New species of the Brachyglutine genus Panabachia Park (Coleoptera, Staphylinidae, Pselaphinae) from Ecuador

225 New species of the Brachyglutine genus Panabachia Park (Coleoptera, Staphylinidae, Pselaphinae) from Ecuador Sofia I. Muñoz-Tobar1, Michael S. Caterino1,2 1 Instituto Nacional de Biodiversidad, Quito, Ecuador 2 Department of Plant & Environmental Sciences, Clemson University, Clemson, SC 29634, USA Corresponding author: Michael S. Caterino ([email protected]) Copyright: © Sofia I. Muñoz-Tobar & Michael S. Caterino. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract Twenty-two new species of Panabachia Park (Coleoptera: Staphylinidae: Pselaphinae: Brachyglutini: Brachyglutina), are described from montane forest and páramo habitats of Ecuador: Panabachia pahuma sp. nov., P. trifecta sp. nov., P. inornata sp. nov., P. amica sp. nov., P. winku sp. nov., P. ayauma sp. nov., P. pastazae sp. nov., P. romeroi sp. nov., P. uktu sp. nov., P. salebrosa sp. nov., P. urbana sp. nov., P. carltoni sp. nov., P. falini sp. nov., P. papallacta sp. nov., P. ananay sp. nov., P. cayambi sp. nov., P. cryptica sp. nov., P. caranqui sp. nov., P. patera sp. nov., P. vigilans sp. nov., P. perdita sp. nov., and P. ambulans sp. nov. Only two species of the genus have been previously described, from Panamanian and Guatemalan lowlands. But, based on unpublished records, the genus is now known to occur from Mexico to Bolivia, and includes several additional undescribed species. These new species exhibit important variability in the diversity in male secondary sexual characters. Images of the habitus, key characters and aedeagus are provided for the new species. Resumen Veintidós nuevas especies de Panabachia Park (Coleoptera: Staphylinidae: Pselaphinae: Brachyglutini: Brachyglutina), de bosque montano y páramo Ecuatoriano, son descritas: Panabachia pahuma sp. nov., P. trifecta sp. nov., P. inornata sp. nov., P. amica sp. nov., P. winku sp. nov., P. ayauma sp. nov., P. pastazae sp. nov., P. romeroi sp. nov., P. uktu sp. nov., P. salebrosa sp. nov., P. urbana sp. nov., P. carltoni sp. nov., P. falini sp. nov., P. papallacta sp. nov., P. ananay sp. nov., P. cayambi sp. nov., P. cryptica sp. nov., P. caranqui sp. nov., P. patera sp. nov., P. vigilans sp. nov., P. perdita sp. nov. y P. ambulans sp. nov. Previamente, solo se habían descrito dos especies, de tierras bajas de Panamá y Guatemala. Sin embargo, el género se registra desde México hasta Bolivia, con varias especies sin describir. Estas nuevas especies presentan importante variabilidad en los caracteres secundarios del macho. En este estudio se proporcionan fotografías del hábito, caracteres claves y del edeago de las nuevas especies. Key words: Ecuador, montane forest, new species, Panabachia, páramo, Staphylinidae Academic editor: Jan Klimaszewski Received: 8 May 2025 Accepted: 26 August 2025 Published: 2 October 2025 ZooBank: https://zoobank. org/09AF0C44-B9DF-473F-977DB00123D955CF Citation: Muñoz-Tobar SI, Caterino MS (2025) New species of the Brachyglutine genus Panabachia Park (Coleoptera, Staphylinidae, Pselaphinae) from Ecuador. ZooKeys 1254: 225–261. https://doi. org/10.3897/zookeys.1254.158319 ZooKeys 1254: 225–261 (2025) DOI: 10.3897/zookeys.1254.158319 226 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia Introduction The neotropical fauna of the rove beetle subfamily Pselaphinae is an incredibly diverse one. With more than 1600 known species (Asenjo et al. 2019) it ranks third in neotropical species richness among subfamilies of Staphylinidae, a family that represents one of, if not the largest family of animals on Earth. Yet these described species represent only the tip of an unfathomably large iceberg, with numerous hyperdiverse countries almost completely unknown. Ecuador, for example, hosts only 16 named species of Pselaphinae (unpublished data), while the much smaller but better-surveyed country Panama is known to host more than 500 (Chandler 1992). Neighboring Colombia is known to have at least 88 species of Pselaphinae (Newton et al. 2005). But surveys of even a single site in lowland Ecuador turned up 178 Pselaphine species, while estimating at least a third more (Carlton et al. 2004). At present, all that can be confidently said about pselaphine diversity in any particular neotropical area is that it is undoubtedly much greater than known. In this paper we explore the diversity in a single pselaphine genus, Panabachia, in a single country, Ecuador. Panabachia was named by Park (1942) for the Panamanian species Bryaxis vulnerata Sharp, based on the unique excavations of the pronotum, distinguishing it from the (purportedly) related genus Reichenbachia. Park did not immediately note that Sharp had suggested a close relationship with his Bryaxis impressicollis, described in the same paper with similar pronotal modifications. But he later moved this species, described from Guatemala, into Panabachia as well (Park 1945). Since that time no additional species have been named or moved to Panabachia, and the genus has only been mentioned in catalogs and checklists. Chandler (in Navarrete-Heredia et al. 2002) gave the distribution of the genus, based on still-undescribed species, as Mexico to Bolivia. Otherwise, little new data has been presented on the genus in many decades. In 2020, as part of a larger study of beetle diversity in high elevation páramo habitats in Ecuador (e.g. Muñoz-Tobar 2019; Muñoz-Tobar and Caterino 2019), Muñoz-Tobar and Caterino (2020) explored the potential species diversity in numerous newly discovered populations of Panabachia beetles. Multilocus sequencing and preliminary morphological study suggested as many as 17 different species, revealing a high-elevation radiation in this poorly known group. Further recent collecting at other Páramo sites as well as some in higher montane forest have revealed additional species. Although it is evident that the species we have found represent only the beginning of a substantial and diverse fauna, it will be valuable to begin to document some of the morphological breadth, as well as call attention to this largely high elevation-restricted diversification. Therefore, below we describe 22 new species of Panabachia for which species status is well-supported and for which males are available. Materials and methods Field sites and sampling Specimens used in this study were obtained mainly from leaf litter samples, from 13 sites across cloud forest and páramo of Ecuador (Fig. 1). The sampling of the sites was part of three research projects focused on describing beetle 227 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia diversity in Ecuador. Collecting permits are the following: 0020-FAU-MA-DPOPNY, MAE-DNG-ARGG-CM-2014-004, and MAATE-DBI-CM--2022-0255. Additional samples were obtained from deposited material at the Instituto Nacional de Biodiversidad/Museo Ecuatoriano de Ciencias Naturales (MECN-EN), Museo de Zoología de la Pontificia Universidad Católica del Ecuador Invertebrados (QCAZ-I), Museo de Zoología de la Universidad San Francisco de Quito (ZSFQ), and Louisiana State Arthropod Museum (LSAM). Morphology was studied using an Olympus stereo-microscope (Olympus SZ61R), genitalia was examined using temporary slide mounts in glycerin, with the use of compound microcopy, as well as photographic images taken with a Macropod Pro 3D (Macroscopic Solutions, CT, USA). The majority of specimens were cleared during DNA extractions, with the use of tissue digestion buffers and proteinase K. The aedeagus was extracted through the abdominal apex, after enzymatic digestion. All extracted specimens were mounted, labelled with extraction numbers, and deposited in public repositories as detailed under ‘type material’ sections below. Measurements Measurements were taken using photographic material generated with a digital camera (The Imaging Source DFK23UX236) adapted to the stereoscope and analyzed in ImageJ (Schneider et al. 2012). At least three individuals per species, females and males, were measured when available (see Table 1). Head length (HL) was measured from the clypeal margin to the upper anterior edge of the neck constriction (ignoring the neck); pronotal length (PnL) was measured along the midline; pronotal width (PnW) was the maximum width, near the midline; elytral length (EL) was measured along the suture from the base of the scutellum to the apex Figure 1. A. Map of the sites where Panabachia were collected in the Ecuadorian Andes. Examples of habitat of Panabachia species: B. Grassland and shrub páramo in Atillo, province of Chimborazo and C Montane forest in El Pahuma Orchid Reserve, province of Pichincha. 228 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia of the suture; elytral width (EW) was the maximum width, total abdomen length (AL) was measured laterally in a straight line from the base of the 1st ventrite to the apex of the last tergite (ignoring telescopy and/or curvature); total length (TL) was calculated as head length + pronotum length + elytral length + abdomen length. Taxonomic account Checklist of the new species (informally grouped by distinctive characters): Short flat median lobe: 1. P. pahuma sp. nov. 2. P. trifecta sp. nov. 3. P. inornata sp. nov. 4. P. amica sp. nov. 5. P. winku sp. nov. 6. P. ayauma sp. nov. 7. P. pastazae sp. nov. 8. P. romeroi sp. nov. 9. P. uktu sp. nov. 10. P. salebrosa sp. nov. 11. P. urbana sp. nov. 12. P. carltoni sp. nov. 13. P. falini sp. nov. Longer, curved median lobe: 14. P. papallacta sp. nov. 15. P. ananay sp. nov. 16. P. cayambi sp. nov. 17. P. cryptica sp. nov. 18. P. caranqui sp. nov. 19. P. patera sp. nov. Table 1. Average (where n > 1) measurements taken in mm, for important body dimensions. nHL PnL PnW EL EW AL TL P. pahuma 10.21 0.31 0.34 0.50 0.33 0.42 1.44 P. trifecta 10.35 0.35 0.37 0.58 0.47 0.30 1.58 P. inornata 3 0.32 0.30 0.38 0.55 0.42 0.45 1.62 P. amica 10.27 0.33 0.46 0.62 0.49 0.42 1.64 P. winku 3 0.22 0.35 0.38 0.58 0.37 0.45 1.59 P. ayauma 3 0.24 0.30 0.37 0.53 0.39 0.36 1.38 P. pastazae 10.24 0.27 0.28 0.40 0.32 0.21 1.12 P. romeroi 3 0.24 0.31 0.31 0.48 0.36 0.33 1.36 P. uktu 4 0.26 0.35 0.37 0.58 0.41 0.38 1.52 P. salebrosa 3 0.27 0.37 0.34 0.62 0.39 0.49 1.63 P. urbana 3 0.20 0.34 0.37 0.47 0.39 0.47 1.49 P. carltoni 20.25 0.30 0.32 0.43 0.33 0.37 1.34 P. falini 20.24 0.35 0.32 0.47 0.35 0.59 1.64 P. papallacta 20.28 0.30 0.41 0.57 0.31 0.44 1.59 P. ananay 3 0.28 0.31 0.34 0.57 0.38 0.41 1.57 P. cayambi 3 0.27 0.28 0.37 0.57 0.39 0.41 1.53 P. cryptica 20.28 0.29 0.33 0.60 0.42 0.40 1.56 P. caranqui 10.28 0.30 0.37 0.60 0.39 0.45 1.63 P. patera 20.26 0.36 0.31 0.48 0.38 0.42 1.52 P. vigilans 10.23 0.38 0.30 0.43 0.34 0.61 1.65 P. perdita 10.21 0.30 0.31 0.49 0.38 0.48 1.48 P. ambulans 10.23 0.40 0.47 0.60 0.38 0.65 1.88 229 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia Genitalia outliers: 20. P. vigilans sp. nov. 21. P. perdita sp. nov. 22. P. ambulans sp. nov. Key to the species of Panabachia (males only) 1 With pronotal modifications including setose pits, depressions, tubercles, or horns; if pronotum unmodified, then ultimate abdominal ventrites slightly depressed or with setose margins (males) ....................................2 – Pronotum lacking modifications and terminal abdominal ventrites simply convex (females) ................................................................not keyed further 2 Pronotum lacking secondary modifications ................................................3 – Pronotum with distinct secondary modifications .......................................7 3 Pronotum widest, angulate in basal 1/2 (Fig. 7G) ......................... P. perdita – Pronotum widest in distal 1/2, sides rounded .............................................4 4 Male flightless (humeri evenly sloped to base); protarsi modified, widened and scaled (Fig. 9); known only from Atillo (Chimborazo Province) ............ ......................................................................................................P. ambulans – Male winged (humeri roundly subquadrate), known from the northern Ecuadorian Andes (Pichincha and Imbabura) .................................................5 5 Dorsum rather densely and uniformly setose; known from Pichincha ........ ...........................................................................................................P. urbana – Parts of dorsum, particularly head and pronotum, sparsely setose; known from Imbabura ...............................................................................................6 6 Male last sternite (Fig. 5O) transverse, convex, sparsely setose, with dense row of 6 flattened setae at middle of apical margin; male apical tergite (Fig. 5N) transverse, slightly depressed in the middle ................P. caranqui – Male last sternite transverse (Fig. 5K, L), setose, with a forked process along the inner basal margin, apical margin sinuous, with denser setae at middle; male apical tergite transverse, apically truncate, setose along posterior margin, slightly emarginate in the middle .................... P. inornata 7 Male pronotum with 1 or closely adjacent pair of median impressions, but lacking secondary anterolateral impressions..............................................8 – Male pronotum with 1 or 2 median impressions in addition to pair of anterolateral discal impressions ....................................................................13 8 Male pronotum with curving median depression and more densely setose regions posterolaterad ..................................................................................9 – Median depression varied in shape, but never with more densely setose regions posterolaterad ................................................................................10 9 Male pronotum with median depression (Fig. 4H) nearly V-shaped, with common median area diverging to anterior corners; pronotum strongly narrowed anterad, nearly with apical collar; densely setose regions at sides larger ..................................................................................... P. romeroi – Male pronotum with median depression (Fig. 4G) transverse, laterally curving only weakly forward; pronotal sides evenly curved anteriorly and posteriorly, without apical collar; densely setose regions at sides of pronotal disk smaller .............................................................................P. pastazae 230 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia 10 Male pronotum (Fig. 4D) widest at base, median depressions located close to posterior margin ..................................................................P. amica – Male pronotum widest near middle, median depressions located in anterior 1/2 of disk ...............................................................................................11 11 Male pronotal sides finely angulate at middle; median depression (Fig. 4M) with paired combs of long, curved setae extending posterad from anterior edge ........................................................................................P. falini – Male pronotal sides rounded; median depression with few or single anterior setae, not with separated pair or clusters of setae ............................12 12 Anteromedian edge of pronotal depression elevated as a median horn (best seen in lateral view); median depression (Fig. 4A) almost subdivided along midline into pair of midlateral depressions ........................P. pahuma – Anteromedian edge of pronotal depression not elevated into median horn; median depression (Fig. 4E) not subdivided at middle, forming a continuous transverse depression................................................................ P. winku 13 Genal carinae well developed behind eyes (Fig. 5J, M) ............................14 – Genae not produced as carinae behind eyes ............................................15 14 Male pronotum with a pair of adjacent median depressions (Fig. 7A), divided by a thin carina, with 3 short distinct rows of setae at the middle and outer edges; posterolateral corners of male pronotum swollen, but not angulate ................................................................................P. papallacta – Male pronotum with larger central median depression (Fig. 7B), with setal patches at lateral corners and divergent tufts on either side of anterior edge; posterolateral corners of male pronotum strongly angulate .............. .......................................................................................................... P. ananay 15 Secondary, anterolateral depressions of male pronotum delimited by distinct, raised carinae .....................................................................................16 – Secondary depressions of male pronotum simple, not delimited by carinae; entire pronotum clearly wider than long ............................................17 16 Secondary depressions of male pronotum (Fig. 4L) small, elongate oval, and located anterolateral to transverse median pronotal depression; pronotum slightly longer than wide, slightly prolonged anteriorly .....P. carltoni – Secondary depressions of male pronotum (Fig. 7F) larger, more nearly circular, and located posterolaterad to more narrowly median pronotal depression .......................................................................................... P. vigilans 17 Median pronotal depression very large, extending onto basal 1/2 of pronotum such that basal median pronotal fovea is within the depression (Figs 4F, 7E) ...........................................................................................................18 – Median pronotal depression more limited in extent, such that the basal median fovea is free, on a convex portion of disk ....................................19 18 Male pronotum (Fig. 7E) wider at midline, sides more or less evenly rounded, broadly longitudinally depressed, with wide, mustache-like comb of setae along anterior edge of depression ......................................P. ayauma – Male pronotum (Fig. 4F) narrow, elongate, subrectangular; median depression nearly as wide as pronotum, anterior edge with small pair of diverging setal clusters .....................................................................................P. patera 19 Male pronotal depression with distinct clusters of setae near its basal midline ..........................................................................................................20 – Male pronotal depression with only sparse setae ....................................21 231 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia 20 Secondary lateral depressions of male pronotum (Fig. 7C) small and located directly at sides of main median depression; entire posterior margin of median pronotal depression fringed with setae ..................... P. cayambi – Secondary lateral depressions of male pronotum (Fig. 7D) located posterolaterad main median depression; posterior margin of median depression with short rows of setae at lateral corners of depression ...P. cryptica 21 Anterior marginal portion of pronotal disk (Fig. 4B) transversely convex, with distinct posterior margin overlapping median and lateral depressions, this swelling relatively densely punctate and setose ......... P. trifecta – Anterior marginal portion of pronotum not modified, lacking distinct posterior marginal ridge and not more densely setose or punctate than rest of pronotal disk ................................................................................................22 22 Male pronotal depression (Fig. 4I) a very small oval present just behind anterior margin, with interior circular carina enclosing small setose region ... ..............................................................................................................P. uktu – Male pronotal depression (Fig. 4J) comprising 2 weakly separated, adjacent, shallow depressions, lacking any carinae ........................P. salebrosa Species descriptions Panabachia Park, 1942 Type species. Bryaxis vulnerata Sharp, 1887, designated by Park (1942). Diagnosis. The genus Panabachia was established principally to recognize the sexually dimorphic male pronotal modification of the type species (Bryaxis vulnerata Sharp, 1887, also once placed in Reichenbachia Raffray 1904]). They are otherwise rather generalized Brachyglutini, with a median longitudinal gular carina, 11-segmented antennae, two basal carinae on the first visible abdominal segment (usually close together and diverging), and subcontiguous mesocoxae. Several new species described herein do not exhibit pronotal dimorphism, but are unambiguously associated by other characters. No male genitalia have ever been described for the genus. The male genitalia among the new species falls into two very distinct classes. One, represented by only two species, has a short broadly rounded aedeagus bearing a transverse basal bridge. The other typically exhibits an elongate median lobe (incorporating a narrow basal bulb), usually thin parameres, and, separately, a highly specialized pair of articulating accessory sclerites in the distal abdominal membrane. Simple forms of these constitute narrow, flat sclerites, but they become elaborated into complex, coiled, and possibly spring-like structures that probably participate in the extrusion/retraction of the genitalia. We cannot be 100% certain which of these major genitalic forms corresponds to the type (P. vulnerata), as its male genitalia has never been illustrated. However, we have dissected a specimen from Costa Rica (LSAM) that appears externally to be very similar to this species (Fig. 2A, B), perhaps conspecific, and its male genitalia represents the more common form, with an elongate median lobe flanked by separate accessory sclerites (Fig. 2C). Among those with distinct accessory sclerites, the male genitalia of the species can be further subdivided into two distinctive forms, one in which the median lobe narrows beyond the basal foramen, and has the apical portion tubular and curving, and one in which the median lobe is shorter and variously flattened beyond 232 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia the basal foramen. These appear to correspond perfectly to the main clades in Muñoz-Tobar and Caterino (2020), discussed further in the discussion section. Panabachia pahuma sp. nov. https://zoobank.org/225651C7-F346-4072-9837-F18C2E5B35F4 Figs 3A, 4A, 5A, 6A Type material. Holotype • ♂ (ZSFQ-i23395): “ECUADOR: Pichincha, 0.0182°N, 78.6372°W, El Pahuma Orchid Res., 28.v-1.vi.2011, FIT, 2200–2400 m. AT1329, M. Caterino, A. Tishechkin” / “Caterino DNA voucher, Ext. MSC-12635, Morphosp. ElP.A.034”; deposited in ZSFQ. Diagnosis. Head (Figs 3A, 5I) broad, slightly shagreened, with rather thick short, appressed white setae; lateral vertexal foveae well developed, median fovea absent; antennal bases slightly elevated, defined posteriorly by fine bent stria at anterolateral margins; eyes protuberant, round, diameter ~2/3 postocular genal width; antennae rather short, antennomere III slightly longer than wide, antennomeres IV–VI beadlike, rounded, VII–IX increasingly transverse, shorter, antennomere X short, crescent-shaped, entirely excavate on inner margin, antennomere XI ~2 × as long as X, slightly concave on inner basal face, continuous with concavity of X, bluntly rounded apically, with distinct, round setose depression on outer apical surface; male pronotum (Fig. 4A) convex in basal 1/2, lacking lateral basal foveae, with distinct but non-setose median basal fovea; anterior 1/2 of pronotum with bilateral depressions, subcontiguous at middle, anterior edge of depression raised into posteriorly directed, anteriorly triangular, acute process; each elytron with three evenly spaced small basal foveae; sutural stria complete, discal stria absent; legs simple; male last sternite (Fig. 5A) broadly and ovally depressed, rugosely textured, with five thick basally arched setae at middle of apical margin; male apical tergite short, transverse, with broad, densely setose band along posterior margin. Aedeagus (Fig. 6A) elongate, with separate accessory sclerites; parameres separate, knobbed at base, converging and fused with median lobe at middle; median lobe narrow, with small, oval basal foramen, distal portion prolonged into a dorsoventrally flattened blade, margins sinuate to uneven, asymmetrical, subtruncate apical margin; each accessory sclerite with thick, round, sclerotized basal knob, with a thin, semi-spiraled process expanded into triangular apical blade. TL 1.44 mm, EW 0.5 mm. Distribution. This species is known only from El Pahuma Orchid reserve, Pichincha, Ecuador. Figure 2. Panabachia near vulnerata (the type of the genus). A. dorsal habitus; B. pronotum; C. aedeagus (one accessory sclerite missing). 233 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia Etymology. This species is named for its type locality. Remarks. The first four species treated here are very similar in genitalic form, having a flattened aedeagus and free accessory sclerites (Fig. 6A-D). The parameres in these species are all free at the base, fusing with the tegmen around its midpoint (which is also the case in a few species that follow these). The tegmen has a rather small basal foramen, remains wide to near the apex, and has one side entire, the other emarginate. In all of them the accessory sclerites are quite large, with curved inner edges and variously emarginated outer ones. Given this similarity, the four show remarkably different male pronotal modifications, with one species (P. inornata) exhibiting none whatsoever, and their identification based on external characters is straightforward. Panabachia pahuma has a unique median ‘horn’ at the anterior margin of the pronotum (Fig. 4A). It is also unusual in occurring lower than most of the others treated in this paper, known from cloud forest habitats (~2500 m) rather than Páramo. There are a few other cloud forest species, but it seems to be a rare excursion from the more typically high elevation habits of this genus. Figure 3. Dorsal habitus of selected Panabachia spp. A. P. pahuma; B. P. pastazae; C. P. romeroi; D. P. salebrosa; E. P. urbana; F. P. carltoni; G. P. patera; H. P. vigilans; I. P. perdita; J. P. ambulans. 240 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia Those females are slightly larger, with a more nearly angulate, anteriorly widest pronotum, and show a slight swelling of the metaventrite, more fully expressed in the male. Future work that is able to sequence fresh males of this species will be necessary to evaluate this suggestion. Panabachia ayauma sp. nov. https://zoobank.org/567B99DC-C051-40A2-A7BB-C49734EBB379 Figs 4F, 6E Type material. Holotype • ♂ (QCAZ-I-278819): “ECUADOR: Chimborazo, El Releche, 3124 m, 00°38.400'S, 78°30.426'W, 8-VII-2016, SIMT256, Berlese, S. Muñoz & A Romero” / “Muñoz DNA Voucher, Ex. SIMT256, Morphosp. Releche14”; deposited in QCAZI. Paratypes (4♀) • 2 same data as holotype (SIMT257, SIMT283) (QCAZ-I-278820 to 278821) • 2: same locality as type, but collected on 22-Jul-2017 (SIMT340, SIMT341) (QCAZ-I-278822 to 278823). Other material. 1: same data as type, 22-Jul-2017 (SIMT342), but sample vial included parts of 1 male and 1 female specimen. It is unclear which was sequenced in Muñoz-Tobar and Caterino (2020) (QCAZ-I-278824). Diagnosis. Head missing (lost during DNA extraction); male pronotum (Fig. 4F) wider than long, widest toward front, sides more or less even rounded, basal margin of disk depressed; most of disk sparsely punctate and with long setae; median portion of disk glabrous, broadly longitudinally depressed from near base to just in front of middle, depression there splits around broad, posteriorly widened and truncate anteromedian ridge that is densely lined with transverse series of diverging setae; anterior depressions on each side of ridge deeply rounded, smooth, with longer setae in irregular series around anterior and lateral edges; laterad these median depressions, each side of pronotum has smaller, round, secondary depression, also with series of setae around lateral margin; elytron with three evenly spaced foveae, no discal stria, apical stria incomplete; male last abdominal ventrite depressed; opposing edge of male last tergite widened, setose; legs simple. Aedeagus (Fig. 6E) broad and flat, with separate accessory sclerites; parameres separate, thin at base, converging and articulated with median lobe basad of middle, free and setose apically; median lobe with broad base, lacking basal apodeme; basal foramen large, ovate; median lobe slightly narrowed toward wide, very shallowly emarginate apex; accessory sclerites moderately asymmetrical, each bipartite, with thinner, flatter median portion and smaller, articulated lateral portion; the median elements weakly spatulate distally, with outer margins knobbed at articulation point; the lateral portions more strongly sclerotized, short, angulate on outer edge, scoop-like on inner apical surface. TL 1.35–1.41 mm, EW 0.37–0.42 mm. Distribution. This species is only known from subpáramo habitats surrounding the Releche Hacienda, province of Chimborazo, Ecuador. Etymology. The name of this species comes from the Kichwa, meaning ‘the one that leads the way’. Remarks. The aedeagus of this species (Fig. 6E), with a broad flattened tegmen, separate parameres, and large accessory sclerites, allies it with all those preceding. The tip of the aedeagus is distinct, being rather simply subtruncate. And the accessory sclerites themselves are subdivided into sclerotized and 241 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia larger laminate portions. Externally the species is distinguished by a unique male pronotum (Fig. 4F), depressed along much of the midline, opening into deep, paired anterior depressions, with a dense transverse setal series at the split. Panabachia pastazae sp. nov. https://zoobank.org/D216F449-064E-46CD-839E-5B86AE4B6D7E Figs 3B, 4G, 6F Type material. Holotype • ♂ (MCEN-EN 40801): “ECUADOR: Tungurahua, -1.3829, -78.2909, Rio Machay Reserve, 2382 m, 12.XI.2024, M.Caterino, sifted litter” / “Caterino DNA voucher, Ext. MSC-13154, Morphosp. Mch.017”; deposited in MECN. Diagnosis. Head (Fig. 3B) broad, setose, with vertexal foveae deep, non-setose, closer to eyes on each side than to each other; eyes protuberant, moderately large, diameter approx. equal to postocular genal length; antennal bases elevated slightly elevated, set off by oblique striae; antennae short, antennomere II globose, III subconical, antennomeres IV–VI beadlike, rounded, VII–IX increasingly transverse, IX 2 × the size of antennomere VIII, transverse, antennomere XI ~2 × as long as X, rounded, densely setose, with small depression on inner apex; male pronotum (Fig. 4G) slightly wider than long, sides evenly rounded; disk convex in basal 1/2, with very fine mediobasal fovea, conspicuously clothed with mostly inwardly directed setae, with small, discrete, rugose areas on either side; anterior portion of pronotal disk with wide, arcuate depression, narrowest at middle, slightly broader at sides where depression extends nearly to each anterior corner, depression with denser setae along anterior margin; pronotal disk in front of depression with setae mostly directed posteriorly toward depression; elytra moderately long, convex, with sides rounded, each with just sutural and single lateral dorsobasal fovea, a distinct sutural stria extending posterad from fovea; male last tergite truncate and setose at apex; prosternum short, anterior prosternal foveae present; legs simple. Aedeagus (Fig. 6F) elongate with separate accessory sclerites; parameres free at base, sinuate, converging to tegmen at middle; tegmen flattened, narrowest at base, with large, elongate oval basal foramen, unevenly widened apically, distal third broadly spatulate, apex rounded and weakly sclerotized; accessory sclerites more or less symmetrical, each comprising small sclerotized basal piece articulated with flattened, curved laminate blade with subacute apex. TL 1.12 mm, EW 0.32 mm. Distribution. This species is only known from montane forest at the Reserve Rio Machay, province of Pastaza, Ecuador. Etymology. The name of the species comes from the Rio Pastaza watershed where it is so far exclusively found. Remarks. This species has only been collected in mid-elevation cloud forest, and not páramo habitats. With only a single example known, we cannot say whether this is an exclusive preference, but it sets it apart from most of the other species here. Its rather simple transversely arcuate anterior pronotal sulcus (Fig. 4G) is unique, as is the presence of only two basal elytral foveae. These characters, along with geography, should make it straightforward to identify without resorting to the also unique, spatulate aedeagus (Fig. 6F). 242 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia Panabachia romeroi sp. nov. https://zoobank.org/0D4FEB19-4FEF-4528-8E36-CDB04224E693 Figs 3C, 4H, 6G Type material. Holotype • ♂ (QCAZ-I-280223): “ECUADOR: Carchi, 0.70587, -77.96642, El Angel, Saladero, 3301 m, 26JUL2016, S. Muñoz & R. Tobar, Ex. Berlese, Páramo de frailejones” / “Muñoz DNA Voucher, Ex. SIMT289, Morphosp. ElAngel17”; deposited in QCAZI. Paratypes (1♂, 1♀) • same data as holotype (SIMT289: QCAZ-I-280224 to 280225). Diagnosis. Head (Fig. 3C) broad, sparsely setose; posterior margin of the head elevated, rounded; two vertexal fovea deep, non-setose, closer to eyes on each side than to each other; eyes protuberant, moderately large, diameter equal to postocular genal length; antennal bases slightly elevated, set off by oblique stria; antennae short, antennomere II globose, III subconical, antennomeres IV–VI beadlike, rounded, VII–IX increasingly transverse, IX 2 × the size of antennomere VIII, transverse, XI ~2 × as long as X, rounded, densely setose, with small depression on inner apex; male pronotum (Fig. 4H) slightly longer than wide, widest just behind middle, strongly narrowed to front, anterior portion of lateral margins slightly concave; disk convex in basal 1/2, with fine mediobasal fovea, mostly smooth, with sparse, fine, setose punctures, slightly denser anteriorly and laterally; anterior portion of disk with widely V-shaped depression, smooth within, the upper arms nearly reaching anterior margin, with long, anteriorly directed setae along posteromedial margin and a denser tuft at the inner base of the V; elytra sparsely setose, each elytron with four evenly spaced foveae, no discal stria, apical stria complete; male last abdominal ventrite entirely depressed, rough, apical margin outwardly subangulate, with recurved setose tooth at apex; opposing surface of male last tergite broadly emarginate, thickened, setose on lower edge; legs simple. Aedeagus (Fig. 6G) broad at base, with wide flat basal rim, flattened, bent abruptly distad ~1/4 from base; parameres free at base, bent inward and fusing with median lobe just distad basal foramen, apices bent in parallel with median lobe, short, free, weakly setose; median lobe narrowing toward apex, with tip upturned, weakly curved, narrowly subacute; accessory sclerites present, free, very reduced, with short, thick basal part and thin, crescent-shaped apical part. TL 1.28–1.41 mm, EW 0.34–0.38 mm. Distribution. This species is only known from páramo habitats of central Carchi province, Ecuador. Etymology. We name this species to acknowledge the assistance in the field of Dr. Andrés Romero-Carvajal, who helped collect several specimens described in this manuscript. Panabachia uktu sp. nov. https://zoobank.org/C3CFF9D9-BDE4-4382-B519-A3EA1630CFA8 Figs 4I, 5B, 6H Type material. Holotype • ♂ (QCAZ-I-280226): ECUADOR: Chimborazo, Páramo de Atillo, 3501 m, 02°11.265'S, 78°31.2601'W, 07-VII-2016, Berlese, S. Muñoz & A. Romero” / “Muñoz DNA Voucher, Ex. SIMT344, Morphosp. Atillo15”; 243 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia deposited in QCAZI. Paratypes (1♂, 3♀) • same data as holotype (SIMT343, SIMT346, SIMT347, SITM349; QCAZ-I-28022 to 2802230). Diagnosis. Head broad, sparsely setose, with lateral vertexal foveae deep, not setose, closer to eyes on each side; median fovea absent; eyes protuberant but not large, diameter ~2/3 of postocular genal width; two gular foveae present; antennal bases elevated, set off by oblique stria; antennae short, antennomere III slightly longer than wide, IV–VI beadlike, rounded, VII–IX increasingly transverse, X 2 × the length of antennomere IX, transverse, XI ~2 × as long as X, rounded, densely setose, with depression on inner apex; male pronotum (Fig. 4I) approx. as long as wide, sides just slightly wider in anterior 1/2, weakly constricted basally; disk weakly depressed along basal margin, posterior 2/3 almost evenly convex, with fine mediobasal fovea; most of disk sparsely and finely punctate, with long, slightly flattened setae in punctures; small, oval median depression present just behind anterior margin, with interior circular carina within, setose within this circle and along upper margin of depression; oblique lateral depressions present extending posterolaterad from median depression, these with denser series of setae along anterior and lateral edges; two anteroprosternal foveae present; each elytron with four basal foveae, three foveae evenly spaced and one disjunct, discal stria absent; apical elytral stria incomplete; wings present; male last sternite (Fig. 5B) flattened, setose along apical margin, with small bifid process on apical margin (this is the external manifestation of a larger, internal, forked lever that articulates outward when genitalia are everted), flanked by modified setae; legs simple. Aedeagus (Fig. 6H) with base curved ventrad, flattening out and widened apically, with separate accessory sclerites; parameres long, connected at sides of basal foramen, free to setose apex; median lobe with basal apodeme curved underneath and slightly distad; basal foramen ovate; median lobe slightly narrowed beyond foramen, then expanded widely to subtruncate, brushy apex, with pads beneath; accessory sclerites complex, asymmetrical, associated with base of median lobe rather than apex; left side with short clubbed process at base, and long thin, apically curving distal process; right side with strong basal arch dividing into two thin, curved distal processes, neither as long as thin process of other side. TL 1.34–1.69 mm, EW 0.32–0.47 mm. Distribution. This species is only known from grassland and shrub páramo around the Atillo lakes, province of Chimborazo, Ecuador. Etymology. The name of this species comes from a Kichwa word meaning hole, referring to the male pronotal modification. Remarks. See the following remarks section for details on separating this from the sympatric and closely related P. salebrosa. Panabachia salebrosa sp. nov. https://zoobank.org/FAD844D4-99A7-41D0-BF9A-B5DF82E364E9 Figs 3D, 4J, 6I Type material. Holotype • ♂ (QCAZ-I-280231): “ECUADOR: Chimborazo, -2.18775, -78.5210, Atillo, 3501 m, 8JUL2017, S. Muñoz & A. Romero, Ex. Berlese” / “Caterino DNA voucher, MSC-13034, Morphosp. Atillo1”; deposited in QCAZI. Paratypes (3♂) • same general locality as holotype, but collected on 7-Jul-2016 (QCAZ-I-280255-280257). 244 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia Diagnosis. Head (Fig. 3D) broad, sparsely setose, with vertexal foveae deep, non-setose, closer to eyes on each side than to each other; median fovea absent; eyes protuberant but not large, diameter ~2/3 of postocular genal width; two gular foveae present; antennal bases elevated, set off by oblique stria; antennae short, antennomere III slightly longer than wide, IV–VI beadlike, rounded, VII–IX increasingly transverse, X 2 × the length of IX, transverse, bluntly acuminate, with setose depression on anterior surface, XI ~2 × as long as X, rounded densely setose with depression on inner apex; male pronotum (Fig. 4J) almost as long as wide, sparsely and finely punctate and setose, setae slightly denser in anterior 1/3, widest behind midline, narrower at apex than base, anterior margin slightly rounded; posterior 1/2 of disk evenly convex, with fine median fovea; bifid depression (broad depression slightly constricted along midline) present in anterior third, anterior edge a little more abrupt than posterior; shallower, slightly oblique lateral impressions present alongside median ones; two anteroprosternal fovea present; each elytron with four basal foveae, evenly spaced; elytra sparsely setose, discal stria absent, apical stria incomplete; wings present, male last sternite flattened, transverse, sparsely setose; male apical tergite transverse, slightly convex in the middle, sparsely setose; legs simple. Aedeagus (Fig. 6I) flat, with separate accessory sclerites; parameres separate, thin at base, briefly fused to median lobe subbasally, then free to apex, expanded; median lobe with basal apodeme distinct, T-shaped; basal foramen small, ovate; median lobe slightly narrowed beyond foramen, then expanded widely to weakly rounded, brushy apex, with setose apices of parameres alongside; accessory sclerites complex, asymmetrical, associated with base of median lobe rather than apex; both with tightly coiled base opening into elongate, sinuate distal processes, one much longer and thinner than other. TL 1.31–1.82 mm, EW 0.37–1.41 mm. Distribution. This species is only known from grassland and shrub páramo around the Atillo lakes, province of Chimborazo, Ecuador. Etymology. The name of this species refers to the ‘uneven’ anterior portion of the male pronotal disk. Remarks. This species is sympatric with P. uktu, and the two exhibit similar male genitalia (Fig. 6H vs Fig. 6I). Yet there are clear differences in the more basally simpler median lobe of P. salebrosa, in its flatter median lobe, and in its more prolonged left accessory sclerite, as well as in minor differences of male pronotal morphology. Panabachia urbana sp. nov. https://zoobank.org/DCE584EF-A69D-4954-ADD5-CD314CCED7AE Figs 3E, 4K, 5E, 6J Type material. Holotype • ♂ (MECN-EN 40874): “ECUADOR: Pichincha, Quito, Cerro Atacazo, -0.347, -78.611, m, Arbustivo quemado, 01-dic-2018, J. Obregón” / “PTA-196” / “Caterino DNA voucher, Ext. MSC-13316, Morphosp. Panabachia_ Atacazo2” / “MECN-EN 40874”; deposited in MECN. Paratypes (2♂, same locality as holotype) • 1: 1-Nov-2018 • 1: 1-Mar-2019 (MECN-EN 40875 to 40876). Diagnosis. Body (Fig. 3E) dark reddish, densely covered with short, semierect setae; head broad, subquadrate, densely setose, with lateral vertexal 245 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia foveae deep, not setose, closer to eyes on each side than to each other; eyes protuberant but not large, diameter slightly greater than postocular genal width; genal carinae not exaggerated; two gular foveae present; antennal bases slightly elevated, set off by oblique stria; antennae short, antennomere I short, cylindrical, II similar in width, ovoid, III–VI, beadlike, rounded, VII–IX increasingly transverse, shorter, X short, XI ~2.5 × as long as X, narrowly rounded, densely setose, with depression on inner apex; no pronotal dimorphism, pronotum (Fig. 4K) slightly wider than long, widest near middle, slightly constricted before the base; median pronotal fovea fine, deeply impressed; each elytron with four basal foveae, evenly spaced; sutural stria complete, discal stria absent; wings absent; 1st abdominal tergite with close, short pair of curved basal carinae; pair of anteroprosternal foveae present; legs simple; 1st male abdominal ventrite (Fig. 5E) weakly emarginate at middle; last male ventrite wide, flattened, densely setose along most of apical 1/2; last male tergite with apical margin wide, setose, similar to opposing ventrite. Aedeagus (Fig. 6J) relatively flat, with basally free parameres and separate accessory sclerites; parameres fused to median lobe at sides of large, elongate oval basal foramen; median lobe with enlarged basal knob, distally widening slightly to asymmetrically bifid apex, with left side emarginate; accessory sclerites slightly asymmetrical, with tightly coiled basal part, and enlarging, laminate distal part with curved and thickened margins. TL 1.38–1.60 mm, EW 0.35–0.41 mm. Distribution. This species is known only from páramo habitats at Cerro Atacazo, province of Pichincha, Ecuador. Etymology. This species is named for its proximity to the Quito metropolitan area. Remarks. This species is sympatric with P. patera, but distinctive in numerous characters. Panabachia urbana lacks male pronotal dimorphisms (Fig. 4K) (and males can be easily recognized by modifications of the last abdominal ventrite; Fig. 5E), and is also lighter in coloration, a bronzy brown, and broader in shape. Panabachia carltoni sp. nov. https://zoobank.org/43F59321-6771-4690-A0D7-8176741C9285 Figs 3F, 4L, 6K Type material. Holotype • ♂ (MECN-EN 42138): “Ecuador: Pichincha Pr. 50 km NW Quito, Reserva Maquipucuna, #13, elv. 1350 m, 20 Dec.1991, light, C. Carlton, R. Leschen” / “Caterino DNA voucher, Ext. MSC-13326, Morphosp. PanMaq1”; deposited in MECN. Paratypes (1♂, 1♀) • same data as holotype (MECN-EN 42139 to 42140). Diagnosis. Body (Fig. 3F) pale brownish-orange, with conspicuous whitish, subrecumbent setae; impunctate; head subquadrate, posterior corners rounded; vertexal foveae deeply impressed; eyes protuberant; antennal bases with very short stria behind; antennae short, basal two antennomeres similar in length, cylindrical, III subconical, IV–VIII short, subquadrate, IX wider, transverse, barely part of club, X–XI conspicuously larger, setose; pronotum broad, sides almost evenly rounded, slightly wider anterad; anterior part of male pronotal disk (Fig. 4L) with transverse depression, bi-arcuate, anterior midpoint 246 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia pointing posterad over middle, bearing few diverging setae at apex, depression glabrous within, setose particularly along posterior margins; laterad depression are very slightly raised, elongate oval disks, finely rugose but not setose within; posterior portion of pronotal disk with median fovea but otherwise unmodified; elytra with four approximately evenly spaced basal foveae; prosternum very short, hypomeron with anterior foveae; male metaventrite slightly swollen, otherwise unmodified; male last abdominal ventrite short, shallowly transversely depressed; margin of last male tergite unmodified. Aedeagus (Fig. 6K) flattened, with basally separate parameres, and separate accessory sclerites; tegmen with apex expanded, membranous across front; accessory sclerites asymmetrical, one with short blunt tip, other with elongate, apically hooked tip. TL 1.32–1.35 mm, EW 0.33 mm. Distribution. This species is only known from the Maquipucuna Reserve, ca 35 km NW of Quito. Although several other species occur very close by (3 species at El Pahuma), none appear to be particularly closely related to this one. Etymology. We name this species to acknowledge the collaboration and assistance (in this and many other projects) of Dr. Christopher Carlton, who helped collect the types of this species. Remarks. There is limited material of this species, and the male dissection is not well sclerotized nor well cleared. Therefore several details are obscured. Nonetheless it does have several unique features, including the apical prolongation of one of the accessory sclerites. The dorsal velum (visible in Fig. 6K) of the tegmen is possibly unique, but possibly also a remnant of an internalized tergite not extracted in other species. Panabachia falini sp. nov. https://zoobank.org/F52768B0-BE91-452A-898D-F5D685F45B2A Figs 4M, 6L Type material. Holotype • ♂ (QCAZ-I-278840): “ECUADOR: Pichincha, Nono, 15.1 km NW, 2000 m, 0°1'58"S, 78°39'19"W, 24–26 OCT 1999; Z. H. Falin, ECU1F99 022, ex: flight intercept trap” / “SM0351637” / “QCAZ-I-278840”; deposited in QCAZI. Other material. 1♀: “ECUADOR: Pichincha, Bellavista Reserve, Ridge Trail, 12 km S Nanegalito, 2250 m, 0°0'54"N, 78°40'56"W, 28 OCT 1999; R. Anderson, ECU1A99 211F, ex. Cloud forest litter” (SM0362958/QCAZ-I-278841). Diagnosis. Body dark orange, very finely setose, setae recumbent, impunctate; head subquadrate, posterior corners rounded, vertex slightly swollen above deeply impressed vertexal foveae; frons with oblique depressions mediad vertexal foveae; antennal bases weakly elevated in front of fine, oblique striae, striae nearly meeting across middle; eyes large, protuberant, diameter ~2 × post-ocular genal length; antennae short, antennomere I slightly curved, ~2 × as long as wide, II subquadrate, ~2/3 length of I, III subconical, slightly longer than wide, IV–VII weakly subquadrate, VIII transverse, shorter, IX–XI forming loose club, XI ~2 × as long as X; male pronotum (Fig. 4M) wider than long, elevated across entire anterior margin, sides angulate at widest point, at sides of anterior elevation; deep triangular impression present behind middle of anterior elevation, with short, dense, transverse cluster of setae projecting backwards from anterior margin just laterad midline, a few longer setae projecting 247 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia anterad from posterior edge of impression; depression shallowly continued in anterolateral, glabrous depressions; extreme sides of pronotal disk with irregularly ovoid, very shallow, glabrous depressions, their outer/lower edges forming lateral pronotal margin; each elytron with four basal fovea, the lateral pair in a common depression, barely distinguishable; no subhumeral fovea or stria; flight wings present; prosternum very short, with anterior prosternal foveae at anterior corner of hypomeron; legs simple, mesotrochanter slightly dentate at middle of posterior edge; male metaventrite setose posteriorly between metacoxae; male last abdominal ventrite densely setose along distal margin, depressed on either side of a thin, prominent median denticle; terminal male tergite with apical margin widened, flattened, and densely setose. Aedeagus (Fig. 6L) slender, base slightly bulbous, with small, oval basal foramen; parameres free for basal 1/4, then fused with median lobe just beyond foramen; median lobe slightly constricted beyond paramere fusion, dorsally arched, with long median part a flattened cylinder, a shorter, slightly widening dorsal process over it, just shorter, and lateral ‘wings’ with long, curved, articulated rods recurving toward middle at apices. TL 1.6–1.67 mm, EW 0.33–0.36 mm. Distribution. This species is known from northeastern montane forest in the province of Pichincha, Ecuador. Etymology. We name this species to acknowledge many years of collaboration and assistance, on this project and numerous others, from Dr. Zachary Falin, collector of the type of this species. Panabachia papallacta sp. nov. https://zoobank.org/FB2E93D3-219C-4A2F-B3B4-B74ECFA1B62B Figs 5J, 7A, 8A Type material. Holotype • ♂ (QCAZ-I-278835): “ECUADOR: Pichincha, -0.30795, -78.23255, La Virgen, 3694 m, 28JUN2016, S. Muñoz, & R. Tobar, Ex. Berlese, Bosque Polylepis y pajonal” / “Muñoz DNA Voucher, Ex. SIMT246, Morphosp. LaVirgen1”; deposited in QCAZI. Paratypes (1♂, 4♀) • 1 same data as holotype (SIMT275) • 4: “ECUADOR: Pichincha, -0.32124, -78.21319, La Virgen, 3942 m, 18JDEC2016, S. Muñoz & R. Tobar, Ex. Berlese, Leaf litter & moss” (SIMT305 to 308) (QCAZ-I-278725 to 278829). Diagnosis. Head broad, with lateral vertexal foveae deep, non-setose, closer to eyes on each side than to each other; median fovea absent; eyes protuberant but not large, diameter ~2/3 postocular genal width; in ventral view genae projected posteroventrad, rounded (Fig. 5J); two gular foveae present; antennal bases slightly elevated, set off by oblique stria; antennae rather short, antennomere III slightly longer than wide, antennomeres IV–VI beadlike, rounded, VII–IX increasingly transverse, shorter, X short, widening slightly to apex, XI ~2 × as long as X, rounded, with setose depression on inner apex; no antennal dimorphism; male pronotum (Fig. 7A) short, ~1.5 × as wide as long, widest at base, unevenly narrowed to weakly emarginate anterior margin; pronotal disk with large pair of close, deep impressions, a thin carina separating them, with three short, distinct rows of setae at middle and outer edges; outside of major depressions, lateral portion of disk with secondary, shallower impressions just anterad middle, lateral margin slightly angulate near their leading edges; posterior 248 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia portion of the pronotum slightly swollen, with fine mediobasal fovea and a sparse posterior marginal setal row; two anteroprosternal foveae present; each elytron with four basal foveae, three foveae evenly spaced and one disjunct; discal stria absent; wings present, reduced in size; legs simple; male last sternite convex, with a forked process at the apical margin, setose patches at either side, basally arched setae at middle of apical margin absent; male apical tergite transverse, slightly depressed in the middle. Aedeagus (Fig. 8A) with elongate median lobe and separate accessory sclerites; parameres apparently fused at base and middle of median lobe, briefly separate subbasally; median lobe narrow, with elongate oval basal foramen, narrowed beyond and curved laterad to rounded apex; accessory sclerites large and elaborate, with strongly sclerotized rim surrounding inner hooked disk. TL 1.56–1.59 mm, EW 0.28–0.34 mm. Distribution. This species is only known from grassland páramo and Polylepis forest of the Papallacta region, province of Pichincha, Ecuador. Etymology. This species is named from the Kichwa word ‘papallacta’, meaning land of potatoes, referring to the highlands where specimens were collected. Remarks. This species and the following (P. ananay) are very closely related, and actually more or less indistinguishable in male genitalia (Fig. 8A vs Fig. 8B). They also share unusual, enlarged posterior genal margins (Fig. 5J, M). However, they are nonetheless clearly separated by genetic data (see Muñoz-Tobar and Caterino 2020) and minor details of pronotal morphology (Fig. 7A vs Fig. 7B). In particular, this species has the lateral angles of the male pronotum further forward and punctate, while the anteromedial cavities are more clearly Figure 7. Pronota of Panabachia spp. A. P. papallacta; B. P. ananay; C. P. cayambi; D. P. cryptica; E. P. patera; F. P. vigilans; G. P. perdita; H. P. ambulans. 249 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia subdivided into larger posterior and smaller anterior depressions. In P. ananay, the main anteromedial cavity is barely divided at all, only by a short rim of setae, with the posterior area smaller than the anterior. It is worth noting that this species is only known from the single locality, whereas P. ananay occurs at other localities in Pichincha, Chimborazo, and Carchi provinces. Panabachia ananay sp. nov. https://zoobank.org/B5F14BE3-1307-4256-8BF0-D9B8B6AA2153 Figs 5M, 7B, 8B Type material. Holotype • ♂ (QCAZ-I-278830): “ECUADOR: Pichincha, -0.18765, -78.54053, Ruco Pichincha, 3897 m, 22JUN2016, S. Muñoz, A. Romero & S. Myers, Ex. Berlese, páramo” / “Muñoz DNA Voucher, Ex. SIMT296, Morphosp. Pich/ Virgen2”; deposited in QCAZI. Paratypes • 2♀ same data as holotype (SIMT250, SIMT266) (QCAZ-I-278831 to 278832). Other material. 4♀: “ECUADOR: Pichincha, -0.30795, -78.23255, La Virgen, 3694 m, 28JUN2016, S. Muñoz, & R. Tobar, Ex. Berlese, Bosque Polylepis y pajonal” (SIMT274, SIMT276, SIMT277, SIMT247) • 1♀: “ECUADOR: Chimborazo, -1.6400, -78.5071, Releche, 3124 m, 08JUL2016, S. Muñoz & A. Romero, Ex.Berlese (SIMT284) • 1♂: “ECUADOR: Carchi, 0.70587, -77.96642, El Angel, Saladero, 3301 m, 26JUL2016, S. Muñoz & R. Tobar, Ex. Berlese, Páramo de frailejones” (SIMT248) (QCAZ-I-278833 to 278834, 280203 to 280206). Figure 8. Aedeagus (dorsal view) of Panabachia spp. A. P. papallacta; B. P. ananay; C. P. cayambi; D. P. cryptica; E. P. caranqui; F. P. patera; G. P. vigilans; H. P. perdita; I. P. ambulans. 256 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia unmodified; elytra short, sides evenly tapered to humeri (wingless), each with four basal foveae, the lateral pair approximate and sharing a depression; 1st male protarsomere (Fig. 9A) enlarged into distinct setose lobe beneath and along lateral margin of 2nd; male 2nd mesotarsomere (Fig. 9B) enlarged, with distinct, blunt ventral tooth, 3rd mesotarsomere thin, short, mesotarsal claw single, very elongate, able to chelate with 2nd tarsomere’s ventral tooth; all tarsal claws unpaired, long; abdomen large; male last ventrite (Fig. 5H) broadly and deeply depressed, densely setose, opposing surface of last tergite similarly depressed and setose. Aedeagus (Fig. 8I) radically asymmetrical, basal foramen oval, parameres indistinct; tegmen with three irregularly trilobed, left lobe long, curved, articulating apically with enlarged, weakly coiled accessory sclerite, this sclerite with a separate, thin distal process; median lobe of tegmen shortest, ending bluntly, free, between enlarged lateral lobes; right lobe of tegmen arising from a large, arcuate basal lamina, tip flattened, curving ventrolaterad, with acute, dorsal subapical spine, tip recurved dorsad, bluntly rounded; second accessory sclerite free near apex of right tegmental lobe, simple, elongate, curved. TL 1.88 mm, EW 0.38 mm. Distribution. This species is only known from grassland and shrub páramo around the Atillo lakes, province of Chimborazo, Ecuador. Etymology. The name of this species means ‘walker’, a reference to its flightlessness. Remarks. This species is unique in numerous respects. It is considerably larger than any other, and the male’s flightlessness (and clearly associated body proportion changes - the bulky abdomen in particular; Fig. 3J) see no parallels in other species. The lack of male pronotal modifications recurs in a number of other species, none of which appear to be closely related to this species or to each other. The male shows tarsal modifications seen in no other species. While a few other males exhibit slightly modified setae on the basal protarsomere(s) (e.g. P. salebrosa), none have the basal tarsomere itself modified, and the enlarged tarsal claws that can chelate with the 2nd mesotarsomere are completely unique. Finally, the aedeagus is very difficult to homologize with any other. Yet, upon closer inspection, it exhibits a basally flattened tegmen, with small oval basal foramen, and it has free accessory sclerites. The tegmen is highly asymmetrical, and subdivided into a trio of lateral lobes, one of which has become closely associated with a highly modified accessory sclerite. The second accessory sclerite appears to have been reduced to a small vestige. Figure 9. A. protarsus and B. mesotarsus of P. ambulans. 257 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia Discussion This study has revealed a surprising diversity of Panabachia species from a remarkably small area, considering that the genus is distributed across montane regions from Costa Rica south to Bolivia. Undoubtedly large numbers of species remain to be discovered as the high Andes and Central American cordillera are better documented through litter sifting techniques. Most of the species documented here are known from single localities, despite fairly thorough surveys by the senior author and others working in the region (e.g. Moret 2005, 2009; Moret et al. 2016) This is not a surprising result, as mountain ecosystems in the northern Andes are characterized by high numbers of endemic species, partially a result of complex geological and paleoclimatic events during the Andes formation (Antonelli et al. 2018; Pérez-Escobar et al. 2022; Romoleroux et al. 2023), as well as influences from adjacent hyperdiverse lower-elevation regions such as Amazonia (Antonelli et al. 2018; de Meyer et al. 2022) and Choco areas (Pérez-Escobar et al. 2019). High rates of diversification have been reported for both páramo (Madriñán et al. 2013; Cortés et al. 2018) and montane forest lineages (Hutter et al. 2017; Karger et al. 2021; Vieu et al. 2022), where species are usually restricted to narrow and fragmented elevational bands. Unfortunately, these diverse upland faunas are increasingly threatened by climate change and changes of land use (Karger et al. 2021; Romoleroux et al. 2023). The main clades highlighted in Muñoz-Tobar and Caterino (2020) correspond well to major morphological groupings based on genitalia. The ‘upper’ (in their fig. 4) clade, corresponding to their numbered species 1–6, includes all those species with a long, curving median lobe, while the lower clade (their species 7–17) corresponds to those species with a shorter, broader, flat median lobe. While we have not had the opportunity to assemble a more complete molecular data set from the larger set of now-known species, we can predict that the as-yet unsequenced species P. pahuma, P. pastazae, P. urbana, P. carltoni, and P. falini will fall out among the latter, while P. patera will be resolved in the former. Determining placements for the morphological outliers, P. perdita, P. vigilans, and P. ambulans is harder to predict, although the very dissimilar aedeagal morphology of the first two would suggest stem or sister group placements. Those species with a longer, curved aedeagus are known only from páramo, while flatter aedeagi are known in both cloud forest and páramo species. We have now also seen new species from Colombia and Venezuela, not described here, that have a flattened aedeagus with separate accessory sclerites, so presumably are members of the ‘flattened aedeagus’ clade. So far the type with an elongated and curved tegmen are limited to the Ecuadorean highlands. Whether this clade extends to higher regions of more distant Andean regions remains to be seen. It is interesting to observe that species lacking male pronotal modifications are found in both major clades. Without even having a hypothetical function for these modifications, it is difficult to speculate on reasons for convergent losses, although if mate recognition were involved, it might be interesting to look more carefully at variation in these characters in communities where more than one Panabachia species was known, such as La Virgen, Atillo, and Mojanda. There are also similar modifications in both clades, with transverse creases, secondary lateral depressions, and median setose fringes, suggesting 258 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia common modes of employment across considerable phylogenetic space. We leave such questions for future students. Studies of alpine beetle faunas in the Andes are in their preliminary stages. But our findings, showing high diversity in a novel beetle lineage from páramo and montane forest in Ecuador, echo those findings available to date. Previous studies have also indicated that tropical alpine ecosystems in the Andes hold a high number of endemic species (Pérez-Escobar et al. 2022; Romoleroux et al. 2023), mostly undescribed among invertebrate groups. Some of the better-studied examples include the ground beetles, for which we have a good understanding of diversity and distribution in Ecuadorian páramo, with more than 224 páramo specialists (Moret 2009; Atiencia-Puca et al. 2023). In a few other groups, such as broad nosed weevils, páramo specialists have been described in the genus Obrieniolus del Río (Coleoptera, Curculionidae, Entiminae) from Peru (del Río and Lanteri 2011), as well as new species in the genus Leschenius del Rio (del Río and Marvaldi 2022) from Ecuadorian páramo. Among the diving beetles, one new species and seven subspecies in the genus Liodessus (Coleoptera, Dytiscidae) from eastern Colombian páramo have been identified (Balke et al. 2023). Similar patterns of high diversity have been described for hover flies from Colombian páramo complexes, with six genera and 37 species endemic to the area of study (Montoya et al. 2021). These studies focused on montane Andean biodiversity clearly indicate a high richness, and show the importance of conserving high elevation ecosystems. Most work to date (including this one) has been limited to particular countries. Integrative regional studies that encompass the range of Andean alpine habitats should be a high priority to reveal broader patterns of diversity and evolution. Acknowledgments We are grateful to numerous individuals and organizations over many years for helping make this study possible. MSC is grateful for assistance with collecting permissions to David Romo and Kelly Swing (both ZSFQ), Joe Meisel and Catherine Woodward (both Ceiba Foundation), Lou Jost (EcoMinga Foundation, and to the Lima Family (El Pahuma Orchid Reserve); for assistance in the field MSC thanks Alexey Tishechkin, Sarah Smith, and Tito Recalde; for assistance in the lab, and during the course of a one-year sabbatical at the Instituto Nacional de Biodiversidad (INaBio) in Quito, MSC is grateful for the help of Diego Inclán, Pablo Jarrín-V, Alex Pazmiño-Palomino, and Ana García-Ruilova. SMT thanks Andres Romero-Carvajal, Shelley Myers, Rosario Tobar, and Corina Argos for their assistance in the field. SMT is grateful to Alvaro Barragan and Fermanda Salazar (PUCE) for assistance with collecting permits, and to Diego Inclan, Alex Pazmiño-Palomino, Ana Garcia-Ruilova, and Marissa Barreno (INaBio) for their assistance in the lab. Diego Cisneros, Margarita Lopez, and Ciara Wirth (ZSFQ) are also acknowledged for their assistance with permissions and specimen deposition. Finally we thank one anonymous reviewer for useful suggestions on the manuscript. Additional information Conflict of interest The authors have declared that no competing interests exist. 259 ZooKeys 1254: 225–261 (2025), DOI: 10.3897/zookeys.1254.158319 Sofia I. Muñoz-Tobar & Michael S. Caterino: Ecuadorian Panabachia Ethical statement No ethical statement was reported. Use of AI No use of AI was reported. Funding No funding was reported. Author contributions Conceptualization: MSC, SIMT. Data curation: SIMT, MSC. Formal analysis: MSC, SIMT. Investigation: SIMT, MSC. Methodology: SIMT, MSC. Writing - original draft: MSC, SIMT. Writing - review and editing: SIMT, MSC. Author ORCIDs Sofia I. Muñoz-Tobar https://orcid.org/0000-0002-4609-3703 Michael S. 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