modified as a thickly sclerotized, brace-like rim similar to tergum VI of S. isaferna, except for its dorsal section which is regularly rounded (not angulate as S. isaferna), apex ending before spiracles; pleural area around spiracle not protruded, flat. Sternum VI transverse, anterior margin thickened, medially concave, laterally sinuate, posterior margin medially concave, laterally rounded; not extended over sternum VII (unlike S. isaferna). Tergum VII elongate-subcylindrical, distally narrower, anterior margin not differentiated, anterolateral corners with short ventral downcurved extensions. Sternum VII elongate-linguiform, anteriorly narrower, with thickened, concave anterior margin which is slightly offset from rest of sclerite, posterior margin with shallow medial concavity, smooth/ bare patch very small, conical in outline, just anterior to marginal notch, medioposterior lobe suborbicular. Tergum VIII similar to that of S. isaferna but overall proportionally narrower. Sternum VIII similar in aspect to that of S. isaferna but proportionally narrower, more elongate. Ostium bursae ringed by thick, stubby, darkly melanized sclerotization with posterior notch, extended internally into very short (shorter than intersegmental membrane SVII-SVIII) conical unmelanized antrum. Ovipositor 2x length of sternum VIII. Bursa copulatrix similar to that of S. isaferna.

  • Biology: Unknown except for the moths being attracted to light and the rather restricted range of elevations of its records, between 200 and 421 m in elevation.

  • Distribution: Presumably endemic to the Galápagos, S. pistillata has been found only on the island of Pinta.

  • Figs 100-108.

    Male phallus of Galápagos Scythris, lateral aspect; arrow indicates position of phallotreme. (100) S. isaferna, slide MIC 4968. (101) S. pistillata, slide JFL 1618. (102) S. galapagensis, slide SRB 218. (103) S. ancystra, slide JFL 1625. (104) S. falcata, slide JFL 1631. (105) S. roquealbeloi, slide JFL 1624. (106) S. forcipata, slide JFL 1637. (107) S. furculata, slide JFL 1632. (108) S. lisae, slide JFL 1626.

    img-z24-1_147.jpg

    Scythris galapagensis
    J.-F. Landry & Bucheli, sp. nov.

    Figs 9-13, 37, 46, 53, 62, 72, 84, 93, 102, 113, 131, 140, 142

  • Material examined

  • Holotype: ♂. 1- “ECU[ADOR]., GALAPAGOS | Santa Fé, Tourist trail | 28.v.1992 | M[ercury]V[apour] L[ight], leg[it]. B. Landry”. 2- “Database # | CNCLEP | 00004160”. 3- “genitalia slide ♂ | SRB 200”. 4- “Barcodes of Life Project | Leg removed | DNA extracted”. 5-“genitalia slide ♂ | SRB 200”. 6- “HOLOTYPE ♂ | Scythris | galapagensis | Landry & Bucheli”. Deposited in MHNG.

  • Paratypes: 33 ♂, 38 ♀ from the Galápagos Islands. – Bainbridge: 1 ♀, Banbridge [sic], Criada en Sesuvium portulacastrum, 30.v.2001 (L. Roque) (CNCLEP00009438). – Española: 6 ♂ (2 dissected, slides SRB 204, SRB 218), Punta Suarez, M[ercury]V[apour]L[ight], 2.v.1992 (B. Landry) (CNCLEP00004125, CNCLEP00004127, C N C L E P 0 0 0 0 4 1 2 8, C N C L E P 0 0 0 0 4 1 3 0, CNCLEP00004131, CNCLEP00004134); 1 ♂, same data except CNCLEP00004126, BOLD DNA # jflandry1340; 1 ♂ (dissected, slide JFL 1619), same data except CNCLEP00004129, BOLD DNA # jflandry1341; 1 ♂, same data except CNCLEP00004132, BOLD DNA # jflandry1337; 1 ♀, same data except CNCLEP00004133, BOLD DNA # jflandry1338. – Pinta: 1 ♂, Playa Ibbetson, 13.iii.1992, MVL (B. Landry) (CNCLEP00004138, BOLD DNA # jflandry1367). – Pinzón: 2 ♀, Playa Escondida, 14 m elev., S 00°35.928' W 90°39.291', 27.iii.2006 (P. Schmitz) (CNCLEP00024706, CNCLEP00024707); 2 ♀, playa Escondida, uvl, 20.iv.2002 (B. Landry, P. Schmitz) (CNCLEP00005981, CNCLEP00005984); 1 ♀ (dissected, slide SRB 216), same data except CNCLEP00005982, BOLD DNA # jflandry1392; 1 ♀, same data except CNCLEP00005983, BOLD DNA # jflandry1393. – San Cristóbal: 1 ♂ (dissected, slide MIC 4963), 2 km SW P[uer]to Baquarizo [sic], MVL, 11.ii.1989 (B. Landry) (CNCLEP00004055); 4 ♂ (3 dissected, slides JFL 1815, JFL 1816, JFL 1841), 1 ♀(dissected, slide JFL 1814), near La Loberia, sea level, GPS: S 00° 55.277' W 089° 36.909', uvl, 24.ii.2005 (B. Landry) (CNCLEP00014092-CNCLEP00014096). – Santa Cruz: 2 ♀ (one dissected, slide MIC 8715), C[harles]D[arwin]R[research]S[tation], Arid zone, MVL, 17.i.1989 (B. Landry) (CNCLEP00004043, CNCLEP00004044); 1 ♂ (dissected, slide MIC 2003), lagoon 2 km E CDRS, reared from Portulaca sp., [collected] 18.i.1989, emerged 26.i.1989 (B. Landry) (CNCLEP00004045, BOLD DNA # jflandry1344); 1 ♀ (dissected, slide MIC 8754), CDRS, Arid zone, MVL, 19.i.1989 (B. Landry) (CNCLEP00004042); 1 ♂ (dissected, slide MIC 2002), same data except CNCLEP00004041, BOLD DNA # jflandry1343); 1 ♂ (dissected, slide MIC 4969), littoral zone, Tortuga Bay, MVL, 29.i.1989 (B. Landry) (CNCLEP00004046, BOLD DNA # jflandry1332); 1 ♀ (dissected, slide SRB 208), E[stacion].C[ientifica].C[harles].D[arwin]., Barranco, MVL, 13.ix.1999 (L. Roque) (CNCLEP00004176, BOLD DNA # jflandry1397); 1 ♂ (dissected, slide SRB 209), same data except 11.xi.1999, CNCLEP00004175; 2 ♂, 3 ♀, Tortuga Bay, UVL, 11.xii.2001 (L. Roque) (CNCLEP00009450-CNCLEP00009454); 1 ♂ (dissected, slide JFL 1800), xii.1968 (R. Perry, Tj. de Vries; B[ritish] M[useum] 1969-17; Ref. No. L. 51) (CNCLEP00005985) (NHMUK). – Santa Fé: 1 ♂, Tourist trail, MVL, 28.v.1992 (B. Landry) (CNCLEP00004159, BOLD DNA # jflandry1331). – Santiago: 1 ♀ (dissected, slide JFL 1817), Costa Sur Este, Minador Sesuvium sp., 13.ii.2002 (R. Boada) (CNCLEP00009449); 2 ♂ (one dissected, slide JFL 1633), 3 ♀, Cerro Inn, MVL, 28.iii.1992 (B. Landry) (CNCLEP00004117, CNCLEP00004119-CNCLEP00004122); 1 ♀, same data except CNCLEP00004118, BOLD DNA # jflandry1336); 1 ♂, same data except CNCLEP00004123, BOLD DNA # jflandry1335). – Seymour Norte: 2 ♀ (one dissected, slide MIC 8714), Arid Zone, MVL, 23.i.1989 (B. Landry) (CNCLEP00004052, CNCLEP00004053); 1 ♀, dissected, slide SRB 222, same data except CNCLEP00004054, BOLD DNA # jflandry1401); 1 ♀, GPS: 17 m elev., S 00°23.935' W 90°17.869', uvl, 22.iii.2004 (B. Landry, L. Roque, P. Schmitz) (CNCLEP00005969, BOLD DNA # jflandry1402); 1 ♀ (dissected, slide SRB 223), GPS: 13 m elev., S 00°24.013' W 90°17.422, uvl, 23.iii.2004 (B. Landry, L. Roque, P. Schmitz) (CNCLEP00005970, BOLD DNA # jflandry1403); 1 ♂ (dissected, slide JFL 1622), 4 ♀ (one dissected, slide SRB 224), [no precise locality], MVL, 29.iii.1992 (B. Landry) (CNCLEP00004098-CNCLEP00004100, CNCLEP00004103, CNCLEP00004104); 1 ♀, same data except CNCLEP00004097, BOLD DNA # jflandry1404); 1 ♀, same data except CNCLEP00004101, BOLD DNA # jflandry1405); 1 ♀ (dissected, slide SRB 225), same data except CNCLEP00004102, BOLD DNA # jflandry1406; 1 ♂, same data except CNCLEP00004110, BOLD DNA # jflandry1346); 1 ♀, same data except CNCLEP00004111, BOLD DNA # jflandry1347); 1 ♂ (dissected, slide JFL 1623), same data except CNCLEP00004113, BOLD DNA # jflandry1345). – South Plaza: 4 ♂ (one dissected, slide JFL 1818), 4 ♀, Plaza Sur, 18 m elev., S 00°34.980' W 90°09.990', uvl, 14.iv.2006 (P. Schmitz) (CNCLEP00024708-CNCLEP00024715).

  • Specimens excluded from the type series: – Genovesa: 3 ♂ (one dissected, slide JFL 742), Tower I[sland]., 25.iii.1935 (Templeton Crocker) (CNCLEP00004177-CNCLEP00004179). Deposited in CAS.

  • Etymology: The new name refers to the Galápagos archipelago, the area of origin of the specimens studied.

  • Diagnosis: In male genitalia, the valva has the apical margin covered with thickened, spiniform setae, the dorsal margin is humped beyond middle, the basal half of the ventral margin slightly and regularly curved; the phallus has a 90° bend in middle and a high, curved dorsal crest with a low rounded lobe at its base; sternum VIII has a narrow, tapered medioposterior projection with a narrow apex of even thickness in lateral aspect. The male genitalia of S. galapagensis resemble in overall appearance those of S. ancystra and, to a lesser extent, those of S. roquealbeloi, but differ in several details which are presented under each species. The male genitalia of S. roquealbeloi differ in the dorsal margin of the valva evenly curved with a short, sharp preapical sinuation, the basal half being broader than the distal half; the medioposterior projection of sternum VIII is straight-sided, apically truncate; the gnathos has the apex finely spinulate; the phallus has a sharp medial bend, broad crest and bifurcated apex.

  • In female genitalia, the sinus vaginalis is on a smooth area which is triangular in outline and nearly flat with the rest of the surface of sternum VIII; the section of colliculum with dense, darkly melanized cuticular folds is short or slightly shorter than the smooth sinus area; on sternum VII the posteromedial smooth patch is hemielliptic, large, extended to the middle of the sclerite and the medioposterior process is wide, about half the width of the posterior margin; the anterior emargination of sternum VI is less than half the length of the plate. In female genitalia, S. galapagensis is most similar to S. ancystra, the latter differs in having the sinus vaginalis conical in outline on an elevated ridge which protrudes from the sternal surface, and the section of colliculum with dense cuticular folds more slender, longer than the sinus ridge; the medioposterior process of sternum VII is small and much narrower than the posterior margin; the anterior emargination of sternum VI is wide and deeper than half the length of the plate.

  • Description: Male (n=34) (Figs 9, 12). Head: occiput and frontoclypeus orange brown, greyish brown, or darker brown, with concolorous or slightly paler lateral tufts of scales projecting dorsomedially from behind antennae. Antenna greyish brown to blackish brown; scape sometimes with white scales at apex; scape and pedicel ventrally white; flagellum with row of light brown scales ventrally; pecten with up to 12 whitish to light brown scales with their bases usually paler. Labial palpus: 1st palpomere white, sometimes with light brown to greyish brown, or all pale greyish brown; 2nd dark greyish brown with scattered white to pale greyish brown scales, mostly white to pale greyish brown at base (rarely to middle of segment), dorsally, and apex; 3rd mostly dark greyish brown, sometimes with scattered white to pale greyish brown scales, white to pale greyish brown at base and dorsally. Haustellum usually white, sometimes dirty white, sometimes with pale greyish brown (rarely dark greyish brown) in middle at base, rarely further along. Thorax various shades of brown, often darker brown at base of tegulae and most of mesothorax or laterally before apex and apically on mesothorax; often paler, orange brown elsewhere. Wings as shown (Figs 9, 12); forewing upperside varying in contrast of pattern and in prominence of the colours on the more contrasting specimens; forewing underside brown, darkening toward apex, with white patch of scales from costa as distal section of retinaculum more or less expressed, sometimes with white scales as well further along costa; forewing length: 3.7-4.6 mm, mean 4.2 ± 0.2, n=34 (holotype: 4.5 mm); hindwing underside with more or less contrasting brown costal half including apex and paler whitish dorsal half from base to tornus. Foreleg coxa greyish brown with white at apex, on inner side, and sometimes scattered elsewhere; femur blackish brown sometimes with light greyish brown scattered; tibia blackish brown with few white scales submedially and postmedially sometimes, and apically; tarsus: 1st tarsomere dark brown with pale greyish brown at base and white at apex; 2nd to 3rd dark brown with white at apex; 4th and 5th dark greyish brown. Midleg coxa satiny white; femur mixed satiny white and dark greyish brown, the latter more extensive toward apex; tibia blackish brown with white spots postmedially and apically; tarsus: 1st tarsomere with extensive white scaling at base and apex, and dark brown in middle; 2nd to 4th dark brown at base more extensive toward distal tarsomeres, white at apex; 5th all dark greyish brown. Hindleg coxa satiny white; femur satiny white with dark greyish brown apex; tibia white with dark to light greyish brown as base, 1/3, and subapically, with crest of hairlike scales white to light brown; tarsus (sometimes brown scaling pale greyish brown): 1st tarsomere with extensive white scaling at base and apex, and dark brown in middle; 2nd to 5th dark brown at base more extensive toward distal tarsomeres, white at apex although often not apparent on 4th and 5th. Abdomen dorsally pale greyish brown, slightly darkening toward distal segments, warmer dark brown on longer scales covering genitalia; ventrally mostly satiny white or ivory white except pale greyish brown on first two sternites and cream to light brown under genitalia. Male abdomen and genitalia (19 preparations) (Figs 37, 46, 53, 62, 72, 84, 93, 102). Tergum VIII transverse with sinuate/concave posterior margin, posterior corners rounded, anterior margin almost straight. Sternum VIII digitiform, symmetrical, base with narrow anterior emargination, medioposterior projection gradually narrowed with evenly curved sides, apex roundly dilated, almost club-shaped and with slight dorsal hump.

  • Tegumen gradually weakly arched, posterior extensions upcurved. Uncus-socii lobes almost flat, inconspicuous in lateral view. Paired ducts relatively long, heavily sclerotized. Gnathos apex with straight, tooth-like dorsoapical process and blunt ventral process, thus appearing bilobate in lateral view. Vinculum trapezoid, posteriorly barely extended under base of valvae. Valva with more or less straight basal half, distal half with dorsal hump and roundly conical apex, ventroapical margin with narrow zone of peg-like setae which are somewhat elongate. Phallus sharply bent down and left at about mid-length, distal portion with prominent ventral, recurved, crest-like process with basal hump, apical portion tapered into thin, elongate point with small terminal hook and with slight corrugation distal of crest. Female (n=38) (Figs 10, 11, 13). Head, thorax and wings as in male, except forewing underside with white patch of scales from costa as distal section of retinaculum more distinctly expressed than in male, and with more white scales further on along the costa; frenulum with 3 bristles. Forewing length: 3.6-5.0 mm, mean 4.3 ± 0.3, n=38. Abdomen dorsally as in male except apical scales usually paler, whitish to light brown; ventrally usually ivory white, sometimes satiny white toward base, sometimes all pale greyish brown.

  • Female abdomen and genitalia (17 preparations) (Figs 113, 131, 140, 142). Abdominal segments I-V of typical construction, moderately sclerotized. Tergum VI strongly sclerotized, a narrow, transverse plate medially narrowed to the point of being almost subdivided into two lateral sections; sides bent ventrally around pleural area and ending in a fold under the lateral margins of sternum VI, with dense, tufted vestiture of firmly set scales. Sternum VI transversely subtrapezoid, anterior margin deeply concave. Tergum VII subquadrate-subcylindrical, posterior margin lined with sparse, fine setae. Sternum VII conical-truncate, about 1.5x length of sternum VI, posteromedial smooth patch hemielliptic, large, extended to middle of sclerite, protruding medioposterior lobe subquadrate, wide, about half the width of posterior margin. Tergum VIII about two-thirds the length of tergum VII, marginosclerite narrow along anterior margin and wider posterolaterally. Anterior apophyses extended from about middle of lateral margin; posterior margin lined with straight row of fine setae, membranous area with very fine longitudinal wrinkles. Sternum VIII membranous, posterior half medially depressed, surface with semiconcentric wrinkles, zone of microtrichia limited to posterior quarter of sternum and not extended into wrinkled zone, with few scattered fine setae on each side of posterior margin. Sinus vaginalis subtriangular in outline, nearly flat with rest of sternal surface. Ostium bursae situated at anterior end of smooth surface of sinus. Antrum shallowly conical around margin of ostium. Colliculum as a darkly melanized gutter with fine, tightly set, obliquely transverse chitinized folds and about as long or slightly shorter than smooth sinus ridge. Anterior apophyses about 1.5x length of tergum VIII, extended to middle of segment VII. Ovipositor about 1.75x length of sternum VIII, membranous, similar to that of S. isaferna in relative length (compared to abdomen) and with similar microsculpture.

  • Biology: Reared from Sesuvium portulacastrum, as a miner on Sesuvium sp., and on Portulaca sp. The first two records may all refer to Sesuvium portulacastrum (L.) L. (Aizoaceae) although the other species of Sesuvium in the Galápagos, S. edmonstonei Hook. f. should not be excluded. Both species are native to and widespread in the Galápagos (Galápagos Species Database, 2024). The genus Portulaca (Portulacaceae) includes five species in the Galápagos (Galápagos Species Database, 2024). The moths are attracted to light and specimens have been collected close to the seashore and up to 850 m in elevation.

  • Distribution: Presumably endemic to the Galápagos, S. galapagensis is the most widely collected of the species, having been found on the islands of Bainbridge, Española, Genovesa, Isabela, Pinta, Pinzón, San Cristóbal, Santa Cruz, Santa Fé, Santiago, Seymour Norte, and South Plaza.

  • Figs 109-112.

    Female abdomen and external genitalia of Galápagos Scythris, ventral aspect. (109) S. isaferna, slide MIC 8762. (110) S. pistillata, slide JFL 1826. (111) S. falcata, slide MIC 8717. (112) S. forcipata, slide MIC 8721.

    img-z26-1_147.jpg

    Figs 113-116.

    Female abdomen and external genitalia of Galápagos Scythris, ventral aspect. (113) S. galapagensis, slide JFL 1834. (114) S. ancystra, slide JFL 1830. (115) S. roquealbeloi, slide MIC 8719. (116) S. cristobalensis, slide MIC 8720.

    img-z29-1_147.jpg

    Figs 117-118.

    Female abdomen and external genitalia of Galápagos Scythris, ventral aspect. (117) S. furculata, slide JFL 1809. (118) S. lisae, slide JFL 1811 (holotype).

    img-z30-1_147.jpg

    Scythris ancystra J.-F. Landry & Bucheli, sp. nov.
    Figs 17-19, 38, 47, 54, 63, 73, 85, 94, 103, 114, 132, 141, 143

  • Material examined

  • Holotype: ♂. 1- “ECU[ADOR], Galápagos, Santa Cruz | NNW Bella Vista, GPS: | 225 m elev[ation]. S 00° 41.293' | W 090° 19.665' | 18.ii.2005, u[ltra]v[iolet] l[ight] | B. Landry & P. Schmitz”. 2- “Database # | CNCLEP | 00014091”. 3- “Barcodes of Life Project | Leg removed | DNA extracted”. 4- “genitalia slide ♂ | JFL 1822”. 5- “HOLOTYPE ♂ | Scythris | ancystra | Landry & Bucheli”. Deposited in MHNG.

  • Paratypes: 15 ♂, 9 ♀ from the Galápagos Islands. – Fernandina: 1♀, North side, 300 m, S 00°20.541' W 091°36.815', U[ltra]V[iolet]L[ight], 12.i.2002 (L. Roque, C. Causton) (CNCLEP00009469); 2 ♂, 1 ♀, SW side, crater rim, reared from, found dead in cage, 12.ii.2005 (L. Roque) (CNCLEP00014101- CNCLEP00014103); 3 ♂, SW side, GPS: 815 m elev[ation]., S 00° 21.270', W 091° 35.341', UVL, 14.ii.2005 (B. Landry & P. Schmitz) (CNCLEP00014080, CNCLEP00014081). – Isabela: 1 ♂ (dissected, slide MIC 4966), 1 km W Puerto Villamil, M[ercury]V[apour]L[ight], 3.iii.1989 (B. Landry) (CNCLEP00004064, BOLD DNA # jflandry1355); 1 ♂ (dissected, slide SRB 219), 2 km W Puerto Villamil, MVL, 5.iii.1989 (B. Landry) (CNCLEP00004065); 1 ♀ (dissected, slide MIC 8689), 3 km N S[an]to Tómas, Agr[iculture]. Zone, MVL, 8.iii.1989 (B. Landry) (CNCLEP00004088); 2 ♂, 11 km N Puerto Villamil, MVL, 9.iii.1989 (B. Landry) (CNCLEP00004067, CNCLEP00004068); 1 ♂ (dissected, slide MIC 2000), same data except CNCLEP00004066, BOLD DNA # jflandry1342); 1 ♂ (dissected, slide JFL 1625), NE slope Alcedo, near shore, GPS: 9 m elev., S 00°23.619' W 90°59.715', uvl, 29.iii.2004 (B. Landry, P. Schmitz) (CNCLEP00005972); 1 ♀, NE slope Alcedo, GPS: 292 m elev., S 00°23.829' W 91°01.957', uvl, 30.iii.2004 (B. Landry, P. Schmitz) (CNCLEP00005949); 3 ♀, NE slope Alcedo, near pega-pega camp, GPS: 483 m elev., S 00° 24.029', W 91° 02.895', uvl, 31.iii.2004 (B. Landry, P. Schmitz) (CNCLEP00005974, CNCLEP00005975, CNCLEP00005978). – Pinzón: 1 ♂, Playa Escondida, UVL, 20.iv.2002 (L. Roque, B. Landry) (CNCLEP00009470). – Santa Cruz: 1 ♂, 1♀, same data as holotype except CNCLEP00014090, CNCLEP00014089; 1 ♀, C[harles]D[arwin]R[research] S[tation], Base of El Barranco, GPS: S 00°44.305' W 90°18.105', uvl, 18.iii.2004 (B. Landry, P. Schmitz) (CNCLEP00005967). – Santiago: 1♂, N side, GPS: 437 m elev., S 00°13.316' W 090°43.808', uvl, 3.iii.2005 (P. Schmitz) (CNCLEP00014097); 1♂, N side, GPS: 527 m elev., S 00°13.690' W 090°44.135', uvl, 5.iii.2005 (P. Schmitz) (CNCLEP00014098). Deposited in CDRS, CNC, and MHNG.

  • Etymology: The new name is derived for the Greek ankistron, meaning fishhook, in reference to the shape of the apex of the phallus.

  • Diagnosis: Scythris ancystra is most similar in genitalia to S. galapagensis and the two species differ in small but constant details. S. ancystra has the dorsal margin of the valva with a lower hump and more pronounced sinuation beyond middle than S. galapagensis, the ventral margin with a slight sinuation in basal half, the phallus broadly arched to 180°, its bulbous crest lacks a basal lobe, there is a small, spiniform process anterior of phallotreme, and sternum VIII has the apex of the medioposterior projection dilated in lateral aspect. In male S. galapagensis, the valva has the dorsal margin with a more pronounced hump, the ventral margin is evenly curved without sinuation; the phallus is bent at 90° in middle, the distal crest is recurved and with a rounded lobe or bump at its base, it lacks a spiniform process anterior of the phallotreme, and the posterior projection of sternum VIII is the same width throughout. In female genitalia, the elevated, smooth ridge of the sinus vaginalis is subconical in outline, the section of colliculum with dense cuticular folds is slender and longer than the smooth sinus ridge; the medioposterior process is small and much narrower than the posterior margin; the anterior emargination of sternum VI is wide and deeper than half the length of the plate.

  • The lateral tufts of scales projecting dorsomedially on the occiput from behind the antennae are orange brown, a colour also found only in S. roquealbeloi.

  • Description: Male (n=16) (Figs 17, 18). Head: occiput appearing orange brown on basal half and dark greyish brown distally, or mostly orange brown or a mixture of both, depending on angle of light; lateral tufts of scales projecting dorsomedially from behind antennae orange brown; frontoclypeus greyish brown mixed with paler orange brown. Antenna blackish brown with purple shine to greyish brown, dorsally without white patch on scape, ventrally white on scape and pedicel and with row of pale brown scales along flagellum; pecten with up to 16 scales of various shades of brown. Labial palpus: 1st palpomere variable shades of brown, often with paler light brown scales dorsally and sometimes ventrally, sometimes all whitish, 2nd mostly dark brown, with paler greyish brown dorsally and sometimes at apex and base, 3rd greyish brown with paler, whitish dorsally. Haustellum uniformly with various shades of greyish brown. Thorax various shades of brown, with tegulae often paler orange brown toward apex, blackish brown with purple shine laterally and apically on mesothorax. Wings as shown (Figs 17, 18); forewing upperside highly variable, with white streaked specimens sometimes with more restricted streak, or with streak orange brown as in many females (Fig. 19); forewing underside dark brown, with greyish brown patch from costa as distal section of retinaculum in most specimens; forewing length: 4.1-5.0, mean 4.5 ± 0.3, n=16 (holotype: 4.4 mm); hindwing underside dark brown with whitish in anal sector, or more extensively whitish on dorsal side of wing until tornus. Foreleg coxa dark to light greyish brown with white mediolaterally and pale greyish brown to white apically; femur dark greyish brown; tibia and tarsomere I dark greyish brown, sometimes with lighter scales at apex; tarsomeres II-V dark greyish brown. Midleg coxa mixed greyish brown and satiny white; femur dark greyish brown with satiny white along ventral margin; tibia dark greyish brown with light brown to white patches postmedially and apically; tarsomeres greyish brown, 1st and sometimes 2nd with white at apex. Hindleg coxa mixed satiny white and pale greyish brown; femur ivory white with dark greyish brown apically; tibia dark greyish brown with satiny white patches postmedially and apically, or lighter greyish brown with less contrasting dirty white patches, with crest of hair-like scales dirty white; tarsomeres dark greyish brown, 1st white at base and apex, 2nd and 3rd sometimes white at apex. Abdomen dorsally pale greyish brown, with slightly warmer brown long hair-like scales over genitalia, or blackish brown on distal two scaled tergites; ventrally white to cream toward apex or all ivory white.

  • Male abdomen and genitalia (8 preparations) (Figs 38, 47, 54, 63, 73, 85, 94, 103). Very similar in most respects to S. galapagensis except as follows: Sternum VIII medioposterior projection with sides more abruptly narrowed and with a more dilated dorsoapical lobe. Valva with dorsal margin nearly straight, barely swollen in distal half, ventral margin angulate in basal half and sinuate in distal half, evenly upcurved apically with slightly more extensive zone of peg-like setae. Phallus more broadly arched at mid-length, dorsal crest more rounded and more dilated, terminal portion straighter and with smaller apical hook; a small, blunt process anterior of phallotreme.

  • Female (n=9) (Fig. 19). Head: occiput and frontoclypeus sometimes darker blackish brown than male in addition to orange brown scales arising from apical margin of occiput; thorax generally darker than male in middle, blackish brown with purple shine except for orange brown tegulae and some scales on basal ‘collarette’; forewing upperside usually mostly blackish brown with purple shine with orange brown longitudinal stripe and sometimes more extensive orange brown along dorsum on basal 1/3 and before tornus; forewing underside dark brown, with white patch from costa as distal section of retinaculum, followed by white scales along costa until about 1/3 wing length; forewing length: 4.0-4.7, mean 4.5 ± 0.2; n=9; hindwing underside dark brown with whitish in anal sector and dorsal side of wing until tornus; frenulum with 3 bristles. Abdomen dorsally pale greyish brown on three basal tergites, darker greyish brown on following tergites, especially distal one; ventrally satiny white to ivory white distally, or ivory white to cream distally.

  • Female abdomen and genitalia (5 preparations) (Figs 114, 132, 141, 143). Very similar in most respect to S. galapagensis except as follows: Anterior emargination of sternum VI wide and deeper than half the length of the plate. Sternum VII with smooth medioposterior zone semi-circular, less extensive, about 0.45x length of sclerite, medioposterior process small and much narrower than posterior margin. Sternum VIII with thicker cuticular folds; sinus vaginalis subconical in outline and elevated, protruding from sternal surface; section of colliculum with dense cuticular folds slenderer, longer than sinus ridge.

  • Biology: Unknown except that the moths are attracted to light and have been collected at a wide range of elevations, from beach sites to the rim of the volcano on Fernandina, which culminates at 1476 m (Wikipedia).

  • Distribution: So far only known from the Galápagos islands of Fernandina, Isabela, Pinzón, Santa Cruz, and Santiago.

  • Remarks: Three specimens collected on Fernandina in 2005 have on their labels “found dead in cage”, suggesting that they were reared, but there is no indication of host plant.

  • Figs 119-125.

    Female abdomen and external genitalia of Galápagos Scythris, dissections photographed in lactic acid. (119) S. isaferna, segments V-ovipositor, lateral aspect, dissection MIC 8762. (120) S. isaferna, segments VI-VII, lateral aspect, dissection JFL 1827. (121) S. pistillata, segments V-ovipositor, lateral aspect, dissection JFL 1826. (122) S. isaferna, sterna VI-VII, ventral aspect, dissection MIC 8762. (123) S. isaferna, terga VI-VII, dorsal aspect, dissection MIC 8762. (124) S. pistillata, sterna VI-VII, ventral aspect, dissection JFL 1826. (125) S. pistillata, terga VI-VII, dorsal aspect, dissection JFL 1826.

    img-z32-1_147.jpg

    Scythris roquealbeloi J.-F. Landry & Bucheli, sp. nov.
    Figs 20-22, 40, 49, 55, 64, 74, 78, 86, 95, 105, 115, 133, 144

  • Material examined

  • Holotype: ♂. 1- “ECU[ADOR], Galápagos, Isabela | NE slope Alcedo, GPS: elev[ation]. | 292 m, S 00° 23.829' W 091° | 01.957', 30.iii.2004, u[ltra]v[iolet]l[ight] | leg[it]. B. Landry, P. Schmitz”. 2- “Database # | CNCLEP | 00005951”. 3- “genitalia slide ♂ | JFL 1627”. 4-“Barcodes of Life Project | Leg removed | DNA extracted”. 5- “genitalia slide ♂ | JFL 1627”. 5- “HOLOTYPE ♂ | Scythris | roquealbeloi | Landry & Bucheli”. Deposited in MHNG.

  • Paratypes: 5 ♂, 9 ♀ from the Galápagos Islands. – Fernandina: 1 ♂, SW side, GPS: 815 m elev., S 00°21.270', W 091°35.341', uvl, 11.ii.2005 (B. Landry, P. Schmitz) (CNCLEP00014077). – Floreana: 1 ♂(dissected, slide JFL 1620, preparation lost), Scalesias near Cerro Pajas, GPS: 329 m elev[ation]., S 01°17.743' W 90° 27.111', u[ltra]v[iolet]l[ight], 12.iv.2004 (P. Schmitz) (CNCLEP00014080). – Isabela: 1 ♂ (dissected, slide JFL 1624), NE slope Alcedo, near shore, GPS: 9 m elev., S 00°23.619' W 90°59.715', u[ltra]v[iolet][light], 29.iii.2004 (B. Landry, P. Schmitz) (CNCLEP00005971, BOLD DNA # jflandry1358). 1 ♀, same data as holotype except CNCLEP00005950, BOLD DNA # jflandry1408; 1 ♀, same data as holotype except CNCLEP00005953, BOLD DNA # jflandry1407; 1 ♀ (dissected, slide JFL 1801), NE slope Alcedo, near pega-pega camp, GPS: 483 m elev., S 00° 24.029', W 91° 02.895', uvl, 31.iii.2004 (B. Landry, P. Schmitz) (CNCLEP00005977); 1 ♂ (dissected, slide JFL 1629), Alcedo, lado [side] NE, 200 m, camp arida alta, uvl, 14.iv.2002 (B. Landry, L. Roque) (CNCLEP00005946); 1 ♀, same data except CNCLEP00005945; 1 ♂ (dissected, slide JFL 1630), Alcedo, lado NE, 400 m, camp pega-pega, uvl, 15.iv.2002 (B. Landry, L. Roque) (CNCLEP00005942, BOLD DNA # jflandry1357); 3 ♀, same data except CNCLEP00005939, CNCLEP00005941, CNCLEP00005944; 1 ♀, same data except CNCLEP00005943, BOLD DNA # jflandry1415; 1 ♀, same data except CNCLEP00005945, BOLD DNA # jflandry1414. Deposited in CNC and MHNG.

  • Etymology: This species is dedicated to lepidopterist Lazaro Roque Albelo, who collected part of the type series and who was very helpful to BL and Patrick Schmitz during their combined four field trips to the Galápagos in the early 2000's.

  • Diagnosis: In male genitalia, the valva has the dorsal margin evenly curved with a short, sharp preapical sinuation, which is narrower than in both S. galapagensis and S. ancystra, the basal half is broader than the distal half, much more so than in S. galapagensis or S. ancystra, with an angulate ventral margin; the gnathos has the apex finely spinulate; the phallus has a sharp medial bend, a broad, massive dorsal crest and bifurcated apex, each bifurcation ending in a small hook or point; the medioposterior projection of sternum VIII is straight-sided and apically truncate (dorsal aspect), not lobate, and proportionally shorter relative to the base than in either S. galapagensis and S. ancystra.

  • In female genitalia, the antrum is a short, transverse, stubby cone, the colliculum is transparent, without dense cuticular folds, and the posteromedial lobe of sternum VII is a thin, transverse band which is almost indistinct; the anterior emargination of sternum VI is narrower than in S. galapagensis and S. ancystra.

  • Description: Male (n=6) (Fig. 20). Head: occiput dark brown, with few orange-brown scales from posterior margin, with orange-brown to paler whitish scales laterally next to antennae; tufts of scales projecting dorsomedially from behind antennae mostly orange brown, sometimes also with paler, cream scales, mostly not touching each other; frontoclypeus dark brown to paler brown, with paler brown scales, especially laterally. Antenna blackish brown, with white underneath scape and pedicel, with row of cream scales along flagellum ventrally; pecten with up to 12 pale brown scales. Labial palpus satiny white with greyish brown patch at base on 1st palpomere, mostly greyish brown with various numbers of white scales dorsally on 2nd and 3rd palpomeres. Haustellum greyish brown medially, satiny white laterally. Thorax mostly dark brown, with some paler brown scales and darker blackish brown spots on mesothorax at apex and laterally before apex. Wings as shown (Fig. 20); forewing upperside brown, with paler line sometimes from base to tornus, with darker brown spot at base on costa, with dark brown streaks, often with purple shine above and below midline, and toward apex; forewing underside brown, gradually darkening toward apex, with white scales on costa as distal end of retinaculum; hindwing underside paler greyish brown on basal half, gradually darker brown toward apex. Forewing length: 3.9-4.4 mm, mean 4.2 ± 0.2, n=6 (holotype: 4.4 mm). Foreleg coxa dark brown with white mostly at apex; femur satiny white at base, dark brown toward apex; tibia dark brown, with few paler scales at apex; tarsus dark brown. Midleg coxa satiny white and greyish brown; femur dark brown with satiny white along ventral margin and at base; tibia dark brown, with few paler greyish brown scales subbasally and postmedially, with few white scales apically; tarsomere I dark brown with few paler scales at apex; tarsomeres II-V dark brown. Hindleg coxa satiny white and greyish brown; femur satiny white with greyish brown at apex; tibia blackish brown to paler greyish brown, with few paler, sometimes including white scales at base of median spurs and usually at apex; tarsomere I blackish brown with white at base and apex; tarsomere II dark brown with white at apex; tarsomeres III-V gradually darker brown, without paler apical scales. Abdomen dorsally greyish brown; ventrally satiny white at base to cream colored toward distal end.

  • Male abdomen and genitalia (6 preparations) (Figs 40, 49, 55, 64, 74, 78, 86, 95, 105). Tergum VIII and sternum VIII similar to those of S. galapagensis except sternum VIII with base more square, anterior emargination more straight-sided and narrower, with thicker strengthening sulci; medioposterior projection nearly straight-sided, very slightly upcurved, apex slightly wider and truncate, not dilated.

  • Lateroposterior extensions of tegumen with apex well sclerotized and upcurved at level of, and posterior to gnathos. Uncus-socii lobes prominently protruded dorsally, bean-shaped in dorsal view. Paired ducts widened, slightly upcurved. Gnathos slender, sharply demarcated from lateroposterior extensions of tegumen by unmelanized zone, apex dorsally with minutely spinulate tooth, ventrally with a rounded process. Valva with broad basal two-thirds, dorsal margin evenly curved with short sinuation before apex; ventral margin in distal half keeled and ridged on outer surface, ventroapical margin with narrow zone of elongate peg-like setae. Phallus markedly bent down in distal half which is oriented anteriorly more so than in the other species, subterminal crest very large, ridged, terminal portion beyond phallotreme bifurcated, each bifurcation ending in a small hook or point.

  • Female (n=9) (Figs 21, 22). Head as in male, sometimes with more orange-colored scales. Thorax as in male, sometimes with more sharply contrasting paler brown apex of tegulae; forewing upperside often with more contrasting pattern of orange-colored stripe from base to tornus, often with white underline, and darker brown scaling with purple shine along costal half and at base along dorsum although often covered here with orange brown; forewing underside dark brown except for white scales from costa as part of retinaculum distally, and paler greyish brown scales along costa nearly until apex of wing and scattered at base; forewing length: 4.1-5.0 mm, mean 4.5 ± 0.3, n=9; hindwing underside paler greyish brown at base, darker brown below costa subbasally and toward apex; frenulum with 3 bristles (one specimen with 2, CNCLEP00005950). Abdomen dorsally mostly greyish brown, with darker brown on last scaled tergite; ventrally all ivory white.

  • Female abdomen and genitalia (2 preparations) (Figs 115, 133, 144). Tergum and sternum VI similar to those of S. galapagensis, sternum VI proportionally slightly longer with narrower conical anterior emargination. Sternum VII with medially sinuate posterior margin at level of smooth zone, the latter small in extent, about 0.2x length of sclerite; anterior margin medially sinuate, posteromedial lobe small, narrowly transverse, barely protruded, almost indistinct. Tergum VIII proportionally wider than that of S. galapagensis, laterodistal portion of marginosclerite wide, dilated, sides of anterior portion converging medially. Sternum VIII with fine and dense transverse wrinkles forming an anteriorly sloping sinus vaginalis, portion around ostium smooth, deeply inset, antrum a short, flat, sclerotized cone with truncate apex projecting ventrally (away from sternal wall); colliculum without cuticular folds; ostium bursae small.

  • Biology: Unknown but for the moths being attracted to light and the 200-815 m range in elevation of its known collecting sites.

  • Distribution: A presumed Galápagos endemic, this species has been found on the islands of Fernandina, Floreana, and Isabela.

  • Figs 126-130.

    Female sterna VI-VII and external genitalia (sternum VIII) of Galápagos Scythris, ventral aspect. (126) S. isaferna, slide MIC 8762. (127) S. pistillata, slide JFL 1826. (128) S. falcata, slide MIC 8717. (129) S. forcipata, slide MIC 8721. (130) S. furculata, slide JFL 1809.

    img-z35-1_147.jpg

    Figs 131-135.

    Female sterna VI-VII and external genitalia (sternum VIII) of Galápagos Scythris, ventral aspect. (131) S. galapagensis, slide JFL 1834. (132) S. ancystra, slide JFL 1830. (133) S. roquealbeloi, slide MIC 8719. (134) S. cristobalensis, slide MIC 8720. (135) S. lisae, slide JFL 1811 (holotype).

    img-z36-1_147.jpg

    Scythris falcata J.-F. Landry & Bucheli, sp. nov.
    Fig 14-16, 42, 48, 56, 65, 75, 79-81, 87, 96, 104, 111, 128, 138

  • Material examined

  • Holotype: ♂. 1- “ECUADOR [sideways on left side] | GALÁPAGOS | Santa Crúz | Los Gemelos | 31.I.1989, M[ercury]V[apour]L[ight] | B. Landry”. 2- “C.N.C. | Genitalia Slide ♂ | MIC 2001”. 3-. “Database # | CNCLEP | 00004047”. 4- “Barcodes of Life Project | University of Guelph | DNA # jflandry1348”. 5- “HOLOTYPE ♂ | Scythris | falcata | Landry & Bucheli”. Deposited in CNC. Paratypes: 11 ♂, 14 ♀ from the Galápagos Islands. – Isabela: 1 ♂ (dissected, slide MIC 8711), 3 km S[an] to Tómas, Agr[iculture]. Zone, M[ercury]V[apour] L[ight], 8.iii.1989 (B. Landry) (CNCLEP00004089, BOLD DNA # jflandry1356); 2 ♂ (one dissected, slide JFL 1821), 2 ♀, V[olcan]. Alcedo, Lado Este [East side] 900 m[e]t[er]s elev[ation]., Ex larva en Althernanthera [sic] helifolia, 6.iv.1999 (L. Roque) (CNCLEP0000946, CNCLEP00009463-CNCLEP00009465); 1 ♀, Alcedo, lado [side] NE, 400 m, camp pega-pega, uvl, 15.iv.2002 (B. Landry, L. Roque) (CNCLEP00005940, BOLD DNA # jflandry1416); 1 ♂ (dissected, slide JFL 1631), Alcedo, lado NE, 700 m, camp guayabillos, u[ltra] v[iolet][light], 16.iv.2002 (B. Landry, L. Roque) (CNCLEP00005935); 1 ♀ (dissected, slide JFL 1804), same data except CNCLEP00005938, BOLD DNA # jflandry1417; 1 ♀, same data except CNCLEP00005936, BOLD DNA # jflandry1418; 1 ♀, same data except CNCLEP00005937; 1 ♂ (dissected, slide BL 1156), ± 15 km N Puerto Villamil, MVL, 25.v.1992 (B. Landry) (CNCLEP00004161, BOLD DNA # jflandry1339); 1 ♀, Alcedo, zona arida alta, UV-FL, 13.x.1999 (L. Roque) (CNCLEP00009432). – San Cristóbal: 1 ♂, antiguo botadero, ca. 4 km SE Puerto Baquerizo, GPS: 169 m elev[ation]., S 00°54.800' W 089°34.574', uvl, 22.ii.2005 (B. Landry) (CNCLEP00014100). – Santa Cruz: 5 ♂, 5 ♀, Los Gemelos, Ex larva minadora de Althernanthera [sic] helifolia, 12.i.1999 (L. Roque No 99.27) (CNCLEP00009455-CNCLEP00009457, CNCLEP00009461 & slide MIC 8712, CNCLEP00009462, CNCLEP00004169, CNCLEP00004170, CNCLEP00009458 & slide MIC 8717-CNCLEP00009460); 1 ♀, same data as holotype except CNCLEP00004048); 1 ♀ (dissected, slide MIC 4962), Finca S[teve]. Devine, MVL, 17.iii.1989 (B. Landry) (CNCLEP00004050, BOLD DNA # jflandry1334). Deposited in CDRS, CNC, and MHNG.

  • Etymology: The name is from the Latin falcatus for sickle, in reference to the sickle-shaped dorsal process of the valva.

  • Diagnosis: In male genitalia, the dorsal margin of the valva is markedly arched and has a sharp, spine-like process at about two-thirds followed by a deep cleft, the ventral margin is sharply sinuate and the distal portion is broadly spatulate and densely covered with spiniform setae; the phallus dorsal crest is conical, angular and the apex is attenuate with a tiny, barely discernable hook; sternum VIII has a broadly digitiform medioposterior projection with a subtruncate or notched apex. The species is most similar in genitalia to S. forcipata; the latter is distinguished by the dorsal margin of valva which is more gently curved, has a blunt spine, and a shallower cleft, narrower and more rounded paddle-like distal portion of valva, the angulate phallus with a rounded crest and very short phallus apex, and linguiform medioposterior projection of sternum VIII with rounded apex.

  • The female genitalia are somewhat similar to those of S. galapagensis except that the smooth medioposterior zone of sternum VII is semi-circular and much smaller, about 0.2x the length of the sclerite (about 0.6x and hemielliptic in S. galapagensis); sternum VIII is more densely wrinkled but the wrinkles are finer and more longitudinal than oblique; the sinus vaginalis is subrectangular in outline and not distinctly delineated, and the section of colliculum with dense cuticular folds is very short and not clearly discernible; the anterior emargination of sternum VI is shallow.

  • This species has an orange-brown head including the tufts of scales projecting dorsomedially from behind antennae. The shiny bronze sheen of the upper surface of the forewing is a feature shared by S. falcata and some specimens of S. ancystra and S. roquealbeloi, although it is more pronounced than in the latter. Despite this, identification of the species must be confirmed by examination of genitalia.

  • Description: Male (n=12) (Figs 14, 15). Head: occiput and frontoclypeus orange brown with bronze shine, including lateral tufts of scales projecting dorsomedially from behind antennae, sometimes with greyish brown. Antenna blackish brown, with lighter greyish brown scape; scape and pedicel ventrally white to cream; flagellum with row of cream scales ventrally; pecten with up to 15 light orange brown to dark brown scales. Labial palpus: 1st palpomere white to light brown, 2nd pale greyish brown with whitish to light brown dorsally and sometimes a white patch apically, 3rd greyish brown with whitish to light brown dorsally. Haustellum scales at base white laterally and light brown along middle, distally light greyish brown. Thorax various shades of brown, usually paler at tip of tegulae. Wings as shown (Figs 14, 15); forewing upperside sometimes with scattered contrasting satiny white or blackish brown scales, the latter shining purple sometimes; forewing underside dark brown, with patch from costa as distal section of retinaculum sometimes contrastingly paler, sometimes with paler scales along costa from tip of retinaculum; forewing length: 4.0-4.8 mm, mean 4.4 ± 0.2, n=12 (holotype: 4.5 mm); hindwing underside light brown, darkening slightly toward apex, sometimes darker along costal half and apex. Foreleg coxa greyish brown at base, light brown elsewhere; femur greyish brown; tibia and palpomere I blackish brown with few paler light brown scales apically; palpomeres II-V greyish brown, sometimes white at apex of 2nd. Midleg coxa satiny white; femur light brown with white along ventral margin and dark greyish brown at apex; tibia light brown with dark greyish brown at base and light to dark greyish brown subapically; tarsomeres I and II greyish brown with white at apex; tarsomeres III-V greyish brown. Hindleg coxa satiny white; femur white with blackish brown apex; tibia ivory white with dark greyish brown patch at base and lighter, sometimes faint, patches at 2/5 and subapically; tarsus greyish brown, with tarsomeres I to IV paler at apex, sometimes with 1st almost completely pale. Abdomen dorsally greyish brown, darkening toward distal segments, with longer scales above genitalia slightly warmer brown; ventrally homogeneously ivory white.

  • Male abdomen and genitalia (6 preparations) (Figs 42, 48, 56, 65, 75, 79-81, 87, 96, 104). Tergum VIII similar to that of S. galapagensis except posterior margin more broadly concave. Sternum VIII with medioposterior projection wider and with straighter sides than in S. galapagensis, weakly upcurved, apex not dilated, subtruncate, slightly notched in some specimens.

  • Uncus-socii and gnathos similar to those of S. galapagensis, almost reaching apex of tegumen extensions, uncus slightly more elevated. Paired ducts relatively long, sclerotized, very apparent. Gnathos apex downcurved, beak-like, ventral process short, barely produced. Valva with obliquely angulate basal articulation, basal half with ventral margin sinuate and dorsal margin arched and produced in middle into a sharp, incurved, spiniform process; distal half roundly spatulate, paddle-like, densely covered with peg-like setae. Phallus more evenly downcurved than that of S. galapagensis, subterminal ventral hump conical, apical portion shorter, straight, with minute terminal hook.

  • Female (n=14) (Fig. 16). Head, thorax, and wings as in male; forewing length: 4.2-4.8 mm, mean 4.5 ± 0.2, n=16; frenulum with 3 bristles. Abdomen dorsally pale greyish brown on first two tergites, usually darker greyish brown on following tergites, sometimes with white scales, with distal scales sometimes paler, ivory white; ventrally homogeneously ivory white to light brown.

  • Female abdomen and genitalia (3 preparations) (Figs 111, 128, 138). Similar in many respects to those of S. galapagensis except as follows: Sternum VI with anterior margin shallowly concave or incurved. Sternum VII conical with smooth posteromedial zone hemicircular, small, no more than 0.2x length of sclerite; posteromedial lobe wider, subrectangular. Tergum VI more extended ventrally over pleural region, ending in subfold under sides of sternum VII and with firmly attached scales. Sternum VIII with obliquely wrinkled, concave sinus vaginalis, without elevated area around ostium bursae, margin of ostium with barely a thin sclerotized rim, antrum with very short thickened/folded section followed anteriorly by short, smooth-walled section before membranous ductus bursae.

  • Biology: Fourteen of the moths studied were reared from larvae leaf mining “Alternanthera helifolia” on Isabela and Santa Cruz islands. This plant name could not be recovered on the International Plant Names Index (IPIN, 2024) or the Galápagos Species Database (2024). We suspect that it refers to Alternanthera halimifolia Standl. ex Pittier. (Amaranthaceae) because of its distribution on both islands (Galápagos Species Database, 2024). However, a misidentification for another Alternanthera species is also possible, and it is also possible that S. falcata larvae feed on several species of this genus. The moths are attracted to light and were found in various habitats from near the shoreline up to 900 m in elevation.

  • Distribution: Presumably endemic to the Galápagos islands and found thus far on Isabela, San Cristóbal, and Santa Cruz.

  • Figs 136-139.

    Female sternum VIII/external genitalia of Galápagos Scythris; images obtained with DIC illumination. (136) S. isaferna, slide MIC 8713. (137) S. pistillata, slide MIC 8718. (138) S. falcata, slide JFL 1804. (139) S. forcipata, slide MIC 8690.

    img-z38-1_147.jpg

    Scythris forcipata J.-F. Landry & Bucheli, sp. nov.
    Figs 29-33, 43, 50, 57, 66, 76, 88, 97, 106, 112, 129, 139

  • Material examined

  • Holotype: ♂. 1- “ECU[ADOR], Galápagos, Fernandina | SW side, GPS: 815 m elev[ation]. | S 00° 21.270', W 091° 35.341' | u[ltra]v[iolet]l[ight], 14.ii.2005 | B. Landry & P. Schmitz”. 2- “Database # | CNCLEP | 00014079”. 3- “Barcodes of Life Project | Leg removed | DNA extracted”. 4- “genitalia slide ♂ | JFL 1823”. 5- “HOLOTYPE ♂ | Scythris | forcipata | Landry & Bucheli”. Deposited in MHNG.

  • Paratypes: 21 ♂, 23 ♀ from the Galápagos Islands. – Española: 1 ♂ (dissected, slide JFL 1634), Bahía Manzanillo, M[ercury]V[apour]L[ight], 25.iv.1992 (B. Landry) (CNCLEP00004136); 1 ♂, same data except 29.iv.1992 and CNCLEP00004135). – Fernandina: 1 ♂, 1 ♀, same data as holotype except CNCLEP00014084 and CNCLEP00014083. – Floreana: 1 ♂ (dissected, slide JFL 1635), close to Loberia, GPS: elev[ation]. 6 m, S 01°17.002' W 90°29.460', 11.iv.2004, u[ltra]v[iolet] l[ight] (P. Schmitz) (CNCLEP00005959). – Isabela: 1 ♂ (dissected, slide MIC 4964), 1 km W Puerto Villamil, MVL, 3.iii.1989 (B. Landry) (CNCLEP00004056); 1 ♂, same data except CNCLEP00004059 and BOLD DNA # jflandry1386; 2 ♀, same data except CNCLEP00004057 and CNCLEP00004058; 1 ♀ (dissected, slide MIC 8690), 2 km W Puerto Villamil, MVL, 5.iii.1989 (B. Landry) (CNCLEP00004061); 1 ♀, same data except dissected (slide MIC 8721); 1 ♀, same data except CNCLEP00004063 and BOLD DNA # jflandry1388; 1 ♂, 11 km N Puerto Villamil, MVL, 9.iii.1989 (B. Landry) (CNCLEP00004069); 1 ♂, 8.5 km N P[uer]to Villamil, MVL, 11.iii.1989 (B. Landry) (CNCLEP00004074); 1 ♂, same data except CNCLEP00004075 and BOLD DNA # jflandry1385; 1 ♂ (dissected, slide MIC 4967), same data except CNCLEP00004080 and BOLD DNA # jflandry1384; 1 ♀, same data except CNCLEP00004077 and BOLD DNA # jflandry1328; 1 ♂ (dissected, slide JFL 1628, preparation lost), NE slope Alcedo, near pega-pega camp, GPS: 483 m elev., S 00° 24.029', W 91° 02.895', uvl, 31.iii.2004 (B. Landry, P. Schmitz) (CNCLEP00005976); 2 ♀, V[olcan] Alcedo, 200 m, Arida alta [zone], luz fluorescente, 12.iv.2001 (L. Roque # 2001-06) (CNCLEP00009447, CNCLEP00009448); 1 ♀, Alcedo, lado [side] NE, low arid zone, bosq[ue]. palo santo, uvl, 18.iv.2002 (B. Landry, L. Roque) (CNCLEP00005954 and BOLD DNA # jflandry1419); 1 ♀, Tagus Cove, MVL, 13.v.1992 (B. Landry) (CNCLEP00004164 and BOLD DNA # jflandry1394. – Pinta: 1 ♂, 200 m elev., MVL, 16.iii.1992 (B. Landry) (CNCLEP00004145 and BOLD DNA # jflandry1353); 1 ♀, 400 m elev., MVL, 18.iii.1992 (B. Landry) (CNCLEP00004147 and BOLD DNA # jflandry1352); 1 ♂, ± 50 m elev., MVL, 20.iii.1992 (B. Landry) (CNCLEP00004149); 1 ♂ (dissected, slide SRB 203, preparation lost), same data except CNCLEP00004150; 1 ♂ (dissected, slide SRB 211, preparation lost), same data except CNCLEP00004151; 1 ♀, same data except CNCLEP00004148; 1 ♂, ± 15 m elev., MVL, 21.iii.1992 (B. Landry) (CNCLEP00004154); 1 ♂ (dissected, slide SRB 212), same data except CNCLEP00004158; 1 ♀, same data except CNCLEP00004155 and BOLD DNA # jflandry1382; 1 ♀, same data except CNCLEP00004157 and BOLD DNA # jflandry1383. – Santa Cruz: 1 ♀ (dissected, slide SRB 217), Tortuga Res[erve]. W S[an]ta Rosa, MVL, 6.ii.1989 (B. Landry) (CNCLEP00004049); 1 ♂ (dissected, slide JFL 1636), E[stacion].C[ientifica].C[harles].D[arwin]., MVL, 4.iii.1992 (B. Landry) (CNCLEP00004090 and BOLD DNA # jflandry1378); 1 ♂, E.C.C.D., uvl, 6.iii.1992 (B. Landry) (CNCLEP00004091 and BOLD DNA # jflandry1333); 1 ♂ (dissected, slide JFL 1637), low agriculture zone, GPS: S 00°42.132' W 90° 19.156', uvl, 13.iii.2004 (B. Landry, P. Schmitz) (CNCLEP00005958); 1 ♀, Finca S[teve]. Devine, MVL, 17.iii.1989 (B. Landry) (CNCLEP00004051); 1 ♀, ECCD, El Barranco, 22 m elev., S 00°44.291' W 90°18.107', uvl, 23.iii.2006 (P. Schmitz); 1 ♀, Finca Vilema, 2 km W Bella Vista, MVL, 1.iv.1992 (B. Landry) (CNCLEP00004096 and BOLD DNA # jflandry1399); 1 ♀, Bahía Conway, MVL, 14.iv.1992 (B. Landry) (CNCLEP00004092); 2 ♂, 1 ♀, E.C.C.D., yellow light, MVL, 4.v.1992 (B. Landry) (CNCLEP00004093-CNCLEP00004095); 1 ♀, Tortuga Bay, Costal [sic] zone, 00°46'59”S, 090°20'45”W, fluorescent light, 23.ix.1996 (L. Roque) (CNCLEP00009424). – Santiago: 1 ♀, Bahía Espumilla, MVL, 4.iv.1992 (B. Landry) (CNCLEP00004135). Deposited in CDRS, CNC, and MHNG.

  • Etymology: The new name is derived from the Latin forceps in reference to the claw-like shape of the valva.

  • Diagnosis: In male genitalia, the dorsal margin of valva has a blunt, slightly curved spiniform process at two-thirds, the ventral margin is deeply sinuate, the distal portion is oval, paddle-like and covered with dense peg-like setae; the phallus is bent at almost a right angle, its dorsal crest is rounded and the apex very short; the medioposterior projection of sternum VIII is linguiform with a rounded apex.

  • In female genitalia, this species is most similar to S. falcata except as follows: sternum VI is semicircular (trapezoid in S. falcata); the medioposterior projection of sternum VII is narrower than the smooth medioposterior zone (as wide as, in S. falcata); and the spinulate zone of sternum VIII is extended into the distal portion of the sinus vaginalis (restricted to a band along the posterior margin in S. falcata).

  • Description: Male (n=22) (Figs 29, 30, 32). Head: occiput dark brown with bronze shine, sometimes with white scales from base laterally, with white to greyish brown tufts of scales projecting dorsomedially from behind antennae; frontoclypeus greyish brown, sometimes with white scales laterally, on ventral margin, and elsewhere. Antenna blackish brown, with satiny white on scape at tip and ventrally, sometimes also on pedicel, with row of ivory white scales ventrally on flagellum; pecten with up to about 12 pale brown to dark brown scales. Labial palpus white on first palpomere, white to pale greyish brown basally, dorsally, and apically on second, and sometimes dorsally on third, otherwise dark brown. Haustellum scaled satiny white to dirty white, with brown at base or more extensively. Thorax dorsally mostly dark brown, sometimes with few paler brown to white scales at base of pronotum. Wings as shown (Figs 29, 30, 32); forewing upperside varying in contrast of pattern, but always with black spot on costa subbasally, underside dark brown, with paler, greyish brown scales on costa at distal end of frenulum and also subapically; hindwing underside dark brown with paler band from frenulum base to middle of dorsum. Forewing length: 4.1-4.7 mm, mean 4.5 ± 0.2, n=22 (holotype: 4.7 mm). Foreleg coxa mixed dark brown and satiny white; femur dark brown with satiny white at base, medially, and sometimes apex; tibia and tarsus dark brown. Midleg coxa white and greyish brown; femur dark brown speckled with white and along ventral margin; tibia blackish brown with three satiny white to ivory stripes submedially (faint), postmedially, and apically; tarsomeres I and II blackish brown with white at base and apex of 1st, and apex of 2nd and 3rd; tarsomeres IV and V dark brown with slightly paler median surfaces as other tarsomeres. Hindleg coxa satiny white and greyish brown; femur satiny white with dark brown at apex; tibia mostly white with dark brown spots subbasally, at 1/3, and subapically, with dorsal crest of hair-like scales white to pale brown dorsally, more homogenously white ventrally; tarsomeres as midleg. Abdomen dorsally mostly greyish brown, but warmer brown on last segment; ventrally all white.

  • Male abdomen and genitalia (11 preparations) (Figs 43, 50, 57, 66, 76, 88, 97, 106). Tergum VIII similar to that of S. galapagensis except posterior emargination proportionally narrower. Sternum VIII with medioposterior projection straight-sided, elongate-linguiform, with rounded apex only slightly upcurved in lateral aspect.

  • Tegumen moderately arched, medioposterior extensions apically extended beyond gnathos and forming a well-sclerotized hump; tegumen-uncus-gnathos extended beyond apex of valvae, longer than in all other species except S. lisae. Uncus-socii lobes dorsally prominent, ovoid, with long, stiff setae. Gnathos base bent down just anterior of paired ducts, the latter prominently sclerotized and relatively long; distal process dentiform with rounded ventral lobe recessed under apex of tegumen extensions. Valva resembling that of S. falcata with dorsal margin bearing a recurved hook in distal third but otherwise nearly straight and hook more or less blunt; ventral margin with single broad sinuation; terminal portion spatulate, with dense cover of short peg-like setae. Phallus with basal two-thirds straight, distal third with sharp right-angled, downward bend in subterminal portion, with a large crest-like hump more or less similar to that of S. galapagensis but straighter, apex tapered into short, dentate point.

  • Female (n=23) (Figs 31, 33). Head as in male, although apparently more homogeneously coloured brown. Thorax as in male; forewing upperside with pattern as in male except for presence of orange-brown scales all along midline or as spots, and sometimes at base and middle on dorsal half; forewing underside with white scales along costa as part of retinaculum and beyond middle of wing; hindwing underside mostly whitish with dark brown band along costa from beyond base until apex; frenulum with 3 bristles; forewing length: 3.7-5.2 mm, mean 4.6 ± 0.3, n=23. Abdomen dorsally greyish brown, with apical row of short ivory to greyish white scales on ultimate scaled tergite; ventrally all white.

  • Female abdomen and genitalia (3 preparations) (Figs 112, 129, 139). Tergum VI similar to that of S. galapagensis with extensive pleuroventral extensions encompassing spiracles, although less protruded than in S. galapagensis, and ending in folds under the edge of sternum VI. Sternum VI hemicircular with straight anterior margin and rounded posterior margin. Tergum VII subquadrate with unmelanized longitudinal zone in middle. Sternum VII broadly conical, smooth zone about 0.2x length of plate, oval; posteromedial lobe transversely narrow, short but prominent. Tergum VIII similar to that of S. galapagensis. Sternum VIII with obliquely wrinkled membranous areas and sinus vaginalis similar to that of S. isaferna, elongate but distally narrowed, microspiculate zone slightly extended into distal part of sinus vaginalis, the latter forming an elongate, narrow trough with smooth surface; antrum a short, transverse rim; colliculum very short, stubby, darkly melanized, without cuticular folds. Ovipositor as in S. galapagensis.

  • Biology: Unknown except for the moths being attracted to light and the collecting localities distributed between the seashore and 815 m in elevation at the type locality.

  • Distribution: Presumably endemic to the Galápagos and found on the islands of Española, Fernandina, Floreana, Isabela, Pinta, Santa Cruz, and Santiago.

  • Figs 140-143.

    Female sternum VIII/external genitalia of Galápagos Scythris; images obtained with DIC illumination. (140) S. galapagensis, slide JFL 1834. (141) S. ancystra, slide JFL 1830. (142) S. galapagensis, slide JFL 1834. (143) S. ancystra, slide JFL 1830.

    img-z40-1_147.jpg

    Scythris furculata
    J.-F. Landry & Bucheli, sp. nov.

    Figs 23-26, 39, 58, 67, 77, 89, 98, 107, 117, 130, 145

  • Material examined

  • Holotype: ♂. 1- “ECU[ADOR], Galápagos, Santa Cruz | low agriculture zone, GPS: | S 00° 42.132' W 90° 19.156' | 13.iii.2004, u[ltra]v[iolet]l[ight] | leg[it]. B. Landry, P. Schmitz”. 2- “Database # | CNCLEP | 00005957”. 3- “Barcodes of Life Project | University of Guelph | DNA # jflandry1381”. 4- “genitalia slide ♂ | JFL 1825”. 5- “HOLOTYPE ♂| Scythris | furculata | Landry & Bucheli”. Deposited in MHNG.

  • Paratypes: 34 ♂, 17 ♀ from the Galápagos Islands. – Baltra: 1 ♀, En Vainas Acacia insulae-jacobi, 3.xi.1999 (C. Causton, D. Ramon) (CNCLEP00009434). – Fernandina: 1 ♂, Cabo Douglas, GPS: S 00°18.269', W 091°39.098', u[ltra]v[iolet]l[ight], 9.ii.2005 (B. Landry, P. Schmitz) (CNCLEP00014099); 1 ♂(dissected, slide JFL 1819), SW side, GPS: 352 m elev., S 00°20.503', W 091°36.969', uvl, 10.ii.2005 (B. Landry, P. Schmitz) (CNCLEP00014086); 3 ♂, SW side, GPS: 815 m elev., S 00°21.270', W 091°35.341', uvl, 11.ii.2005 (B. Landry, P. Schmitz) (CNCLEP00014075, CNCLEP00014076, CNCLEP00014078). – Isabela: 1 ♂ (dissected, slide MIC 4965), 3 km N S[an]to Tómas, Agr[iculture]. Zone, MVL, 8.iii.1989 (B. Landry) (CNCLEP00004087); 1 ♂ (dissected, slide JFL 1632), V[olcan]. Darwin, 300 m elev., MVL, 15.v.1992 (B. Landry) (CNCLEP00004163 and BOLD DNA # jflandry1369). – Pinta: 1 ♂, Playa Ibbeston, MVL, 13.iii.1992 (B. Landry) (CNCLEP00004137 and BOLD DNA # jflandry1368); 1 ♂, same data except CNCLEP00004139 and BOLD DNA # jflandry1366; 1 ♂ (dissected, slide SRB 201, preparation lost), same data except CNCLEP00004140; 1 ♂ (dissected, slide SRB 202), same data except CNCLEP00004141; 1 ♂ (dissected, slide SRB 221), Playa Ibbeston, MVL, 14.iii.1992 (B. Landry) (CNCLEP00004142); 1 ♀, same data except CNCLEP00004143; 1 ♂ (dissected, slide SRB 220), arid zone, M[ercury]V[apour]L[ight], 15.iii.1992 (B. Landry) (CNCLEP00004144); 1 ♀(dissected, slide JFL 1802), ± 50 m elev., MVL, 20.iii.1992 (B. Landry) (CNCLEP00004152); 1 ♂(dissected, slide SRB 210, preparation lost), ± 15 m elev., MVL, 21.iii.1992 (B. Landry) (CNCLEP00004153); 1 ♀ (dissected, slide MIC 8755), same data except CNCLEP00004156. – San Cristóbal: 1 ♀, near Loberia, GPS: elev[ation]. 14 m, S 00°55.149' W 89°36.897', uvl, 16.iii.2004 (B. Landry, P. Schmitz) (CNCLEP00005961, BOLD DNA # jflandry1391); 1 ♀, same data except CNCLEP00005962. – Santa Cruz: 1 ♀, transition zone, recently cut road, GPS: S 00°42.528' W 90°18.849', uvl, 12.iii.2004 (B. Landry, P. Schmitz) (CNCLEP00005960, BOLD DNA # jflandry1380); 2 ♀, same data as holotype except CNCLEP00005955, CNCLEP00005956; 1 ♂(dissected, slide JFL 1638), C[harles]D[arwin]R[esearch] S[tation], wall of Invert[ebrate]s Lab., GPS: 11 m elev., S 00°44.478' W 90°18.132', uvl, 19.iii.2004 (B. Landry, P. Schmitz) (CNCLEP00005968); 1 ♂, CDRS, Barranco, UVL, 12.ix.2001 (L. Roque) (CNCLEP00009428); 1 ♂(dissected, slide SRB 206), CDRS, Barranco, MVL, 7.x.1999 (L. Roque) (CNCLEP00004171); 1 ♂ (dissected, slide SRB 205), same data except CNCLEP00004173; 1 ♀, same data except CNCLEP00004172, BOLD DNA # jflandry1396; 1 ♂, CDRS, Barranco, UVL, 23.x.2001 (L. Roque) (CNCLEP00009429); 1 ♂ (dissected, slide SRB 207), C.D.R.S., Barranco, MVL, 11.xi.1999 (L. Roque) (CNCLEP00004174); 1 ♂, C.D.R.S., Barranco, MVL, 17.xi.1999 (L. Roque) (CNCLEP00009437); 2 ♀, same data except CNCLEP00009435, CNCLEP00009436; 4 ♂, 1 ♀, C.D.R.S., Barranco, UVL, 23.xi.2001 (L. Roque) (CNCLEP00009439-CNCLEP00009443); 1 ♂, 2 ♀, C.D.R.S., Barranco, UVL, 10.xii.2001 (L. Roque) (CNCLEP00009444-CNCLEP00009446). – Santa Fé: 1 ♀, Larva en Waltheria ovata, 5.xi.2000 (A. Mieles). – Seymour Norte: 1 ♂, MVL, 29.iii.1992 (B. Landry) (CNCLEP00004105, BOLD DNA # jflandry1360); 1 ♂, same data except CNCLEP00004106, BOLD DNA # jflandry1361; 1 ♂, same data except CNCLEP00004107; 1 ♂ (dissected, slide JFL 1621), same data except CNCLEP00004108, BOLD DNA # jflandry1359; 1 ♂ (dissected, slide SRB 226, preparation lost), same data except CNCLEP00004112; 1 ♂, same data except CNCLEP00004114, BOLD DNA # jflandry1363; 1 ♂, same data except CNCLEP00004115, BOLD DNA # jflandry1364; 1 ♂, same data except CNCLEP00004116, BOLD DNA # jflandry1362; 1 ♀ (dissected, slide JFL 1809), same data except CNCLEP00004109, BOLD DNA # jflandry1365. Deposited in the CDRS, CNC, and MHNG.

  • Etymology: The new name is based on the Latin furcula, meaning small fork, in reference to the forked apex of the phallus.

  • Diagnosis: In male genitalia, the phallus has a markedly bifurcate distal portion with a shorter and smaller dorsal furcation and wider and bigger ventral one; sternum VIII is subquadrate and lacks the medioposterior projection in place of which there is only a small bump, its sides are flat, not dorsally upbent as in the other species; the valva is medially narrowed with a wider, flared distal portion covered with short, peg-like setae; the lateroposterior extensions of the tegumen end in a ventral, recurved process with a small terminal, forked hook. This teguminal process is reminiscent of a gnathos in appearance but is actually ventral to the true gnathos. In female genitalia, sternum VII is subcordiform in outline, its anterolateral angles have short extensions recessed into pockets around the spiracles, its posterior margin is broadly concave, thickly rimmed, and the medioposterior lobe is a thickly sclerotized, subquadrate plate with a concave surface and is set under the margin of the sternal plate. The ostium bursae is bordered by a small half-ring sclerotization anteriorly.

  • Description: Male (n=35) (Figs 23, 26). Head: occiput dark brown, with bronze to more rarely purple shine, with few white to pale brown scales laterally next to base of antennae; tufts of scales projecting dorsomedially from behind antennae mostly dark brown, usually with some pale brown to dirty white scales; frontoclypeus dark brown, as occiput, with white scales showing on ventral margin and white to pale brown laterally. Antenna blackish brown, with satiny white on scape and pedicel ventrally, with row of ivory white scales ventrally on flagellum; pecten with up to about 15 pale to dark brown scales. Labial palpus satiny white on 1st palpomere, 2nd palpomere varying from mostly dark brown with white dorsally to mostly white with dark brown toward apex laterally, 3rd palpomere dark brown with paler, sometimes white scales dorsally. Haustellum scales satiny white and dark greyish brown. Thorax mostly blackish brown, often with few scattered orange-brown scales, sometimes with shining darker spots apically and laterally before apex, surrounded by paler brown scales. Wings as shown (Figs 23, 26); forewing upperside varying in pattern contrast, sometimes with orange-brown scales along midline on basal half and toward dorsum postmedially, sometimes with more or less apparent purple to blue shine, especially toward tornus; forewing underside with few white scales projecting from costa as distal part of retinaculum; hindwing underside with wide stripe of paler greyish brown to white along middle. Forewing length: 3.5-4.5 mm, mean 4.0 ± 0.2, n=35 (holotype: 4.3 mm). Foreleg coxa satiny white with greyish brown; femur satiny white with blackish brown especially toward apex; tibia blackish brown with small white spots medially and apically; tarsus dark brown with white to greyish brown at apex of tarsomeres I and II. Midleg coxa satiny white with greyish brown at base; femur mostly dark brown with white along ventral margin and scattered elsewhere; tibia blackish brown with faint to well-formed submedial greyish to white spot and satiny white to ivory white bands postmedially and apically; tarsomere I blackish brown with white apically; tarsomere II-V paler, greyish brown, with white at tip of tarsomere II. Hindleg coxa satiny white and greyish brown; femur satiny white with blackish brown at apex; tibia mostly white with blackish brown spots subbasally, at 1/3, and subapically, with dorsal crest of hair-like scales white to pale brown dorsally, more homogenously white ventrally; tarsomere I blackish brown with white at base and apically; tarsomeres II-IV dark brown with few white scales apically; tarsomere V dark brown without white apically. Abdomen dorsally greyish brown, with slightly warmer brown hair-like scales over genitalia; ventrally all white.

  • Male abdomen and genitalia (15 preparations) (Figs 39, 58, 67, 77, 89, 98, 107). Tergum VIII narrowly transverse, anterior and posterior margins almost straight, sides rounded. Sternum VIII subquadrate, anterior margin with shallow medial V-shaped emargination less than 0.2x length of sternum, posterior sulci markedly developed, medioposterior projection undeveloped or reduced as a small bump.

  • Tegumen proportionally shorter than in other Galápagos species, with tegumen-uncus-gnathos barely reaching two-thirds of valvae; lateroposterior extensions more heavily sclerotized distally, with ventral recurved process and small terminal, forked hook; dorsally with well-sclerotized hump just below apex of gnathos. Uncus-socii lobes low, more or less flat, with fewer stiff setae than other species. Gnathos distal process tooth-like or beak-like. Valva with ventral margin sinuate, dorsal margin evenly upcurved, distal portion spatulate or paddle-like, with zone of short, stubby peg-like setae which have a heavier aspect than in previous species (S. galapagensis, S. ancystra, S. roquealbeloi, S. forcipata). Phallus markedly arched, distal portion with large bifurcation, ventral arm of bifurcation heavier than dorsal arm, apically pointed, dorsal arm shorter, curved and with minute apical hook; phallotreme wide.

  • Female (n=17) (Figs 24, 25). Head and thorax as in male; forewing pattern dorsally as in male, usually with orange-brown scales placed as in male; forewing underside with white scales along costa as part of retinaculum and beyond, until middle of wing; forewing length: 3.5-4.5 mm, mean 4.0 ± 0.3, n=17; hindwing underside with ivory white along costa until about 1/3 wing length, dark brown on costal 1/3 from beyond base to apex, whitish grey on rest of wing; frenulum with 3 bristles. Abdomen dorsally greyish brown at base, sometimes gradually becoming darker, sometimes to blackish brown, ultimate scaled segment sometimes with some scales basally paler brown or all scales paler brown; ventrally all ivory white.

  • Female abdomen and genitalia (3 preparations) (Figs 117, 130, 145). Tergum VI narrowly transverse, half the width of tergum V, anterior margin a thick rim, posterior margin evenly concave. Sternum VI transversely rectangular with thick anterior rim, posterior margin nearly straight. Tergum VII slightly trapezoid, anterior margin with thickly rimmed sides not noticeably expanded over pleural region. Sternum VII subcordiform in outline, anterolateral angles with short extensions recessed into pockets around spiracles, posterior margin broadly concave, thickly rimmed, smooth zone restricted to narrow crescent; medioposterior lobe a thickly sclerotized plate subquadrate in outline, with concave surface and set vertically under margin of sternum VII (perpendicular to surface of sterna VII-VIII). Tergum VIII similar to that of S. galapagensis. Sternum VIII shorter than sternum VII, sinus vaginalis with oblique wrinkles laterally, medial surface concave with spiculate zone limited to posterior margin; sclerotization around ostium bursae forming half-ring anteriorly; ostium bursae large, very distinct, oriented ventrally; antrum very short, indistinct. Ovipositor similar to that of S. galapagensis.

  • Biology: One specimen was reared from a pod of “Acacia insulae-jacobi” while another was reared form a larva on “Waltheria ovata”. Now known as Acacia insulae-iacobi (L. Riley) Seigler & Ebinger (Fabaceae), the former plant is native to the Galápagos where it is reported from the islands of San Cristóbal, Santa Cruz and Santiago at a wide range of elevations (Galápagos Species Database 2024). Waltheria ovata Cav. is a shrub of the family Malvaceae; it is native to the Galápagos islands where it is found on Santa Fé and all of the larger islands (Galápagos Species Database 2024). The wide taxonomic discrepancy in host plants is suspicious and it is possible that the larva found on Waltheria ovata may not have developed on that plant but found there incidentally (there is no note on whether the larva was feeding). However, polyphagy has been observed in the sand-dune dwelling Areniscythris brachypteris Powell, including such diverse plants as Asteraceae, Fabaceae, Hydrophyllaceae, and Lamiaceae at the same site, as long as the plants grew in conditions suitable for the larvae (Powell, 1976).

  • Distribution: This new species is presumably endemic to the Galápagos, where it is found on the islands of Baltra, Fernandina, Isabela, Pinta, San Cristóbal, Santa Cruz, Santa Fé, and Seymour Norte.

  • Figs 144-147.

    Female sternum VIII/external genitalia of Galápagos Scythris; images obtained with DIC illumination. (144) S. roquealbeloi, slide MIC 8719. (145) S. furculata, slide MIC 8755. (146) S. cristobalensis, slide MIC 8720. (147) S. lisae, slide JFL 1811.

    img-z43-1_147.jpg

    Figs. 148-149.

    Detail of microsculpture of male valva and female ovipositor, obtained with DIC illumination. (148) Scythris pistillata, slide JFL 1812; enlarged view at 200x (center) and 400x (right) of distal portion of male valva showing granular microsculpture. (149) S. isaferna, slide MIC 8713; enlarged view (400x) of intersegmental membrane SVIII-SIX (ovipositor) showing transverse rows of microspinules.

    img-z44-1_147.jpg

    Fig. 150.

    Maximum-likelihood phylogram from an unpartitioned analysis of the barcode region of COI of Scythris from the Galápagos Islands. Representatives of the type species S. limbella from North America were used as an outgroup. SH-aLRT values / ultrafast bootstrap values to the left of the corresponding node. Node colours: green, SH-aLRT ≥ 80 and UFB ≥ 95; yellow, either SH-aLRT ≥ 80 or UFB ≥ 95; red, SH-aLRT < 80 and UFB < 95. Branch terminal labels include BOLD process Ids, species, and BOLD sample Ids. Red labels indicate holotypes. Details of BOLD:AAB0053 in triangle in Figs 151-152.

    img-z46-1_147.jpg

    Scythris lisae J.-F. Landry & Bucheli, sp. nov.
    Figs 34, 41, 59, 68, 71, 90, 99, 108, 118, 135, 147

  • Material examined

  • Holotype: ♀. 1- “ECU[ADOR], Galápagos, Isabela | NE slope Alcedo, GPS: elev[ation]. | 292 m, S 00° 23.829' W 91° | 01.957', 30.iii.2004, u[ltra]v[iolet]l[ight], | leg[it]. B. Landry, P. Schmitz”. 2- “Database # | CNCLEP | 00005952”. 3- “Barcodes of Life Project | University of Guelph | DNA # jflandry1389”. 4- “genitalia slide ♂ | JFL 1811”. 5- “HOLOTYPE | Scythris | lisae | Landry & Bucheli”. Deposited in MHNG.

  • Paratypes: 3 ♂, 1 ♀ from the Galápagos Island of Isabela: 1 ♂ (dissected, slide JFL 1626), NE slope Alcedo, near pega-pega camp, GPS: elev[ation]. 483 m, S 00°24.029' W 91°02.895', 31.iii.2004, u[ltra]v[iolet] l[ight] (B. Landry, P. Schmitz) (CNCLEP00005979); 1 ♂(dissected, slide JFL 1810), V[olcán]. Alcedo, Pega-Pega [camp], U[ltra]V[iolet]-F[luoresent]L[ight]., 14.x.1999 (L. Roque) (CNCLEP00009433); 1 ♂ (dissected, slide JFL 1824), 1 ♀ (dissected, slide JFL 1806), V. Alcedo, Zona Arida alta, UV-FL, 13.x.1999 (L. Roque) (CNCLEP00009430, CNCLEP00009431). Deposited in CDRS and MHNG.

  • Etymology: The species is dedicated to biologist Lisa Bartels, colleague of JFL at the CNC, for her outstanding technical assistance with this project and many others.

  • Diagnosis: In male genitalia, the lateroposterior extensions of the tegumen are asymmetrical, bent leftward (best seen in apical view), heavily sclerotized, and with the apex bearing a dense cluster of short spines; the phallus is proportionally short, sickle-shaped, with a dorsal, slit-like phallotreme; the valva has a dilated basal half, and a narrow, finger-like distal half with short, dense, peg-like setae.

  • In female genitalia, the surface of sternum VIII is entirely sclerotized and forms a complete subrectangular plate around the ostium bursae, its basal two-thirds bearing transverse wrinkles laterally; the posterior margin of sternum VII is widely excavated and encircling the base of sternum VIII, its medioposterior lobe is prominent, axe-like but thinly sclerotized; the antrum is darkly melanized and extended into a long colliculum. The female genitalia of S. lisae have the most sclerotized genital segments (VI-VIII) of the Galápagos species, with the most prominent ostium bursae and antrum. It is the only species with a fully sclerotized sternum VIII. The female frenulum has two bristles in contrast to the three bristles in all other species (exceptionally two in a few specimens of S. isaferna and one specimen of S. roquealbeloi).

  • Description: Male (n=3). Head: occiput black with blue shine, with white, brown, and black tufts of scales projecting dorsomedially from behind antennae; white scaled on eye margin, dark blackish brown on frontoclypeus, with a few white scales laterally and on ventral margin. Antenna blackish brown, with few white scales on basal segments, e.g., apex of scape; pecten incomplete (1-4 scales) on available specimens, pale brown. Labial palpus white on first palpomere, base and tip of second, and sometimes dorsally on third, otherwise dark brown. Haustellum white scaled. Thorax dorsally mostly dark brown, with few paler brown scales on patagia and sometimes at base of pronotum. Wings as in female (Fig. 34); forewing upperside with purplish shine towards apex, with pattern more or less contrasted; forewing underside dark brown, with broad white band along costa until middle of wing; hindwing underside mostly pale greyish brown, with dark brown band along costa from beyond base until apex. Forewing length: 3.3-3.8 mm, mean 3.7 ± 0.2, n=3. Foreleg coxa mostly white, brown medially; femur mostly brown, with white scales at base and scattered laterally; tibia and tarsus dark brown. Midleg coxa white; femur white at base, dark brown towards apex; tibia dark brown with three white to cream stripes submedially, postmedially, and apically; tarsomeres I and II blackish brown with white at base and apex of 1st, and apex of 2nd; tarsomeres III-V dark brown. Hindleg coxa creamy white; femur satiny white with dark brown at apex; tibia dark brown with white at base, subbasally, medially, apically, ventrally and on spurs, with dorsal crest of hair-like scales white to brown, with white hair-like scales ventrally as well, before median spurs; tarsomeres as midleg, with slightly paler scales at apices of 3rd and 4th. Abdomen dorsally dark brown, with white to greyish brown on first two tergites; ventrally mostly white, with orange brown on antepenultimate sternite.

  • Male abdomen and genitalia (3 preparations) (Figs 41, 59, 68, 71, 90, 99, 108). Tergum VIII with deep, evenly concave posterior emargination, lateroposterior corners angulate. Sternum VIII more or less flat, with shallow and broad anteromedian emargination, medioposterior projection conical with evenly curved sides, short apex barely reaching basal half of valvae. Tegumen-uncus-gnathos complex elongate, only slightly arched, extended beyond apex of valvae; medioposterior extensions heavily sclerotized distally, apex asymmetrical, skewed to the left, armed with dense, short spines and teeth. Uncus-socii lobes elongate but flat, not dorsally prominent. Gnathos distal process beak-like, downcurved. Valva with dilated basal half, dorsal margin curved with small bump beyond middle, ventral margin shallowly sinuate, distal half much narrower than basal half, zone of peg-like setae covering dorsoapical portion, pegs short and stubby. Phallus proportionally short, shortest of all species, arched, sickle-shaped, with heavy widened base, medio-dorsally with small tooth, distal half attenuate, phallotreme slit-like and on dorsal side of phallus.

  • Female (n=2) (Fig. 34). Head and thorax as in male; forewing length: 3.8-4.2 mm, mean 4.0 ± 0.3, n=2 (holotype: 3.7 mm); forewing upperside and underside with pattern as in male; hindwing frenulum with 2 bristles. Abdomen dorsally greyish brown on basal sternite, darker blackish brown on 2nd, blackish brown on next tergites, with apical row of short white scales on ultimate scaled tergite; ventrally all white.

  • Female abdomen and genitalia (2 preparations) (Figs 118, 135, 147). Tergum VI narrowly transverse, sides ventrally recurved around pleural region and surrounding spiracles which are protruded/ humped. Sternum VI also narrowly transverse, anterior margin more or less straight, posterior margin slightly concave, thickened, overlapping anterior margin of sternum VII. Tergum VII transversely subrectangular, 2x length of tergum VI, posterior margin overlapping basal third of tergum VIII. Sternum VII transverse, heavily sclerotized, sides bent and expanded dorsally to form half-cylinder around base of sternum VIII; anterior margin medially with smooth, semi-elliptic area partially recessed under posterior margin of sternum VI; posterior margin thick, beaded at right angle with plate surface, posteromedial lobe spatulate, proportionally longer than in all other Galápagos species but more membranous (or less sclerotized; it stains readily with chlorazol black). Tergum VIII about as long as tergum VII, with lateroposterior portion of marginosclerite expanded, broad. Sternum VIII markedly sclerotized, elongate-subquadrate, lateroposterior angles rounded, basal two-thirds with transverse wrinkles/creases laterally, with a few scattered setae, posterior margin medially very finely spinulate. Ostium bursae situated anteromedially on smooth hump, wide, prominent; antrum/colliculum darkly sclerotized, forming tube extended inwardly at steep angle, as long as sternum VII, anteriorly tapered to membranous ductus bursae. Ovipositor similar to that of other Galápagos species.

  • Biology: Unknown except that the moths are attracted to light and were collected at mid elevations, between 200 (upper arid zone) and 483 m in elevation.

  • Distribution: Presumably endemic to the Galápagos, this species has been found only on Isabela Island, on the slope of Alcedo Volcano.

  • Fig. 151.

    Continued from Fig. 150, BOLD: A A B 0 0 5 3 . Details of triangle-clade (a) on right side, details of triangle-clade (b) in Fig. 152.

    img-z47-2_147.jpg

    Fig. 152.

    Continued from Fig. 151, BOLD:AAB0053, details of clade (b).

    img-z49-1_147.jpg

    Scythris cristobalensis
    J.-F. Landry & B. Landry, sp. nov.

    Figs 27, 28, 116, 134, 146

  • Material examined

  • Holotype: ♀. 1- “ECU[ADOR], Galápagos, San Cristóbal | near Loberia, GPS: elev[ation]. 14 m | S 00° 55.149' W 89° 36.897' | 16.iii.2004, u[ltra]v[iolet] l[ight] | leg[it]. B. Landry, P. Schmitz”. 2- “Database # | CNC LEP | 00005966”. 3- “Barcodes of Life Project | University of Guelph | DNA # jflandry1390”. 4-“genitalia slide ♀ | SRB 215”. 5- “HOLOTYPE ♂ | Scythris | cristobalensis | J.-F. Landry & B. Landry”. Deposited in MHNG.

  • Paratypes: 3 ♀ with same data as holotype, except without a ‘jflandry’ DNA number: CNCLEP00005963, CNCLEP00005964 (genitalia slide MIC 8720), CNCLEP00005965 (genitalia slide JFL 1803). Deposited in CNC and MHNG.

  • Etymology: The new name refers to the occurrence of the species only on the island of San Cristóbal.

  • Diagnosis: In external characters, especially forewing pattern, this species is confusingly similar to most of its Galápagos congeners except S. falcata, but differs by its smaller size (FW length 3.7-4.0 mm). In female genitalia, the sclerotization of sternum VIII is suborbicular and extended around the perimeter of the sternum, encircling the sinus vaginalis and ostium bursae except for a narrow gap medioposteriorly; the posterior margin of sternum VII is slightly sinuate but not emarginate, the medioposterior lobe is reduced to a thin band, indistinct; the antrum is slightly excavated and has very dense, darkly melanized cuticular folds posterior to the ostium; sternum VI is narrow, transversely subtrapezoid without anterior emargination. The male is unknown.

  • Description: Female (n=4) (Figs 27, 28). Head: occiput dark brown, with bronze shine, sometimes with white scales laterally next to antennae; tufts of scales projecting dorsomedially from behind antennae mostly dark brown, with few white to greyish brown scales; frontoclypeus dark brown, with white scales mostly laterally. Antenna blackish brown, with satiny white on scape ventrally, with row of white scales ventrally on flagellum; pecten with up to about 8 pale brown scales. Labial palpus with 1st segment white, 2nd segment white at base, dark brown toward apex with white dorsally, 3rd segment dark brown with white dorsally. Haustellum scaled satiny white laterally, dark brown medially. Thorax dorsally mostly dark brown, with few pale brown scales laterally at apex of mesothorax and tips of patagia. Wings as shown (Figs 27, 28); forewing upperside varying in contrast of pattern, sometimes with few orange-brown scales along midline and at tornus; forewing underside with white scales along costa as part of retinaculum and beyond middle of wing, darker brown toward apex; hindwing underside with ivory white along costa until about 1/3 wing length, dark brown on costal 1/3 from beyond base to apex, whitish grey on rest of wing; frenulum with 3 bristles. Forewing length: 3.7-4.0, mean 3.8 ± 0.1, n=4 (holotype: 3.8 mm). Foreleg coxa satiny white with dark brown; femur dark brown with white at base and along ventral margin; tibia and tarsus dark brown with white to greyish brown at apex of tibia and tarsomeres I and II. Midleg coxa satiny white with some dark brown; femur white except blackish brown apex; tibia blackish brown with three satiny white to ivory stripes submedially (faint), postmedially, and apically; tarsomeres I and II blackish brown with white at base and apex of 1st, and apex of 2nd, ivory white medially on 1st; tarsomeres III-V dark brown with slightly paler median surfaces. Hindleg coxa satiny white and greyish brown; femur satiny white except for blackish brown apex; tibia mostly white with blackish brown spots subbasally, at 1/3, and subapically, with dorsal crest of hair-like scales white to pale brown dorsally, more homogenously white ventrally; tarsomere I blackish brown with white at base and apically; tarsomeres II-IV dark brown with few white to pale greyish brown scales apically; tarsomere V dark brown without paler apical scales. Abdomen dorsally greyish brown, darker from 2nd tergite to apex; ventrally ivory white.

  • Female abdomen and genitalia (3 preparations) (Figs 116, 134, 146). Tergum VI evenly and narrowly transverse, sides ventrally recurved around pleural region, posteroventral angles expanded around spiracles and ending in folds under sides of sternum VI, folds covered with firmly set scales in tuft-like arrangements. Sternum VI narrowly and transversely subtrapezoid with straight anterior and posterior margins. Tergum VII subrectangular, 2x length of tergum VI. Sternum VII subrectangular, 2x length of sternum VI, anterior margin beaded; posterior margin slightly sinuate, thickly rimmed and beaded, recessed; smooth zone narrow, crescentic; posteromedial lobe reduced to thin band. Tergum VIII proportionally broader than S. galapagensis and most other species (except S. lisae); marginopleural sclerite anteriorly angulate, posteriorly widened. Sternum VIII suborbicular in outline, lateral portions sclerotized with few cuticular wrinkles or grooves laterally, encircling sinus vaginalis and ostium bursae. Ostium bursae small but distinct, on a slightly excavated antrum with very dense, darkly melanized cuticular folds posterior to ostium which seem to be part of the sternal wall (or closely appressed to it). Ovipositor 2.6x length of sternum VIII, with fine transverse microsculpture.

  • Biology: Unknown except for the collecting site situated near the seashore and the moths being attracted to ultraviolet light.

  • Distribution: Presumably endemic to the Galápagos, this species is possibly also restricted to San Cristóbal Island, the oldest of the emergent islands of the archipelago.

  • Remarks: The male is yet unknown.

  • DISCUSSION

    The group of ten Scythris species described here are considered to constitute a monophyletic clade. It is well supported by morphological traits and DNA barcodes. Evidence of monophyly of the Galápagos Scythris is the putatively autapomorphic modification of the posterolateral walls of the tegumen which are extended caudally beyond the apex of the fused uncus-socii-gnathos complex. The gnathos itself is reduced and fused to both the uncus-socii and the tegumen extensions, and appears to sit on top of the latter. This configuration appears to be unique within the family as far as is known. Monophyly suggests that the Galápagos Scythris may have diversified from a single founding ancestor. As such, they would represent the second-largest radiation known in the Lepidoptera of the Galápagos, following that of Galagete Landry (Autostichidae) with 12 species (Schmitz et al., 2007), and only the fourth recognised such radiation if smaller ones are considered, that is Utetheisa Hübner (Erebidae: Arctiinae) (Roque-Albelo & Landry, 2009) and La Błeszynski (Crambidae: Crambinae) (Landry & Léger, 2024), with five species each. Monophyly of the Galápagos Scythris is also supported by the molecular evidence, forming a single, well-supported clade. It is notable that this clade was similarly recovered by Nupponen & Sihvonen (2022) with their analysis of New World Scythrididae covering a much greater taxon sample.

    DNA barcodes were useful in detecting one species (S. cristobalensis) which was originally missed across a series of similar-looking specimens from the same collecting event. Despite its confusing superficial similarity with some S. furculata specimens, S. cristobalensis is very distinct in female genitalia. Thus, characterisation of its COI sequence led to its diagnostic morphological features being subsequently recognised.

    Low barcode divergences may be indicative of relatively recent divergence. Significant barcode divergence has been generally effective to identify Lepidoptera species as well as in detecting cryptic species in complexes of confusingly uniform groups (for ex.: Mutanen et al., 2014; Huemer & Mutanen, 2015; Sohn et al., 2015; Kozlov et al., 2016; Tabell et al., 2018; Landry et al., 2023, Kaila & Huemer, 2024), or for exploring unknown faunas (for ex., Lees et al., 2013; Mutanen et al., 2013). Such results were achieved by combining morphological and other evidence with DNA barcode data in a process of reciprocal illumination. However, exceptions have been found in which well-supported complexes of closely related species recognised on the basis of extensive morphological and biological evidence had uninformative barcodes (Kaila & Ståhls, 2006; Schmidt & Sperling, 2008), heeding warnings and caveats against using barcodes as sole or primary source of evidence for taxonomic decisions (Wahlberg et al., 2003; Mutanen et al., 2016; Phillips et al., 2019). In such situations, increased sampling and investigating other lines of genetic evidence focusing on nuclear genes have provided a useful complement to morphological and biological evidence. Unfortunately, data from nuclear genes were not available for the present study, and are something to be pursued in future investigations. Distributional patterns (Table 2) generally do not reflect single-island endemicity with seven species occurring on two or more islands. It is possible that the apparent geographical restriction of some species reflects incomplete sampling. In Galagete (Autostichidae), the other genus of microlepidoptera to have undergone a significant radiation on the Galápagos from a single ancestor, the distribution pattern of species was not correlated with the age of individual islands but was hypothesised to have resulted from stochastic colonisation (Schmitz et al., 2007).

    The recent descriptions of 23 new species of Scythrididae from South America (Nupponen & Sihvonen, 2022; Heppner, 2010a) represented an increase of 65% from historically described taxa for that region. The work of Nupponen & Sihvonen (2022) revealed several species with highly derived or modified genitalia which currently defy clear classification, in addition to a few species (S. fluvialis Meyrick, S. directiphallella group) that shared a generalised male genitalia configuration partially similar to the Galápagos Scythris (sternum VIII, overall symmetry), although all possess a gnathos separate from the uncus-socii with both structures making the most caudal part of the genitalia, and a distally unmodified tegumen. The male genitalia of the S. directiphallella group were noted to resemble those of the Afrotropical genus Haploscythris Viette in some respects (bilobed uncus, divided valva, V-shaped anterior margin of vinculum), but data are insufficient to infer possible relationship (Nupponen & Sihvonen, 2022). It is possible that these character states are homoplasious or represent groundplan conditions at the family level.

    It must be underlined that phylogenetic relationships and generic delimitation in the Scythrididae remain very poorly understood. Phylogenetic and biogeographic inferences are further impeded by the large amount of undescribed diversity which has been detected in existing collections and awaits study (Landry, 1991). Relationships of the Galápagos species with taxa from continental South America is a possible hypothesis, although one currently not supported by any evidence. Dispersalist models have traditionally been invoked for the origin of Galápagos biota (Heads & Grehan, 2021). However, several similar biogeographic patterns of Galápagos endemics involving various organisms (insects, spiders, mites, amphipods, crustaceans, polychaete worms, birds, fishes, plants) have been shown to be coincident with complex tectonic patterns, with vicariance of metapopulations of widespread ancestors providing a more convincing hypothesis of origin (Heads & Grehan, 2021). Insights into the relationships of the Galápagos Scythrididae will require more studies of Neotropical species involving not only South America but also Central America and the islands of the Caribbean Plateau, as well as deeper phylogenetic analyses.

    ACKNOWLEDGEMENTS

    We thank Stewart B. Peck, retired from Carleton University, Ottawa, Canada, for organizing the collecting permits and the 1989 and 1992 field trips which BL was invited to join; this was made possible with financial support from the Natural Sciences and Engineering Research Council of Canada for field research on arthropod evolution. BL is grateful to the authorities and staff of the Galápagos National Park Directorate and the Charles Darwin Research Station for allowing field work, for providing the necessary permits, and for logistical support. The help of Lazaro Roque Albelo, in particular, was greatly appreciated during BL's and Patrick Schmitz's collecting efforts between 2002 and 2006, including for companionship in the field, hospitality, and for contributing many specimens to this study. In addition, we are thankful to Stewart Peck, as well as Novarino Castillo, Joyce Cook, Charlotte Causton, Moraima Inca, José Loaiza, Ricardo Palma, Patrick Schmitz, Brad Sinclair, and Eduardo Vilema for field companionship and help to BL. For the loan of material in their care we thank Norm Penny and Paul Arnaud Jr. at the CAS, as well as Klaus Sattler and David Lees at the NHMUK. For financing his visit to the NHMUK in 2000, BL thanks the Galápagos Conservation Trust, London, U.K. The City of Geneva, through its Muséum d'histoire naturelle, is acknowledged for funding BL's collecting expeditions to the Galápagos in the early 2000s. At the CNC, JFL thanks Lisa Bartels for her technical assistance in barcoding preparations, data checking, specimen photography, image editing, and plate preparation; Ahmed Badiss and Andrea Brauner assisted with specimen databasing; Adam Brunke advised and assisted with the molecular analysis.

    SRB is grateful to the Department of Entomology at The Ohio State University for funds provided to travel to the CNC in February of 2005. A portion of this research was paid for by the National Science Foundation Grant #0415061.

    JFL is indebted to Paul Hebert and the team at the Center for Biodiversity Genomics, University of Guelph, Ontario, Canada for their generous assistance in the production of DNA barcodes. Funding for DNA barcoding and sequence analysis was partly provided by the Government of Canada through the National Science and Engineering Research Council, Genome Canada, the Ontario Genomics Institute in support of the International Barcode of Life project, and Agriculture & Agri-Food Canada. Finally, we are grateful to Doug Yanega, Department of Entomology, University of California at Riverside and commissioner on the International Commission on Zoological Nomenclature, for his help in clarifying the nomenclatural status of taxonomic names in thesis dissertations. Lauri Kaila and Oleksiy Bidzilya made constructive comments on the manuscript.

    This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited (see  https://creativecommons.org/licenses/by/4.0/).

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