In recent years, the study of wild silkmoths (Lepidoptera: Saturniidae) has increased exponentially due to the intense sampling effort and the use of molecular evidence for species delimitation, which led to the description of numerous new taxa especially from the Neotropic. Given these rapid advances, the checklist of the Colombian Saturniidae needs to be updated to cover the taxonomy, distribution, and diversity of these moths in the country. After an extensive review of literature, data repositories, and collections, an updated and comprehensive list of Saturniidae from Colombia is presented, including their occurrence status in each Colombian department. The checklist includes 7 subfamilies, 55 genera, and 790 taxa (766 in species rank) of Saturniidae in Colombia. Current distribution data show that the genus Winbrechlinia, the subgenus Darylesia, 379 species, and 18 subspecies are endemic to Colombia. Moreover, a dichotomic key to the Colombian subfamilies is provided. A few taxonomic changes are proposed based on a thorough taxonomic revision of the Colombian taxa. This revision also addresses the issue of outdated species names reported in the first checklist of Colombian Saturniidae (Amarillo-Suárez 2000) and excludes old records of taxa that are considered dubious for Colombia based on new evidence. By presenting an updated list of Colombian species, including the newly described taxa, this study aims at eliminating confusion stemming from outdated names and provides a useful resource for researching and conservating Saturniidae in Colombia. We wish to offer a common reference for future studies on the biodiversity and biogeography of moths in the Neotropical realm.

Bombycoidea, distribution, endemism, Neotropic, South America

“This great diversity of entirely American groups in the Saturniidae […] suggests that the group as a whole arose in the Western Hemisphere and no doubt in the American tropics.”

(Michener 1952: 371)

The Saturniidae, known as wild silkmoths, represent the largest family within the Bombycoidea superfamily and are found almost worldwide (Kitching et al. 2018). However, they are most diverse in the Neotropic region, where they originated (Regier et al. 2008; Rougerie et al. 2022). A global list of valid names for Saturniidae includes 3,454 species in 180 genera (Kitching et al. 2018), of which nearly 2,400 species are estimated to occur in the Neotropic (Decaëns et al. 2021). In the Neotropic, the Saturniidae family is divided into seven subfamilies: Arsenurinae, Ceratocampinae, Cercophaninae, Hemileucinae, Hirpidinae, Oxyteninae, and Saturniinae, the latter being the only cosmopolitan, while the others are exclusively from the New World (Lemaire and Minet 1998; Rougerie et al. 2022). Lemaire revised four subfamilies in America: Saturniinae (Lemaire 1978b), Arsenurinae (Lemaire 1980), Ceratocampinae (Lemaire 1988b), and Hemileucinae (Lemaire 2002). Before publishing his latest three-volume monograph (Lemaire 2002), he compiled a preliminary list of 921 species for the Neotropic (Lemaire 1996). Amarillo-Suárez (2000) reported a total of 183 species in the first checklist of the Colombian Saturniidae, underestimating the diversity of the Hemileucinae, which was later revised by Lemaire (2002). However, both Lemaire (1996) and Amarillo-Suárez (2000) excluded the subfamilies Cercophaninae and Oxyteninae from their checklists. These most basal subfamilies were described as families by Jordan (1924) and then assigned to Saturniidae based on morphological characters (Minet 1994), later confirmed by phylogenetic evidence (Regier et al. 2008). Furthermore, the diversity of Cercophaninae and Oxyteninae has increased enormously due to recent species descriptions (Brechlin and Meister 2014b; Brechlin et al. 2014; Brechlin 2020e, 2021j, 2021o, 2021e, 2023i).

Despite the growing interest and popularity of the Saturniidae (Janzen 1984; Howse and Wolfe 2011; Meister 2011; Basset et al. 2017; Rubin et al. 2018), the literature dealing with the distribution of Colombian Saturniidae is limited. A few ecological studies included lists for specific Colombian localities: Río Ñambí Natural Reserve, Barbacoas, Nariño (Amarillo-Suárez 1997a); San José del Palmar, Chocó (Decaëns et al. 2003b); Tambito Reserve, El Tambo, Cauca (Muñoz and Amarillo-Suárez 2010); Gorgona Island National Park, Guapí, Cauca (Calero-Mejía et al. 2014); and Utría National Park, Chocó (Prada-Lara et al. 2019) in the Chocó biogeographic region; Albania, Caquetá, in the Orinoquía region (Racheli and Vinciguerra 2005); and Arcabuco and Quipama, Boyacá, in the Andean region (Decaëns et al. 2007). However, these local checklists have lost their validity due to recent taxonomic advances.

In the last decade, approximately 1,500 new species and subspecies of Saturniidae have been described globally (Kitching et al. 2018). This enormous number is mainly due to DNA studies (Hebert et al. 2003; Padial et al. 2010). Despite being controversially discussed (Will and Rubinoff 2004; Will et al. 2005; Peigler 2013), DNA barcoding is now widely recognized as a tool for revealing cryptic Lepidoptera species (Decaëns and Rougerie 2008; Vaglia et al. 2008; Gibbs 2009; Hausmann et al. 2009; Decaëns et al. 2021; Moraes et al. 2021). Today, integrative taxonomy combines morphological features, geographic distribution, COI barcode studies (Silva-Brandão et al. 2009 provide an extensive review on the subject), and nuclear markers to increase resolution (Rougerie et al. 2012). Unsurprisingly, many of the newly described Saturniidae are distributed in Colombia due to the variety of ecosystems (van der Hammen and Rangel 1997), biodiversity hotspots (Myers et al. 2000), and the recent sampling boosting, especially at high elevations, in previously inaccessible localities, given the limitations due to the control of the territory by various armed groups (Brechlin 2016f; Murillo-Sandoval et al. 2020). Since 2008, many descriptions of neotropical Saturniidae have been published in the Entomo-Satsphingia journal, including two major revisions of the genera Hylesia Hübner, 1820 (Brechlin et al. 2016a) and Janiodes Jordan, 1924 (Brechlin 2020e). The most striking result was the description of the genus Winbrechlinia Brechlin, 2016, endemic to the cloud forests and páramos of the Sierra Nevada of Santa Marta in northern Colombia (Brechlin 2016f, 2018m, 2020d).

The geographical complexity of Colombia makes its fauna extraordinarily diverse and highly endemic (Bernal and Lynch 2008; Avendaño et al. 2021; Bota-Sierra et al. 2021; Pérez-Escobar et al. 2022). The topography is characterized by three main parallel Andean mountain ranges, known as Cordilleras, located within the “Tropical Andes” biodiversity hotspot (Myers et al. 2000). The Andean region is bordered to the east by the Orinoquía and Amazon regions and to the west by the Chocó biogeographic region, another biodiversity hotspot (Myers et al. 2000). The three Andean mountain ranges are separated by the two large, major streams of Colombia, the Cauca River, which flows northward between the Western and Central Cordilleras, and the Magdalena River, which divides the Central and Eastern Cordilleras. After emerging from the Colombian Massif, these two rivers join and descend to the Caribbean Sea. On the Caribbean’s margin there is the Sierra Nevada de Santa Marta, whose highest elevations are the tallest peaks (5775 m) in Colombia. This more recently formed area (Gómez et al. 2021) is also an essential hotspot of biodiversity that hosts many endemic species (Myers et al. 2000; Brechlin 2016f). The geological history of Colombia has allowed the discontinuous isolation of species, favoring allopatric speciation (Vargas et al. 2023) through the increase in the availability of new niches (Purser 2015; Hazzi et al. 2018). Notably, páramos have been described as “islands” with flickering connectivity (Flantua et al. 2019) and the fastest diversification rate (Madriñán et al. 2013). For these geographical reasons, Colombia probably has one of the most diverse and endemic Saturniidae faunas in tropical America (Lemaire and Venedictoff 1989; Decaëns and Rougerie 2008; Jiménez-Bolívar et al. 2021).

In light of the advances made in the last two decades, the first checklist of the Colombian Saturniidae (Amarillo-Suárez 2000) is outdated. No systematic work has been carried out to date to review the taxonomy and distribution of the Colombian Saturniidae taxa, considering the contributions of checklists for some specific Colombian localities and including the vast cryptic diversity revealed by the use of DNA barcoding and the recent tremendous sampling effort. At the subfamilies level, the diversity of Cercophaninae and Oxyteninae has previously been underestimated since the contribution of these subfamilies is reviewed here for the first time, taking as a starting point the preliminary checklist by Comoglio and Brechlin (2021). Likewise, considering the separation of Hirpidinae from Hemileucinae (Rougerie et al. 2022), updating the taxonomic key for subfamilies is necessary.

This work aims to present an updated checklist of the known Saturniidae from Colombia, considering the many new descriptions and revalidation of taxa, and to clarify the taxonomic confusion that these taxonomic advances may have produced. An updated dichotomous key was also elaborated for the subfamilies distributed in the country. Furthermore, a few taxonomic changes are proposed and discussed, following the criteria of delimitation of species currently used in the literature (Brechlin and Meister 2011a, 2011c; Brechlin et al. 2016a; Bénéluz 2021; Brechlin 2022i). Based on an extensive review of literature, data repositories, and collections, this checklist is the first one for Colombia (published on 6 August 2021 as a preprint) which also includes the subfamilies Cercophaninae and Oxyteninae, and that extensively covers the Hemileucinae. Species and subspecies considered endemic for the country are highlighted here, thanks to an extensive review of the distribution data of all the Neotropical taxa considered in this study. Taxa excluded from our Colombian checklist are discussed in detail, comparing the present knowledge with the old records found in the literature. In addition, taxa with potential distribution in Colombia, but whose presence has not been confirmed yet, are discussed on the basis of their current distribution data. After the preprint of this study (Comoglio and Brechlin 2021) was made available, another checklist of the Colombian Saturniidae was published by Jiménez-Bolívar et al. (2021), whose results are thoroughly reviewed here.

This checklist is mainly the product of recent sampling efforts, which have led to the description of many Saturniidae taxa for Colombia, and a literature review of articles, species descriptions, taxonomic revisions, and records available on BOLD (Hebert and Ratnasingham 2007) until 15 June 2023. In addition, both authors contributed with data from the collections they are currently curating to add new records which have not been previously published elsewhere. The first author examined specimens in the Entomological Collection of the “C.J. Marinkelle” Natural History Museum, Universidad de los Andes, Bogotá, Colombia (ANDES-E). The second author conducted a comprehensive review of his Research Collection, Pasewalk, Germany (CRBP), which includes type material of more than 530 Colombian taxa and is extensively barcoded (“BC-RBP” in BOLD). The specimens deposited in the main national collections, such as the Institute of Natural Sciences, National University of Colombia, Bogotá (ICN-MHN); the “Francisco Luis Gallego” Entomological Museum, National University of Colombia, Medellín (MEFLG); the Javeriano Museum of Natural History “Lorenzo Uribe, S.J.”, Pontificia Universidad Javeriana, Bogotá (MPUJ), and the National Taxonomic Collection of Insects “Luis María Murillo”, Mosquera (CTNI), were examined by Amarillo-Suárez (2000), Clavijo-Giraldo and Uribe (2019), and Jiménez-Bolívar et al. (2021). However, some of these collection records are provided ignoring the current nomenclature that is extensively reviewed and discussed in this paper. Some interesting records are found in the entomological collection of the Alexander von Humboldt Institute, Villa de Leyva, Colombia (IAvH-E), and have DNA barcodes in BOLD for some additional taxa. Barcodes are essential for the verifiable identification of specimens, especially those belonging to species complexes (Decaëns and Rougerie 2008; Decaëns et al. 2021). The implications of using raw data available in repositories without curation or expert identification, such as some records shown in Jiménez-Bolívar et al. (2021), have been debatable (Zizka et al. 2020) and are also discussed in this work, which is also a tool to improve the identification of Colombian Saturniidae taxa.

The dichotomous key to subfamilies covers only Colombian species (e.g., Janiodes species for the Cercophaninae). It is based on morphological studies of adults achieved by Michener (1952) and Lemaire and Minet (1998), with the addition of the analysis of the adult morphology of the Arsenurinae (de Camargo et al. 2009), Ceratocampinae (Balcázar-Lara and Wolfe 1997), Cercophaninae (Jordan 1924; Minet 1994), Hemileucinae (Lemaire 2002), and Hirpidinae (Rougerie et al. 2022). The morphological characters used in the literature were systematized and corroborated by direct examination of the specimens in the reviewed collections. Finally, an updated dichotomous key for Colombia was created, modifying the last global subfamily key presented by Lemaire and Minet (1998) and including the recently described subfamily Hirpidinae (Rougerie et al. 2022).

The taxa’s higher classification, names, and authority follow the Bombycoidea global checklist (Kitching et al. 2018) with some additions due to the most recent descriptions and revalidation of old taxa names found in the literature, and especially the recent phylogenomic analysis by Rougerie et al. (2022). The main list is shown alphabetically, ordered by subfamilies, tribes, subtribes, genera, subgenera, species, and subspecies. The occurrence in each Colombian department is provided for each species and subspecies. The Colombian departments are abbreviated as follows: Amazonas (Am), Antioquia (An), Arauca (Ar), Boyacá (By), Caldas (Cl), Caquetá (Ca), Casanare (Cn), Cauca (Cc), Cesar (Ce), Cundinamarca (Cu), Chocó (Ch), Guainía (Gn), Guaviare (Gv), Huila (Hu), La Guajira (Gj), Magdalena (Ma), Meta (Me), Nariño (Na), Norte de Santander (NS), Quindío (Qu), Putumayo (Pu), Risaralda (Ri), Santander (St), Tolima (To), Vaupés (Va), Valle del Cauca (Vl), and Vichada (Vi). Endemic species and subspecies are highlighted. Taxa previously reported in Colombia that were excluded from the list are discussed separately. According to their current distribution data, a list of taxa with potential occurrence in Colombia is also presented.

A total of 843 specimens in ANDES-E and more than 15,000 specimens in CRBP were examined, and 2,854 barcodes of Colombian Saturniidae specimens were retrieved from BOLD. To show the updated checklist, it is necessary to propose some taxonomic changes that are summarized here and then discussed, compared with previous studies, and interpreted in light of the criteria currently used to delimit species. An identification key for the 7 Saturniidae subfamilies distributed in Colombia is also presented as a result of this review. An updated national list for the family is then presented, including the distribution of the taxa in the Colombian departments, highlighting the endemic species, and reporting the current evidence as a barcode and related bibliographic references. The taxa excluded from this updated list are discussed below, comparing our results with the old records found in the literature. In addition, species with a possible distribution in Colombia, whose presence has not yet been confirmed, are discussed based on their current distribution data.

During the taxonomic review carried out in this study, we found taxa whose delimitation criteria between species have lost their validity and taxa that need to be revalidated conservatively, according to current knowledge. The proposed taxonomic changes are reflected in the checklist, and their validity and interpretation are discussed in detail below. The proposed taxonomic changes are summarized here. The following taxa, hitherto treated as subspecies, are raised to species status: Arsenura lemairei L. Racheli & T. Racheli, 1998, stat. nov. from A. thomsoni Schaus, 1906, Copiopteryx banghaasi Draudt, 1930, stat. nov. from C. semiramis (Cramer, 1775), and Rhescyntis norax Druce, 1897, stat. nov. from R. hippodamia Druce, 1897. A new combination is proposed in this context: Copiopteryx banghaasi andensis (Lemaire, 1974), comb. nov. In addition, Bathyphlebia aglia gschwandneri Schawerda, 1925, stat. nov. is removed from its synonymy with B. a. aglia C. Felder & R. Felder, 1874 and here treated as a subspecies of the latter. Grammopelta cervina Rothschild, 1907, stat. rev. and Copaxa ignescens Lemaire, 1978, stat. rev. are reinstated and removed from their current synonymies with G. lineata (Schaus, 1906) and C. niepelti Draudt, 1929, respectively. Furthermore, Rothschildia equatorialis bogotana Rothschild, 1907, stat. rev., comb. nov. is reinstated as a subspecies, but now of equatorialis Rothschild, 1907 instead of orizaba (Westwood, 1853). A new synonymy is proposed: Rhescyntis hippodamia colombiana (Bouvier, 1927), syn. nov. is now treated as a subjective junior synonym of R. norax Druce, 1897.

Key to subfamilies

A dichotomous key for the seven subfamilies of Saturniidae found in Colombia is presented below, excluding the genera of these subfamilies not found until now in Colombia. The mentioned characters account only for the external morphology. Therefore, identification at the subfamily level is generally immediate.

1	Male with antennal flagellum dorsally scaled to the apex and lateroventral orientation of the rami; bipectinate antennae in both sexes; proboscis present (Fig. 1A)	Oxyteninae Jordan, 1924
–	Male with antennal flagellum unscaled, at least for most of its length; lateral or laterodorsal orientation of the rami	2
2	Hindwing with strongly indicated crossvein (R) between Sc and upper edge of the discal cell; ventrally spined tarsi; forewings with at least one dark discocellular spot; small to medium size	3
–	Hindwing with crossvein nearly always absent or faintly indicated; hindwings with tails, hyaline discal spots, or eyespots; proboscis always absent	4
3	Yellow proboscis, strong and coiled up; bipectinate antennae in males, simple in females; males with a large foretibial epiphysis, whose median area looks internally notched; butterfly-like body shape and pierid-like wing shape (Fig. 1B)	Cercophaninae Jordan, 1924
–	Proboscis absent; orange quadripectinate antennae in males, simple in females; frons slightly convex at sides; orange-brown coloration	Hirpidinae Rougerie, 2022
4	Presence of solid bristles on pilifers or the clypeal margin between pilifers; hindwings usually with tails, longer in males; dull brown coloration; medium to large size (Fig. 1C)	Arsenurinae Jordan, 1922
–	Pilifers and clypeal margin without bristles	5
5	Frons convex at sides so that lateral sutures are hidden in an anterior view and antennal cones (short ventral protuberances on flagellomeres) simple; distal section of antenna devoid of rami; general body shape sphingid-like (Fig. 1D)	Ceratocampinae Harris, 1841
–	Frons flat at sides or, if convex, antennal cones multiple	6
6	Antennae, when quadripectinate, with bases of rami invariably well separated; thorax with the anterior area of mesoscutum lacking middorsal projection; in the forewing, when the discal cell is closed, the base of M1 arising closer to M2 than Rs or about midway between M2 and Rs; segments of labial palpi not fused; hyaline discal spots on both forewings and hindwings (Fig. 1E)	Saturniinae Boisduval, 1837
–	Antennae, when quadripectinate, with apical ramus of a segment usually adjacent to the basal ramus of next segment; if rami separate, mesoscutum with an anterior middorsal projection or forewing with the base of M1 distinctly closer to Rs than to M2 (or even stalked with Rs); antennal cones present and simple; labial palpi occasionally fused; hindwings usually with eyespots (Fig. 1F, G)	Hemileucinae Grote & Robinson, 1866

Figure 1. 

Representative taxa of the diversity of the Colombian Saturniidae subfamilies A Therinia terminalis (Oxyteninae) B Janiodes lavcarchensis (Cercophaninae) C Copiopteryx jehovah (Arsenurinae) D Eacles niepelti (Ceratocampinae) E Copaxa sapatoza (Saturniinae) F Automeris alticarchensis (Hemileucinae) G Winbrechlinia sinjaevi (Hemileucinae).