Marzo-2017-prueba

Revisión de los Elmidae (Coleoptera: Dryopoidea) de Chile

Marcos Ferrú a, b, *, William Shepard c

a Universidad Católica de la Santísima Concepción, Alonso de Ribera 2850, Concepción, Chile
b Centro de Muestreo y Análisis Biológico, Luis Acevedo 456 Chiguayante Bíobío-Concepción, Chile
c University of California, Essig Museum of Entomology, 1101 Valley Life Sciences Bldg., Berkeley, California, 94720 USA

*Corresponding author: marcos.ferru@gmail.com (M. Ferrú)

Received: 12 August 2025; accepted: 17 December 2025

Abstract

The family Elmidae is one of several families of aquatic beetles in the superfamily Dryopoidea. The Elmidae occur throughout the world (except Antarctica), with approximately 150 genera and 1,500 species. Elmidae have been studied as part of the taxa used to assess water quality. There are no keys to identify larvae of Chilean species, and to identify adults it is necessary to remove the male genitalia. Research carried out in Chile has included Elmidae in descriptions, lists and reviews of aquatic beetles in general. Until now, there has been no review dedicated entirely to Chilean Elmidae. In Chile, after this review there are 2 subfamilies, 7 genera and 18 species currently recognized. The species Austrelmis woytkowskii (Hinton), which has been cited in error for Chile, is eliminated from the Chilean list and the species Austrelmis sp. 1 is included, as is Austrolimnius elatus Hinton. Expanded distributions of some species are included.

Keywords: Elmidae; Elminae; Larainae; Diversity; Distribution

Resumen

La familia Elmidae es una de varias familias de escarabajos acuáticos perteneciente a la superfamilia Dryopoidea. Los Elmidae se encuentran en todo el mundo (excepto en la Antártida), con aproximadamente 150 géneros y 1,500 especies. Los Elmidae han sido estudiados como parte de los taxones utilizados para evaluar la calidad del agua. No existen claves para identificar larvas de las especies chilenas y para identificar adultos es necesario extraer los genitales de los machos. Investigaciones realizadas en Chile han incluido a los Elmidae en descripciones, listas y revisiones de escarabajos acuáticos en general. Hasta ahora, no había una revisión dedicada completamente a los Elmidae chilenos. En Chile, después de esta revisión hay 2 subfamilias, 7 géneros y 18 especies actualmente reconocidas. La especie Austrelmis woytkowskii (Hinton), que ha sido citada por error para Chile se elimina de la lista chilena y se incluye la especie Austrelmis sp. 1, al igual que Austrolimnius elatus Hinton. Se incluyen la ampliación de distribuciones de algunas especies.

Palabras clave: Elmidae; Elminae; Larainae; Diversidad; Distribución

Introduction

The family Elmidae (Coleoptera: Dryopoidea) colloquially known as riffle beetles is a group of globally distributed beetles (except for Antarctica) found in clean, permanent waters. Elmidae is divided into 2 subfamilies (Elminae and Larainae), with approximately 150 genera and 1,500 species Jäch et al. (2016) and more taxa are described every year. In Chile, the Elmidae have 2 large faunal divisions. This is a fauna with continental characteristics that shares genera, such as Heterelmis Sharp and Microcylloepus Hinton, with other South American countries. The other faunal area is the Southern Cone, which contains genera shared with New Zealand and Australia, or has genera only shared with Argentina, as is the case of the genera Hydora Anon. [Broun], Luchoelmis Spangler and Staines and Stethelmis Hinton. The different species of Elmidae have similar life cycles. Eggs are laid on submerged substrates such as stones or wood, with hatching occurring in approximately 2 weeks. Pupation lasts about 2 weeks. After about 2 weeks of pupation, the adults emerge, some fly to find new habitats before re-entering the water, while others just re-enter the stream from which they came. Adults can live approximately 10 years (Brown, 1974). Larvae and adults of Elmidae often co-occur in appropriate microhabitats, such as sand, gravel, leaf bundles, aquatic macrophytes, wood, and rocks in clear water, etc. The main requirement for Elmidae is uncontaminated water with high levels of dissolved oxygen, up to moderately polluted environments González-Córdoba et al. (2020). Most Elmidae consume fine particulate organic matter that they browse from the surface or find in the interstitial spaces between gravel particles. This material is usually algae, fungal hyphae, and crushed leaves or wood. The few species that feed on decaying and submerged wood take several years to develop due to the low levels of nutrients wood presents. Reviews of elmid biology are in Brown (1987) and Elliott (2008). Elmidae breathe with tracheal gills (larvae) or plastrons (adults). Both systems require relatively high levels of dissolved oxygen (Brown, 1987; Kodada et al., 2016). Elmidae have been found to be very susceptible to contaminants such as surfactants in soap, petroleum products, accordiong to Brown (1984), Elliott (2008), Lafuente et al. (2019) and anything that removes oxygen from the water. Therefore, their presence in a stream is a sign of uncontaminated water during the lifespan of the Elmidae. Thus, Elmidae are often targeted during water quality assessment programs, along with Ephemeroptera, Plecoptera and Trichoptera, all of which are also susceptible to contaminants (Fierro et al., 2012). When these studies are carried out, it is very important that the sampling considers the microhabitat requirements of the different species. The new global catalog of Elmidae has helped to understand generic and species distributions (Jäch et al., 2016). An identification key for adult genera of South American Elmidae has been developed by Manzo (2005) and an identification key for most larvae of South American Elmidae genera has been developed by Manzo and Archangelsky (2008), Pasos et al. (2018). These have been important scientific advances since Elmidae are an integral part of the ecology of streams and rivers and are often collected during sampling (Fierro et al., 2012). None the less despite its importance there are no keys to identify the larvae to species, and generally to identify the adults it is necessary to extract the male genitalia. The first Elmidae described in Chile are found in 2 articles by Germain (1854, 1892). This was followed by articles published by Champion (1918), Hinton (1941, 1945), Janssens (1957), Spangler and Brown (1981) and Spangler and Staines (2002). Some of the species described in these papers are now found in different genera. Other papers dealing with Chilean dryopoid beetles include authors like Lataste (1897 a, b), Dajos (1974), Solervicens (1974), Shepard (2016) and Shepard and Barr (2023). Nonetheless, the information is scattered and Elmidae continues to be a relatively unknown group. This study will facilitate further efforts on Chilean Elmidae by bringing together information and providing new distribution data.

Materials and methods

The collection of elmid larvae and adults usually involves aquatic nets that are placed downstream of the sampling area where the substrates are disturbed. Elmidae then drifts in the current and into the net. The specimens are collected from the net and preserved in 70% or 95% ethyl alcohol. Some of the net contents collected by WDS were entirely preserved in 95% alcohol bottles and then processed in the laboratory. This makes it easier to collect smaller adults (such as Austrolimnius) and younger larvae. After collecting at each site, ecological characteristics were recorded on field data sheets. Collection locality maps were produced with QGIS and used all known localities, published and new. Locality data are presented primarily as they occur on the specimen labels. A forward slash “/” indicates separate lines in a label and a double forward slash “//” indicates separate labels. Information included by “( )” is below the locality labels. Additional data provided here, but not in the labels, is indicated by brackets “[ ]”. The “WDS-A- ____” numbers refer to WDS data sheets. Collection locality maps were made using ArcGIS and used all known localities, published and new. Additional records of the specimens were obtained from the collections: National Museum of History of Chile (Santiago) and the Museum of the Department of Zoology of the University of Concepción, as well as from published literature on the Elmidae. Abbreviations cited in the text include the following: National Museum of Natural History, Washington, DC, USA (USNM); Natural History Museum, London, England (NHML); Royal Belgian Institute of Natural Sciences, Brusseels, Belgium (RBINS); William D. Shepard (WDS).

Results

As a result of this work, we can consider the diversity of Elmidae to include 18 species, 7 genera and 2 subfamilies. The species Austrelmis woytkowskii (Hinton, 1937) is eliminated because the analysis of samples collected from various streams in northern Chile shows no evidence of its presence. The species Heterelmis trivialis (Germain, 1892) is also eliminated. One of the reasons for this decision is the loss of the type material. Another reason is the fragility of the stream where it was cited; this stream is small and urbanized and tends to have a considerable decrease in its water flow in summer. This species was searched for at the type locality on 2 occasions, thus corroborating its presence in Chile, but it has not been found or reported since its description. In these samplings, the other species described by Germain and cited for the area were found. In this work, the species Austrolimnius elatus, eliminated by Elgueta and Arriagada (1989), is reestablished; this is supported by the type material found. Finally, it is important to mention that despite the limited progress in species richness shown in this work, there are numerous new localities for the different species present in Chile. A list of the Elmidae known from Chile is provided (Table 1).

Key to the genera of adult Elmidae known from Chile extracted from Manzo and Archangelsky (2008) and Passos et al. (2018). 1 Body densely pubescent, without plastron; transverse procoxae, with exposed trochantins; adults fly easily from water or net — Larainae Hydora (Fig. 19). 1’ Body not pubescent, with plastron; procoxa globosa, with unexposed trochantins; adults do not fly from water or net — Elminae (2). 2 Thorax without sublateral carinae (Figs. 27, 30, 32) — 3. 2’ Thorax with sublateral carinae — 4. 3 Tarsal claws with basal teeth (Fig. 31) — Stethelmis. 3’ Tarsal claws without basal teeth — Luchoelmis. 4 Epipleura with 1 or 2 longitudinal lines of granules — Austrolimnius. 4’ Epipleura without longitudinal lines of granules — 5. 5 Pronotum with complete deep transverse groove (Fig. 29) — Neoelmis. 5’ Pronotum without deep transverse groove complete — 6. 6 Pronotum with circular anterior depression, with V-shaped carina in ½ basal, with sublateral elevation (Fig. 28) — Microcylloepus. 6’ Pronotum without depressions; sublateral carina restricted to anterior half (Figs. 25, 26) — Austrelmis.

Generic reviews

This section is reproduced from the article PDF; the genus-by-genus accounts (with new distributional records and material examined) are available in full in the linked PDF.

Discussion

The Chilean Elmidae include 2 subfamilies, 7 genera and 17 species. However, many additional species are known but not yet described. Many geographic areas remain unexplored, so additional species are likely to be found. Endemism is relatively high in the Chilean Elmidae, 82.3% (14 of 17 species) of the species are endemic to Chile. The only Chilean Elmidae not endemic to Chile are Austrolimnius elatus, A. nycteloides, and Hydora annectens.

The number of genera and species present in the South American countries on the Pacific slope decreases from north to south (Ecuador to Chile) (Table 2). Partly this is because the northernmost areas are closer to the equator, where temperatures are more favorable to elmid evolution. Also, partly this is due to Ecuador and Peru having areas located on the east front of the Andes near to the Amazon where the evolution of the Elmidae has gone in a different direction (Shepard unpublished data). Chile lacks an Amazonian component.

In Chile, the elmid fauna has been negatively impacted by coastal and Andean orogenies, especially in the north. This is mainly due to the isolation and desertification that characterizes this area (Shepard unpublished data). The only 2 genera present there are Austrelmis and Microcylloepus. Microcylloepus elsewhere has been able to maintain populations in desert environments where small springs or seeps occur, so it is not surprising that it would be found here, while Austrelmis is a very typical inhabitant of high mountain habitats throughout the Andes.

The Southern Cone area of Chile has genera that were thought to be restricted to this area, such as Austrolimnius, Luchoelmis, and Stethelmis. However, Austrolimnius and Stethelmis have now been collected somewhat farther north, Coquimbo in the case of Austrolimnius and Atacama in the case of Stethelmis, nonetheless their diversity remains in the Southern Cone area.

Two collection attempts in the type locality of Heterelmis trivialis failed to find extant populations. It is not known why this population might have disappeared. Likewise, attempts to find populations of Austrelmis woytkowskii were unsuccessful. These findings reinforce the necessity to amplify sampling focused on aquatic coleoptera and Elmidae specifically, in order to determine if these populations are truly extinct or only minimally abundant.

Acknowledgements

We thank all the people who made this work possible: Walter Sielfeld, for the support on the ground and collection of specimens; Mario Elgueta (National Museum of Natural History); Margarita Ruiz de Gamboa (CEMABIO); Pablo Fierro (Universidad del Valdivia) for his help, contribution of copies and ideas; the specialists Verónica Manzo (University of Tucumán, Argentina) and María Passos (Federal University of the State of Rio de Janeiro, Brazil) for their bibliography. And, finally, the Laboratory of Water Quality Bioindicators EULA-Chile, for their contribution in specimens and land. Additionally, we thank the following people for providing photographs of type specimens and labels under their care: Pol Limbourg (RBINS), Keita Matsumoto (NHML) and Charyn Micheli (USNM).

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Redescripción de Stagmomantis hebardi (Mantodea: Mantidae), con la descripción de la hembra

Aarón Emilio Vásquez-Quintero a, Iker Cubillos-Macías b, Erick Omar Martínez-Luque c, Jovana M. Jasso-Martínez b y Fernando Varela-Hernández a, *

a Universidad Autónoma del Estado de Morelos, Escuela de Estudios Superiores del Jicarero, Laboratorio de Sistemática Molecular, Carretera Galeana-Tequesquitengo s/n, Col. El Jicarero, 62909 Jojutla, Morelos, Mexico
b Universidad Nacional Autónoma de México, Instituto de Biología, Colección Nacional de Insectos, Tercer Circuito Exterior s/n, Cd. Universitaria, Copilco El Alto, Coyoacán, Apartado postal 70-233, 04510 Mexico City, Mexico
c Universidad Autónoma de Querétaro, Facultad de Ciencias Naturales, Av. de las Ciencias s/n, Juriquilla, 76230 Querétaro, Querétaro, Mexico

*Corresponding author: fernando.varela@uaem.mx (F. Varela-Hernández)

Received: 02 September 2025; accepted: 08 December 2025

Abstract

The genus Stagmomantis Saussure, 1869 comprises a group of New World mantids whose taxonomic delimitation and composition have been challenging due to complex morphological variation. Stagmomantis hebardi Rehn, 1935, a species distributed in the Neotropical region and the Mexican Transition Zone, was originally described based on both a female and a male specimen. However, the female was subsequently recognized as belonging to a different species within the subgenus Auromantis. In this study, we provide the redescription of the male and the description of the female of S. hebardi. Descriptions are based on external morphology and both sexes internal genitalia characters. The conspecificity of the male and female is supported by molecular evidence provided in previous studies and further validated by biological observations from captive rearing. The corkscrew-like pda (posterior process of the ventral phallomere) in the male genitalia and the lobe-like structure between the fifth and sixth tergites in the female abdomen are diagnostic characters that distinguish this species from others of the genus Stagmomantis.

Keywords: External morphology; Internal genitalia; Mantids; Taxonomic description

Resumen

El género Stagmomantis Saussure, 1869 es un grupo de mantis del Nuevo Mundo cuya delimitación y composición han sido desafiantes debido a su compleja variación morfológica. Stagmomantis hebardi Rehn, 1935, una especie distribuida en la región Neotropical y la Zona de Transición Mexicana, fue descrita originalmente con base en un macho y una hembra. Más tarde, se reconoció que la hembra pertenecía a una especie diferente del subgénero Auromantis. En este estudio, proporcionamos la redescripción del macho y la descripción de la hembra de S. hebardi. Ambas descripciones se basan en caracteres de morfología externa y de la genitalia de ambos sexos. La asociación específica entre el macho y la hembra está respaldada por análisis moleculares previos y validada por observación directa en cautiverio. El pda (proceso posterior del falómero ventral) en forma de sacacorchos en la genitalia del macho y la estructura en forma de lóbulo entre los terguitos quinto y sexto en el abdomen de la hembra son caracteres clave para separar a esta especie de otras del género Stagmomantis.

Palabras clave: Morfología externa; Genitalia interna; Mantis; Descripción taxonómica

Introduction

The order Mantodea (praying mantises) comprises about 2,500 species worldwide, distributed in 16 superfamilies, 29 families, 60 subfamilies, and 436 genera (Ehrmann, 2002; Otte & Spearman, 2005; Schwarz & Roy, 2019). Mantidae Latreille, 1802 is the most species-rich family, containing approximately half of the described species (Patel & Singh, 2016; Svenson & Whiting, 2004, 2009); however, the limits and phylogenetic relationships within Mantidae remain unclear. Several groups previously defined based on morphological data were later recovered as non-monophyletic by analyses using mitochondrial and nuclear genes (Svenson & Whiting, 2004, 2009), as well as mitochondrial genomes (Wang et al., 2022; Xu et al., 2021).

Stagmomantis Saussure, 1869 is a New World genus belonging to the subfamily Stagmomantinae Giglio-Tos, 1919. It is one of the most species-rich genera within Mantidae, with 25 recognized species (Anderson, 2025). Species delimitation within Stagmomantis has been challenging due to significant morphological variation among its members, rendering the number of valid species debatable (Agudelo-Rondón et al., 2007; Anderson, 2020a, b, 2021; Ehrmann, 2002; Maxwell, 2014; Otte & Spearman, 2005; Terra, 1995). For instance, Maxwell (2014) did not recognize S. conspurcata Serville, 1869 as a valid species, treating it as a synonym of S. carolina (Johansson, 1763). Later, Anderson (2020b) revalidated S. conspurcata, arguing that sufficient morphological differences —such as body size and male hindwing coloration— justified its recognition as a distinct species. This case exemplifies the taxonomic instability present within Stagmomantis, which has important implications, such as constant changes in the reported geographic distribution patterns of its species.

In 1935, Rehn described Stagmomantis hebardi —named after the American entomologist Morgan Hebard— based on a female and a male specimen from Sinaloa, Mexico. Hebard (1923) had previously identified these specimens as S. limbata Hahn, 1835 and S. tolteca Saussure, 1861, respectively. In his description, Rehn included images of the dorsal habitus of both the female and male, as well as an illustration of the male’s internal genitalia. The taxonomic status of S. hebardi, based on the male-female association established by Rehn, has remained unchallenged for decades.

Recently, a molecular analysis grouped together a female and a male specimen previously identified as S. carolina (Varela-Hernández et al., 2022). In addition to the molecular data, that study included high-resolution images of the dorsal habitus. Subsequent comparison of those specimens with the descriptions and illustrations in Rehn (1935) revealed that the male assigned to S. carolina corresponded to S. hebardi. Crucially, the morphological traits of the female described as S. hebardi by Rehn (1935) did not match those of the female specimen molecularly associated with the male in Varela-Hernández et al. (2022).

Based on this evidence, we consider that Rehn (1935) erroneously associated the sexes of S. hebardi, establishing a heterospecific type series. Consequently, the main objectives of this study are to correctly associate both sexes of S. hebardi, to redescribe the male, and to formally describe the female of S. hebardi (Rehn, 1935). This work is supported by the aforementioned molecular evidence and is based on external morphology as well as a comprehensive analysis of male and female genitalia.

Materials and methods

A total of 48 pinned specimens identified as Stagmomantis hebardi were examined (29 males and 15 females, see “Taxonomic summary”). These specimens are deposited at the Colección Nacional de Insectos, Instituto de Biología, Universidad Nacional Autónoma de México (CNIN-IBUNAM) and the Colección Entomológica de la Universidad Autónoma del Estado de Morelos, México (CEUAEM-MAN). Additionally, we examined high-resolution images of the holotype (Fig. 1) and allotype (Fig. 2) deposited at the Academy of Natural Sciences of Philadelphia (ANSP), USA.

Observations and measurements were conducted using an Olympus SZ stereomicroscope. Specimens identification at genus and species level followed the keys and descriptions provided by Rehn (1935), Agudelo-Rondón and Chica (2002), Maxwell (2014), and Anderson (2021). Specimens of both sexes were photographed from multiple angles using a Canon EOS Rebel T5 digital camera equipped with an EF 100 mm f/2.8 Macro USM lens. Camera control was managed via EOS Utility software; images were subsequently stacked and edited using Zerene Stacker and Photoshop CC 2019.

Terminalia of both sexes were softened in hot water (80-100 °C) for 15-20 min, following the protocol of Ferraz et al. (2023). Genitalia were then cleared in a 10% KOH solution at 40 °C for 20-30 min (Brannoch et al., 2017). The extraction of female genitalia followed the method described by Rodrigues and Cancello (2016). Cleared genitalia were rinsed twice in distilled water and stored in 70% ethanol. Remaining soft tissue was removed, and structures were photographed suspended in antibacterial gel to maintain their position (Su, 2016). Abbreviations for all measurements and genitalic structures are provided in Table 1.

To visualize the geographical distribution of the examined specimens, a map was generated using ArcGIS software, utilizing the State Geostatistical Framework, scale 1:250,000 (INEGI, 2024).

Description

Stagmomantis hebardi Rehn, 1935

Diagnosis. Male: distinguished by the corkscrew-like posterior process of the ventral phallomere (pda) (Fig. 3). Female: prosternum (inter-coxal area) blackish; anterior coxae armed with 3-6 blackish spines; presence of a protrusible lobe-like structure located between abdominal tergites 5 and 6 (Fig. 4).

Identification. Males: antennae longer than pronotum. Tubercle present between antennifer and compound eyes. Wings longer than abdomen; fore- and hindwings hyaline; costal area of forewings opaque with brown spots. Genitalia with the posterior process of ventral phallomere (pda) corkscrew-shaped. Females: abdominal tergites 4-6 dorsoventrally flattened; protrusible lobe-like structure present between tergites 5 and 6. Supragenital plate wider than long with well-developed longitudinal medial keel. Forewings opaque, greenish or brownish, with brown spots; hindwings opaque, anteriorly yellowish with brown spots near posterior margin, or yellowish with colorless spots posteriorly. Both sexes: ocellar tubercles absent; ocelli flush with surface of frons. Lower frons smooth, wider than long. Pronotum long and slender; lateral margins denticulate; dorsal surface smooth. Forecoxae approximately 3/4 the length of pronotum; anterior and posterior margins denticulate; forecoxal lobes convergent. Hearing organ of DK type.

Male (Fig. 5). Measurements (Table 2). Head: head capsule (Fig. 6a) with oval eyes; juxtaocular bulges undeveloped; ocellar tubercles undeveloped. Anterior margin of vertex straight, depressed near parietal sulcus. Ocelli larger than in females. A small tubercle present between antennifer and compound eye. Antennae filiform, longer than pronotum, reaching anterior portion of the mesothorax; pedicel nearly as long as scape. Frons wider than long, with an M-shaped depression in the middle of the transverse carina. Thorax: pronotum, supracoxal dilation slightly pronounced; supracoxal groove well-marked. Lateral margins of prozone denticulate; lateral margins of metazone denticulate, denticles decreasing in size posteriorly. In ventral view, inter-coxal area dark-colored. Legs: forelegs, spination formula: F = 4DS/12-15AvS/4PvS; T = 9-14AvS/9-11PsV. Forecoxae armed with 3-5 blunt, often blackish spines, small denticles interspersed between them; coxal lobes convergent (Fig. 6b). Forefemur relatively long, slender; femoral brush located at the level of last 3 posteroventral spines; tibial spur groove located at approximately the middle of femur; first discoidal spine smaller than the others; third discoidal spine longer than the others; blackish spot present at the level of first anteroventral spine (AvS). Foretarsus: distal portion of each segment darkened (Fig. 6c, d). Wings: slightly surpassing tip of abdomen. Forewings: hyaline, with scattered brown spots in discoidal area; stigma covered by dark brown patch; anterior margin wide at base, narrowing posteriorly; apex rounded. Hindwings: hyaline with a dark patch at apex; main central longitudinal veins feature a series of spotted brownish lines running along posterior margin to about half the distance to anterior margin (Fig. 5). Abdomen: slender, cylindrical. Subgenital plate longer than wide. Supragenital plate wider than long; apex rounded, with marked longitudinal medial keel. Cerci elongate; cercomeres cylindrical, subequal in length; distal cercomere smaller, conical, rounded at apex.

Genitalia: left phallomere, sclerite L4B almost rectangular, longer than wide, with left margin projected anteriorly (Fig. 7a). Anterior lobe of phalloid apophysis (aafa) linguiform, sometimes oval, distally dentate. Posterior lobe of phalloid apophysis (pafa) elongated, apically rounded, dentate, bearing 3-6 ventral denticles. Posteromesal lobe (loa) oval, bearing short setae. Posterior process (paa) robust, sometimes slender. Right phallomere: triangular (Fig. 7b); posterior apex rounded with short setae. Dextral extension (bm) thick, slightly elongate; anterior extension of sclerite R3 (an) elongate, slender. Process anteromesal to pia (pva) short, thick, smooth with rounded apex; process posterolateral to pva (pia) short, smooth. Ventral phallomere: sclerite L4A oval (Fig. 7c, d). Secondary spine (ssp) well-sclerotized, triangular. Posterior process (pda) corkscrew-shaped, twisted left or right, with apex angled to slightly flattened.

Female (Fig. 8). Measurements (Table 3). Head: head capsule with oval eyes; juxtaocular bulges absent; ocellar tubercles absent; ocelli smaller than in males. Anterior margin of vertex straight; vertex slightly depressed near parietal sulcus. Antennae filiform; scape almost twice as long as pedicel. Frons wider than long in frontal view, with M-shaped depression in the middle of transverse carina (Fig. 9a). Thorax: pronotum, supracoxal dilation slightly pronounced; supracoxal groove well-marked. Lateral margins of prozone denticulate; lateral margins of metazone denticulate, becoming smooth distally; medial keel of pronotum well-developed. In ventral view, inter-coxal area brownish. Legs: forelegs, spination formula: F = 4DS/14-16AvS/4PvS; T = 12-14AvS/8-10PvS. Forecoxae (Fig. 9b) armed with 3-6 apically blunt spines, often brownish, denticles interspersed among them; forecoxal lobes convergent. Forefemur relatively long, slender; femoral brush located at the level of last 3 posteroventral spines; tibial spur groove located at approximately half the length of femur; first discoidal spine smaller than the others; third discoidal spine longer than the others; with blackish spot at the level of first anteroventral spine (AvS). Foretarsus: distal portion of each segment blackish (Fig. 9c, d). Wings (fig. 10a-c): reaching approximately 3/4 the length of abdomen. Forewings: anterior margin wide at base, narrowing distally; color variable, ranging from brown to light hues; stigma darker than rest of wing. Hindwings fully colored; color variable, ranging from yellowish at base to middle area, with dark spots distally, to yellowish with hyaline spot; apex of hindwing yellowish with greenish portions. Abdomen: dorsoventrally flattened, reaching maximum width between 4th and 6th tergites, with prominent, protrusible lobe-like structure between 5th and 6th tergites. Supragenital plate wider than long, rounded at the apex, with a marked longitudinal medial keel. Cerci elongate; cercomeres cylindrical, subequal in length; distal cercomere smaller, conical, apically rounded.

Genitalia (Fig. 11): gonoplacs (gl9), simple, elongate, apically rounded, bearing short setae throughout, with longer setae apically. Gonapophysis: gonapophysis 9 (gp9) membranous, elongate, shorter than gl9. Gonapophysis 8 (gp8) elongate, bearing short setae; in lateral view, distinct furrow near ventral margin; lateral margins sinuous depressed near base of apical lobe (gpal8). Apical lobe of gonapophysis 8 (gpal8) glove-shaped; dorsal portion laterally compressed, distally rounded; ventral portion short, elongate; surface covered with short setae, longer setae on inner surface. Medial outgrowth of gonapophysis 8 (gpmo8) well-developed. Coxa 8 (CX8): acuminate, broad, rounded at base, tapering distally; divided into dorsal and ventral regions by a marked depression, a prominent keel extending through middle of depression. Dorsal region of CX8 elongate lateral wider, more rounded than dorsal region. Caudogyne (CG8) weakly sclerotized; dorsal margin rounded, sinuous; ventral margin depressed at the middle (M-shaped). Spermathecal bulge (spb) well-sclerotized. Accessory gland supporting lobe (agsl) smooth, weakly sclerotized. Ventral segmental fold 7 (vf7) membranous, rounded, with sinuous margins. Vestibular sclerite (VS7) weakly sclerotized, rhomboidal in shape, with ventro-medial corner rounded.

Taxonomic summary

Holotype: 1 ♀, Mexico. Sinaloa: Venvidio, 2-6.VIII.1918, type # 8988 (ANSP). Allotype: 1 ♀, Sinaloa: Los Mochis, 25.XII.1918 (ANSP). Additional material examined: Mexico. Estado de México: 1 ♂, Nepantla, Finca Badoni, 18°59’0.8” N, 98°50’44.9” W, 2,020 m asl, 05.X.2023 (CEUAEM-MAN-135); 1 ♀, Malinalco, Chalma, 118°55’56.3” N, 99°26’11” W, 29.XI.1980 (IBUNAM: CNIN: MA909). Guerrero: 10 specimens (5 ♂, 5 ♀). Five ♂: Acahuizotla, 17°28’8.9” N, 99°25’10.3” W, 18.XI.1983 (IBUNAM: CNIN: MA644, MA645); Iguala, Km 3 desv. a Tepecoacuilco, 18°17’32” N, 99°28’58” W, 14.X.2006 (IBUNAM: CNIN: MA724); Iguala, Tuxpan, 2 km a Microondas, 18°23’34” N, 99°28’27” W, 14.X.2006 (IBUNAM: CNIN: MA813); Tecpan, La Laguna, Km 15 San Luis, 17°13’14.9” N, 100°37’9.3” W, 1,425 m asl, 20.VII.2005 (IBUNAM: CNIN: MA816). Five ♀: Atenango del Río, Km 36 Huitzuco-Atenango, 18°8’32” N, 99°7’49” W, 200 m asl, 14.X.2006 (IBUNAM: CNIN: MA823); Guerrero, 22.IX.2022 (CEUAEM-MAN-159, CEUAEM-MAN-160); Guerrero, 2024 (CEUAEM-MAN-161, CEUAEM-MAN-162). Jalisco: 13 specimens (11 ♂, 2 ♀). Eleven ♂: La Huerta, Estación de Biología Chamela (UNAM), 19°29’55.2” N, 105°2’39.5” W, 01.VIII.1981 (IBUNAM: CNIN: MA426); 10.V.1980 (IBUNAM: CNIN: MA802, MA804); 24.V.1980 (IBUNAM: CNIN: MA803); 10.VIII.1981 (IBUNAM: CNIN: MA805-MA810); 01.IV.1981 (IBUNAM: CNIN: MA811). Two ♀: La Huerta, Estación de Biología Chamela (UNAM), 19°29’55.2” N, 105°2’39.5” W, 14.X.1990 (IBUNAM: CNIN: MA723); 08.VIII.1975 (IBUNAM: CNIN: MA1560). Morelos: 23 specimens (12 ♂, 11 ♀). 12 ♂: Jojutla, EESJ, 18°36’49.8” N, 99°13’50.2” W, 957 m asl: 08.III.2022 (CEUAEM-MAN-013); 07.X.2018 (CEUAEM-MAN-014); 21.XI.2018 (CEUAEM-MAN-015); 20.XI.2019 (CEUAEM-MAN-021); Jojutla, 18°36’41.6” N, 99°10’44.7” W, 894 m asl, 21.X.2023 (CEUAEM-MAN-140); Morelos, 18°39’3.9” N, 99°2’20.5” W, 16.XI.1972 (IBUNAM: CNIN: MA519, MA520); Tepalcingo, Sierra de Huautla, 18°32’31.3” N, 98°56’9.4” W, 12.VI.2016 (IBUNAM: CNIN: MA943-MA946). Eleven ♀: Zacatepec, Galeana, 18°38’24.1” N, 99°12’39.6” W, 934 m asl, 22.IX.2019 (CEUAEM-MAN-017); Jojutla, EESJ, 18°36’49.8” N, 99°13’50.2” W, 957 m asl: 28.XI.2019 (CEUAEM-MAN-018); 22.XI.2019 (CEUAEM-MAN-019); 25.IX.2020 (CEUAEM-MAN-109); 13.XI.2020 (CEUAEM-MAN-110); 04.X.2023 (CEUAEM-MAN-129); Tehuixtla, 18°33’16.8” N, 99°16’12.8” W, 26.XII.2019 (CEUAEM-MAN-072); Yautepec, 18°53’12.8” N, 99°3’47.8” W, 1,216 m asl, 17.X.2023 (CEUAEM-MAN-138); Jojutla, 18°36’41.6” N, 99°10’44.7” W, 894 m asl, 27.X.2023 (CEUAEM-MAN-139); Cuautla, 18°48’59.4” N, 98°56’56” W, XI.1954 (IBUNAM: CNIN: MA905); Morelos, 18°39’3.9” N, 99°2’20.5” W, IV.1981 (IBUNAM: CNIN: MA906). Nayarit: 1 ♂, Bahía de Banderas, San Rafael, Arroyo de la Virgen, P. H. Aguamilpa, 20°50’19.5” N, 105°16’44.4” W, 734 m asl, 01.XI.1991 (IBUNAM: CNIN: MA824).

Geographic distribution: Stagmomantis hebardi is distributed in the states of Estado de México, Guerrero, Jalisco, Morelos, Nayarit, and Sinaloa (Fig. 12).

Remarks

Taxonomic delimitation within Stagmomantis based on male morphology has been a subject of study for decades. Notably, Saussure and Zehntner (1894) and Rehn (1935) considered the males of S. carolina and S. tolteca to be indistinguishable, grouping them within the “carolina group,” which currently corresponds to the subgenus Stagmomantis.

Morphological differences between the female originally described by Rehn (1935) and the females analyzed in this study are summarized in Table 4. Key distinctions include: a) the pronotum is proportionately wider in Rehn’s female compared to the slender pronotum in our specimens; b) wing shape and coloration differ significantly; Rehn’s specimen has wider forewings with a small, oval, light-colored stigma lacking a surrounding patch, whereas our specimens exhibit a stigma covered by a dark patch and surrounded by a white area covering nearly the proximal half of the wing; c) the abdomen in Rehn’s female is widely oval, whereas it is subfusiform (widening at the distal segments) in the females observed in this study; and d) the lobe-like structure between the 5th and 6th abdominal tergites, observed in all examined specimens, appears to be a diagnostic feature and a potential autapomorphy for this species.

Although Stagmomantis tolteca and S. hebardi are morphologically similar, significant differences in male genitalia provide reliable diagnostic characters. As noted by Arteaga-Blanco et al. (2016), the posterior process of the ventral phallomere (pda) in S. tolteca is strongly sclerotized, wide at the base, tapering distally into a spine, and directed sinistrally (Fig. 13a, blue). Conversely, in S. hebardi, the pda is corkscrew-shaped, twisted either sinistrally or dextrally (Figs. 1, 5, 13b, blue). Further comparisons revealed differences in other genitalic structures: the aafa in S. tolteca is triangular with an angulate apex (Fig. 13a, yellow), whereas it is rounded in S. hebardi (Figs. 5, 13b, yellow); the ssp in S. tolteca is more developed with a slightly angulate inferior margin (Fig. 13a, purple), compared to the small and triangular ssp in S. hebardi (Figs. 5, 13b, purple); finally, the pafa in S. tolteca is elongate and curved near the middle, with the apex directed inward (Fig. 13a, green), while the pafa of S. hebardi projects laterally, is less curved, and has a more rounded apex (Figs. 5, 13b, green). The paa is morphologically similar in both species (Fig. 13, red). Rehn’s (1935) illustrations of the male genitalia correspond more closely to the pda morphology described in the present study than to that illustrated in Arteaga-Blanco et al. (2016). Thus, the sclerotized structure of the pda serves as a robust diagnostic character to separate both species and supports the identification of our male specimens as S. hebardi, in accordance with Rehn (1935).

A remarkable feature observed in this study is the presence of a lobe-like structure on the female abdomen. While comparable structures are rare in Mantodea, a few instances have been documented. For example, Robinson and Robinson (1979) described a terminal abdominal structure in females of Acanthops falcata Stål, 1877, identifying it as a pheromone-releasing gland. Similarly, Schwarz and Glaw (2021) described a Y-shaped protrusible structure near the abdominal apex of Stenophylla lobivertex Lombardo, 2000, hypothesizing that it represents a pheromone-releasing gland more efficient than simpler structures performing the same function.

The precise function of the lobe-like structure identified in this study remains unknown; however, several hypotheses can be proposed. According to Robinson and Robinson (1979) and Schwarz and Glaw (2021), such structures may function as sex pheromone glands, as reported for A. falcata and S. lobivertex. However, we observed this structure in both sexes during the early developmental stages of S. hebardi, which suggests it may possess an alternative or additional function. Another hypothesis involves the release of prey-attracting pheromones (kairomones), similar to the aggressive mimicry observed in bolas spiders (Mastophora Holmberg, 1876), which mimic lepidopteran sex pheromones to capture prey (Yeargan & Quate, 1997). Finally, this structure might enhance crypsis, similar to strategies observed in the subfamily Vatinae. However, unlike Vatinae, which are characterized by fixed cuticular lobes on the legs or abdominal margins, the structure in S. hebardi is distinctively protrusible.

Acknowledgements

We thank Cristina Mayorga-Martínez for her support in reviewing the CNIN-IBUNAM; Michael R. Maxwell, Henrique M. Rodrigues, and Lohitashwa Garikipati for their comments on an earlier version of this work; Jason D. Weintraub for providing us high resolution images of the holotype and allotype of Stagmomantis hebardi. Aarón Emilio Vásquez Quintero was supported by a scholarship from the Secretaría de Ciencia, Humanidades, Tecnología e Innovación (Secihti; CVU 1315459).

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