﻿Mapping the distribution of armored harvestmen (Opiliones, Laniatores) in Colombia: updated list of species, taxonomic contributions, and insight of diversity in protected areas

﻿Abstract Colombia is a biodiverse country with 1,486 protected areas covering almost 50 million hectares, yet little is known about the biodiversity they harbor, particularly in terms of harvestmen (Arachnida: Opiliones). This study provides a comprehensive updated summary of the armored harvestmen (Laniatores) species found in Colombia with a list of 173 nominal species, focusing on the Laniatores fauna found in protected areas and its diversity is detailed and analyzed. Maps with all records associated with ecoregions and protected areas in Colombia are presented. Additionally, three new Laniatores species are described here: Phalangodusandresisp. nov. from the department of Cundinamarca, and Ventrifurcaphallainasp. nov. and Ampycellafortunatasp. nov. from the department of Valle del Cauca; and a new family assignment is proposed for Paraphalangodus Roewer, 1915, placing it in the family Nomoclastidae. Information available on Laniatores in the National System of Protected Areas is still scarce and promoting strategies to facilitate the regulatory procedures for collecting specimens in these areas and increasing investment in basic science projects, are suggested to improve the understanding and study of the Laniatores fauna and other invertebrates in Colombia. Finally, a chronicle and timeline set of figures of species of Laniatores from Colombia, described by various authors during three periods, is given.


Introduction
Colombia is one of the most biodiverse countries in the world (Andrade-C 2011). This diversity is represented and legally protected in 1,486 conservation units (covering 49,884,326.77 hectares), under the figure of National System of Protected Areas (RUNAP 2022). These areas allow ecological self-regulation, their ecosystems have not been substantially altered by human exploitation or occupation, and their biodiversity is under management (CONPES 2021). Although Colombia is one of the countries in South America with the largest number of areas under conservation (CONPES 2021), knowledge about the species it harbors is scarce, especially in terms of harvestmen (Arachnida: Opiliones). Colombia ranks third in South America in terms of the number of recorded Opiliones species, only behind Brazil (1,010) and Venezuela (392) (Villarreal et al. 2021a). The first effort to list the arachnid species of Colombia was made by Flórez and Sánchez-C (1995), summarizing 77 species of harvestmen. Later, all records and distribution data on Colombian Laniatores, which constitute the vast majority of Opiliones in the country, were compiled by Kury (2003) into an excellent New World Laniatores catalog. Taxonomic studies on harvestmen have become increasingly frequent in the Neotropics, especially in Colombia, where numerous genera and species have been recently described or recorded (e.g., Villarreal and Rodríguez 2006;García 2014;Villarreal and García 2016;Pinzón and Pinto-da-Rocha 2020;Villarreal and García 2021). More recently, a general diagnosis of the current knowledge of the group in Colombia was published (Perafán et al. 2013), recording a total of 162 Opiliones species in the country. Minor local inventory works mention 170 (García and Medrano 2015) and 186 (Barriga et al. 2019) species of Opiliones in Colombia. Only two opilionofaunal inventory and/or ecological studies have been conducted in Colombia to our knowledge so far (García and Medrano 2015;De Moya et al. 2021). García and Medrano (2015) carried out the guide of Opiliones of the Reserva Natural Rio Ñambí, a protected area on the coastal foothills (department of Nariño) (CONPES 2021), recording 12 families and a wealth of undescribed species. More recently, De Moya et al. (2021) studied the diversity of Opiliones by altitude in Sierra Nevada de Santa Marta (department of Magdalena), an area of high conservation importance due to its significant climatic variability and high rates of endemism, recording nine families. Studies focused on analyzing the composition of neotropical harvestmen in protected areas are not frequent. These have been carried out mainly in Brazil, among them, Soares and Soares (1970) describe seven species for the Itatiaia Biological Station (Rio de Janeiro); Adis et al. (2002) record seven species for the Ducke Reserve (Amazonia), and Bragagnolo and Pinto-da-Rocha (2003) recorded 52 species in the Serra dos Órgãos National Park (Rio de Janeiro). A recent precedent was made by Guerrero (2019), who recorded 17 species from 30 protected areas in the province of Buenos Aires (Argentina). However, like the situation in Colombia, he concludes that many important areas still remain for the conservation of diversity that have yet to be surveyed to understand their harvestmen composition.
Since the publication of the New World Laniatores catalog by Kury (2003), more than 20 years ago, Colombia has undergone numerous changes and additions to its opilionofauna. This is partly due to the appearance of a growing group of enthusiastic local and regional arachnology researchers who have turned their interest to the fauna of this country, as well as the expansion of its biological collections and collaboration with other renowned institutions. Currently, Colombia has 162 species of armored harvestmen (Opiliones: Laniatores) recorded, belonging to 12 families of which 46 species have been recorded or described after 2003. Therefore, we consider it opportune to provide an updated and detailed summary of the taxonomic changes in the Colombian laniatorean fauna after 2003.
The purpose of this paper was to provide an updated list of species and summarize the taxonomic changes and additions that occurred after 2003. Additionally, we aim to relate the distribution of Colombian species with the biogeographical areas proposed by WWF, discuss the presence of species in the country's National System of Protected Areas (NSPA), and present the descriptions of two new cranaid species from the departments of Cundinamarca and Valle del Cauca as well as one new gonyleptid species from the department of Valle del Cauca.

Taxonomic background for Phalangodus, Ventrifurca, and Ampycella of Colombia
The neotropical family Cranaidae has attracted attention from taxonomists in recent years; nevertheless, the definition of generic or suprageneric groups in this family remains unsatisfactory Villarreal and García 2016). Although some groups have been reviewed, leading to the resolution of generic synonyms, new combinations or redefinitions, improper placements, as well as descriptions of new species (e.g., Orrico and Kury 2009;Pinto-da-Rocha and Bonaldo 2011;Hara et al. 2014;Villarreal et al. 2015;Villarreal and García 2016;Hara et al. 2017), much work remains to be done.
Only three works have analyzed the phylogenetic relationships within the family Cranaidae and the genus Phalangodus Gervais, 1842 has only been studied in a phylogenetic context in an analysis of the familial relationships of Gonyleptoidea , where it appears as sister group to the other four representatives of the family. Despite this, the taxonomy of Phalangodus is relatively well understood, with a total of six species described, five of them known from Colombia (Villarreal and García 2016). A similar case exists for the genus Ventrifurca Roewer, 1913, which was recently reviewed , revealing some synonyms and the discovery of a new species from the department of Quindío. Currently, three species of Ventrifurca are known from Colombia, distributed in the western Andes.
Two species have been recorded from Colombia, belonging to the family Ampycidae, Ampycus telifer (Butler, 1873), and Licornus tama Villarreal & Kury, 2012(García 2014. This family lacks taxonomic revisionary work, and its genera are poorly delimited or defined. Recent works have contributed to better understanding the diversity of some genera within Ampycidae (e.g., Tourinho and Mendes 2014;Hara et al. 2017) or described new species (Villarreal and Kury 2012). The genus Ampycella Roewer, 1929 includes two Andean species from Ecuador , vaguely described and illustrated and whose genital morphology is unknown. This paper describes three species: two new cranaid (Phalangodus and Ventrifurca) and one ampycid (Ampycella, including the first image of the male genitalia for the genus), collected in the departments of Cundinamarca in the Central Mountain Range in the Colombian Andes and the department of Valle del Cauca in Chocó Biogeographic region.

Taxonomy
Individuals of each species were photographed using a Leica M205C stereoscope attached to a Leica DFC450 digital camera, a Wild Heerbrugg stereoscope attached to a Nikon COOLPIX P900 and a Wild Heerbrugg microscope attached to a HAYEAR 2307 digital camera. The multiple resultant images at different focal planes were combined with Combine ZP Suite software (Hadley 2015) to increase the depth of field and were thereafter edited in Photoshop CC 2014 software. Drawings of the species were made using Illustrator CC 2017 and Inkscape 1.2.2 softwares (Harrington 2004(Harrington -2005. To color descriptions the standard names of the 267 Color Centroids of the NBS/IBCC Color System were used as named in Centore (2016). Male genitalia were studied using standard methods for this structure (Acosta et al. 2007).
Morphological terminology and patterns of taxonomic description follow Villarreal and García (2016) with slight modifications; integumentary ornamentation follows DaSilva and Gnaspini (2010); terminology for chaetotaxy of penis ventral plate follows Kury and Villarreal (2015) with the modifications proposed by Villarreal and García (2016) for the genus Phalangodus; the ovipositor morphology follows Villarreal and García (2016); terminology of dorsal scutum outline types follows Kury and Medrano (2016).
The first-order administrative divisions of Colombia (departments) are underlined. Maps were made using ArcGIS ® 10.1 software (ESRI 2022). Colored areas represent WWF Terrestrial Eco-regions of the World (Olson et al. 2001), here abbreviated as WWF.

Species inventory
A comprehensive list of all described Laniatores species occurring in Colombia was compiled, based on the bibliography published until April 2023, including all valid species of Laniatores recorded for Colombia. The family and superfamily classification follows Kury et al. (2022). The logonymic information is updated only for Colombian species that were described, recorded, or underwent nomenclatural changes after New World Laniatores catalog (Kury 2003). The taxonomic references in the logonymy of the listed species were not considered in the References section.
A list of all protected areas under the NSPA jurisdiction, which harbor records of Laniatores species, was compiled. Only areas with some form of public regulation for their protection were considered. For each protected area, the families and species of armored harvestmen recorded were listed (Table 1). The source of the species data can be seen in the Suppl. material 1. In the species list, we utilize two types of dashes: the n-dash (-), used to indicate the reference of the original description, and the m-dash (-) to introduce each new subsequent bibliographic reference following the original description. After each species name, the m-dash is placed to separate a new citation.

Species descriptions
Male holotype (ICN-Ao-1908). Dorsum (Figs 1A,B,3A,B). DS outline type alpha, with bulge longitudinally asymmetric widest at scutal groove II, lateral borders with granules only on the middle region. Carapace with few granules on the anterolateral and posterior region. Ocularium high, without median depression, with a paramedian pair of sharp tubercles and granules close to the eyes. Integumentary dome of ozopore raised and conspicuous. Abdominal scutum well delimited, divided into four well-marked scutal areas; I divided into left and right halves by invasion of the scutal area II; I and II granulated, with a pair of conspicuous medial tubercles, one tubercle on each side; III with a pair of paramedian acuminate spines and densely granulated; IV divided, with a row of six or seven granules on each side. Posterior border of the DS slightly convex and with a row of granules. Free tergites I-III with a row of granules.
Venter. Coxa I with a row of large tubercles of different sizes; II longer than I and III, with two median rows of low tubercles, the anterior one more conspicuous; III densely covered with irregular rows of small tubercles and with the posterior border sigmoid; IV strongly backward, with a median row of tubercles in the medium area and lateral border, and small tubercles densely distributed. Stigmatic area with a row of small tubercles on posterior border and minute granules sparsely distributed. Stigmata large, oval, and oblique. Free sternites with a row of small granules.
Chelicerae (Figs 2A,3C). Segment I with well-defined bulla, with dorsomesal tubercles and a row of three large tubercles in the ectal region. Segment II swol- len. Fixed finger with a proximal narrow and low tooth, a large gap and three subdistal median teeth. Movable finger with a proximal wide and low tooth, one large tooth and the distal inner surface irregularly dentate. Mesal side of the base of the fixed finger and near the base of the movable finger with setiferous tubercles of different sizes.
Pedipalps (Figs 2B, C, 3D-F). Trochanter with a dorsal pair of paramedian tubercles. Ventrally with large bifid tubercle in distal portion. Femur slightly compressed, dorsally curved, and ventrally straight in lateral view, with dorsoectal distal row of granules, one dorsal, and one ventral row of large forward projected tubercles (the apicalmost of the ventral row bifid and thicker than the remaining), the ventrodistal portion unarmed. Mesal and ectal faces without large tubercles. Patella short, cylindrical, and curved, with a dorsal row of short tubercles and small dorsodistal granules. Tibia dorsally granulated; tibia mesal IiIi (3>1>2=4), ectal IiiIi (4>1>5>2>3). Tarsus dorsally granulated, tibia with spines only in the distal portion, mesal Ii, ectal IiIiI (3>1>5>4>2). Claw proximally swollen. Legs (Figs 1A, 2D-F, 3B, 3G-J). Coxa I and III smooth; II with one dorsal tubercle; IV with one dorsodistal domed large tubercle, and a row of large tubercles in the prolateral border. Trochanters I-III unarmed; II with a prolateral row of small tubercles; III with one retrodistal tubercle and one group of small prolateral tubercles; IV with small prolateral granules sparsely distributed and one row of small tubercles in the retrolateral border. Femora I-III straight and with longitudinal rows of granules; IV sub-straight, densely granulated, ventrally with one large subdistal tubercle hook-like shaped and one distal rounded trifid tubercle. Ratio Fe IV/DSL = 1.85. Patellae I-IV granulated. Tibiae I-V straight and densely granulated, unarmed. Metatarsi I to IV with rows of small granules, I-III unarmed, IV with a ventro-distal border pair of spines. Tarsi III and IV with subparallel smooth claws and tarsal process. Penis (Fig. 4). VP subsquare, with proximal and distal portion delimited by a medial constriction, and lateral rounded lobes in the proximal portion; distal border slightly concave. Glans + Stylus columnar, glans with some folds at the base, stylus normally thickened, shorter than the glans, substraight; stylar caps ring-shaped. MS-A/B groups indistinguishable from each other, located on the proximal portion of the VP, with an increase in setae and composed of three to four pairs; MS-C/D groups located in the distal portion of the VP, separated from MS-A/B by a small gap, the setae rearranged into two irregular rows, one Female (ICN-Ao-1909) (Fig. 1C, D). Differing from male by: ocularium slightly narrower; carapace shorter; coda wider (DS outline type alpha-keyhole); tubercles of area III slightly higher. Chelicerae non-hypertelic, with movable finger thinner. Pedipalpal femur lower and thinner in lateral view. Stigmatic area shorter. Genital operculum wider. Trochanters III and IV narrower; femur IV thinner, without large ventral-subdistal tubercle with hook-like shape and without a ventrodistal rounded trifid tubercle. For color differences, see the color description of the male.
Ovipositor. Dorsal lobes (dl) and ventral lobes (vl) rounded, with four and two pairs of large, acuminated, single-tipped setae, respectively. The dl with three pairs of dorsal setae (ds) distally located and one pair basally located. Lateral region of the ovipositor with one pair of dorso-lateral setae (dls).
Distribution. Known only from the type locality. Etymology. The species is named in honor of our colleague and friend, the arachnologist Andrés F. García, who has greatly enriched the field's knowledge of Opiliones in Colombia and described the vast majority of species within the genus Phalangodus; moreover, he was the collector of the type series for this species. Diagnosis. Ventrifurca phallaina sp. nov. can be distinguished from V. abnormis by the scarce ornamentation of the mesotergal areas I to III; also, can be distinguished from all other species of the genus by the ornamentation pattern of the yellow tubercles on dorsal scutum, restricted to the lateral of carapace and posterolateral portion of DS (Figs 5A, B, 6A, B, 9A, B); the presence of a yellow spot on the anterolateral portion of the carapace; the shape and size of posteroventral projections on the stigmatic area, simple and short (Figs 5C, 6C), instead of bifid and globular (V. albipustulata) or large and digitiform or curve (V. caffeinica and V. abnormis); and the absence of yellow tubercles on the mesotergal area III, behind the paired spines (present in V. albipustulata and V. caffeinica). The genital morphology is very similar to V. albipustulata, with slight differences in relation to the shape of the concavity of the truncus (Fig. 8). with yellow rounded tubercles from carapace to area I, following outline of scutum. Carapace with three or four tubercles on each side of the anterior border. Ocularium low, armed with a paramedian pair of acuminate forward inclined tubercles and some small tubercles near the eyes. Integumentary dome of ozopore raised and conspicuous. Mesotergum well delimited, divided into three well-marked areas: area I divided into two roughly ellipsoidal halves by longitudinal groove, with a paramedian pair of large tubercles and four small tubercles; II penetrating subtly into I, with a median transverse row of eight small tubercles; III with a pair of paramedian acuminate high subparallel spines, located near the posterior margin; vestigial groove between III and IV present, partially blurred and located just behind the spines. Posterior border of DS straight. Free tergites I-III with a pair of paramedian large granules, I to II with one to three smaller laterodistal granules on each side; III armed with a posterior row of 11 tubercles, the paramedian pair much stouter.
Venter (Figs 5B, C, 6B, C). Coxa I with a row of tubercles of different size; coxa II longer than coxae I and III, with a cluster of distal tubercles of different size; coxa IV strongly backward. Stigmatic area with minute granules sparsely distributed, posterior border with posterodistal process very small. Stigmata large and oval.
Chelicerae (Figs 5D,6D). Segment I with well-defined bulla, with a row of four or five tubercles in ectal region. Segment II slightly swollen, with a row of large setiferous tubercles near to the base of the movable finger and a few mesal setiferous tubercles. Fixed finger with three medial teeth, decreasing in size. Movable finger with one large and wide medial tooth, accompanied by two flat, low teeth on each side.
Pedipalps (Figs 6E,F,7). Trochanter with tubercles, in dorsal and ventral region. Femur cylindrical, slightly elongated, armed with a longitudinal row of four tubercles in dorsal side and three tubercles in ventral side. Patella short, cylindrical, and curved, with row of tubercles in dorsal view, one ectodistal and two parallel rows of tubercles in mesal view tubercles.
Legs (Figs 5A-C, 6A, B, G, H). Coxae I and II with one anterior row of large tubercles; III with two rows and IV with two diagonal rows. Trochanter I with two prolateral, two retrolateral and four ventral tubercles; II with two prolateral, two retrolateral and five ventral tubercles; III with four prolateral, two retrolateral and four ventral tubercles; IV with one prolateral, one retrolateral and five ventral tubercles, more conspicuous than those from the other trochanters. Tubercles of trochanter IV more conspicuous than the others. Femora I to III straight, with longitudinal rows of granules; IV curved in its proximal portion, with seven longitudinal rows of tubercles, the retrolateral row with large proximal tubercles, and one large subdistal tubercle on the same row. Patellae I-IV with small tubercles. Tibiae and metatarsi I-V straight and granulate, unarmed. Claws III and IV smooth, with two minute proximal ventral protuberances. Ratio Fe IV/DSL = 1.35. Tarsal process present. Tarsal counts 7(3)-7(3), 14(3)-14(3), 7-7, 7-7.
Penis (Fig. 8). Ventral plate (VP) with concave lateral margins, with subdistal constriction and the distal corners ear-shaped, apical wide u-shaped cleft. Glans + Stylus complex surpassing the VP, glans columnar, with folds at the base, stylus normally thickened, sub-straight; stylar caps foliar-shaped, short. MS-A1-A2 straight, located at the distal portion of the VP; MS-B absent; MS-C1-C3 curve, close to each other, and located in the distal part of the VP; MS-D1 larger than MS-C group, curve and located medially to MS-C, MS-D2 absent; MS-E1-E2 small, located on the flange of the VP. Truncus with ventral concavity next to the VP.
Distribution. Known only from the type locality. Etymology. Phallaina is a Greek word that means whale. It is used in apposition as a reference to the humpbacked shape of the species in lateral view.
Diagnosis. Outline of DS type alpha; four mesotergal areas well defined and unarmed; DS with small yellowish tubercle on the lateral margins (Fig. 10A, B, F). Free tergites II and III with a posteromedial spine (Fig. 11B). Penis (only known to A. fortunata sp. nov.): VP sub-rectangular with medial constriction, giving it a guitar-shaped appearance, and distal border without cleft; MS-A1-A3 in the medial portion of the VP; MS-B absent; MS-C1-C3 distally located, similar to each other; MS-D1 as large as MS-A similar to each other, medially located and pointing distally. Stylus clearly differentiated from the glans, slightly curved in the distal portion (Fig. 12). Diagnosis. Ampycella fortunata sp. nov. can be distinguished from the other two species of the genus by the size ratio between the spines on the free tergites: spine on free tergite II larger than the spine on free tergite III (Figs 10A, B, F, 11A, B) (vs III larger than II). Additionally, it can be distinguished from A. frizzellae by having a low ocularium with two small tubercles (vs high ocularium with two erect spines) and the entire scutal area II (vs divided into three lobes); and from A. spiniventris for having the anterolateral part of scutal area II projected forwards, anterior to the middle part (vs anterolateral parts similar to the medial portion), and having longer dorsal spines on the coxa IV.

Ampycella fortunata
Description Male holotype (CIUQ-020635). Dorsum (Figs 10A, B, 11A, B). DS outline type alpha. Dorsal scutum widest at scutal area II; lateral borders of dorsal scutum with yellowish granules only on the middle region. Carapace mostly smooth, with a group of granules on the posterior region. Ocularium located slightly posterior to the ozopores, with a paramedian pair of small tubercles. Mesotergum well delimited, divided into four well-marked areas: I divided medially into left and right halves, with a medial large tubercle on each half and one small lateral tubercle and two anterior granules on each side; II and III entire, with a paramedian pair of large tubercles and two to three pairs on each side; IV divided into two halves, with four tubercles on each side. Posterior border of the DS substraight, with a row of tubercles. Free tergites I with a row of tubercles; II and III with a medial large spine (spine in II larger than in III) and some lateral tubercles, some of them yellowish colored.
Venter (Figs 10B, C, 11B). Stigmatic area with the posterior border convex, unarmed, only with minute granules sparsely distributed. Stigmata large, oval, and subparallel. Coxa I with two rows of large tubercles of different sizes; coxa II longer than coxae I and III, II and III with two rows of low tubercles, the anterior ones more conspicuous; coxa IV strongly backward and widened, conspicuously wider than the anterior coxae, with large tubercles in the posterior and lateral regions. Free sternites each with a row of small tubercles, the lateral larger.
Chelicerae (Fig. 11A). Chelicerae not swollen. Fixed finger with five teeth. Movable finger with ten teeth, one large tooth and distal inner surface dentate. Mesal side of the base of the fixed finger and near the base of the movable finger with setiferous tubercles of different sizes.
Female (CIUQ-020636) (Fig. 10F). Distinguished from male by carapace broader and the coda wider, genital operculum wider, spines on coxae IV present but less conspicuous than those found in males. and distal spine of femur smaller than in males. Distribution. Known only from the type locality. Etymology. fortunata is a Latin word that means happy, lucky, and blessed. It is used in reference to the meaning of the name of the type locality Buenaventura.
Remarks. Among the known genitalia, only Licornus tama Kury, 2012 andHernandarioides plana F.O. Pickard-Cambridge, 1905 have an elongated ventral plate. However, both genera have a subtle distal cleft in the distal border of the VP, which is not the typical deep cleft seen in the subfamily (e.g., Hexabunus Roewer, 1913, Hutamaia Soares & Soares, 1977, Thaumatocranaus Roewer, 1932. Ampycella fortunata sp. nov. shares the presence of an elongated ventral plate, but it is unique in not having a distal cleft. Additionally, a pair of large, apically pointing MS-D is observed in Ampycella fortunata, which is also observed in some of the genera whose genitalia is known (e.g., Hexabunus, Hernandarioides Pickard-Cambridge, 1905).

Species list recorded or described from Colombia
Only those species described, registered, or modified after 2003 are provided with a detailed logonymy here. For the remaining species, refer to the catalog by Kury (2003). When possible, we added information about the WWF ecoregion where the species is found according to available records.
Remarks. Nueva Granada was a historical region of northern South America. The Viceroyalty of New Granada (1717-1723, 1739-1810, 1815-1822) was a political and administrative entity established by Spain in the 18 th century that encompassed a large part of the Andean region of South America, including present-day Colombia, Ecuador, Panama, and Venezuela. It was governed by a viceroy appointed by the Spanish crown. After gaining independence from Spain in 1819, the Republic of New Granada (1831-1858) was established as a sovereign state in the northern Andean region of South America, encompassing present-day mainly Colombia and Panama. The republic underwent various changes in its name and territorial boundaries.

Remarks.
The type locality for this species is Moa River in the state of Acre, in Brazil. It was recorded for Colombia by Villarreal-M. and Pinto-da-Rocha (2006).

Rationale for exclusions
Two species, Cosmetus flavopictus Simon, 1880 and Stygnoplus forcipatus have traditionally been associated with the list of Colombian species, which we do not consider in our list. Cosmetus flavopictus was described for a locality in Darien, extrapolated by Kury (2003) as Colombian territory. Here, we follow the interpretation given by Medrano et al. (2021b) and consider not citing it as part of the Colombian species. Stygnoplus forcipatus Koch, 1845 was originally described without any locality data other than "Columb." (Koch 1845). Subsequently, it was recorded from Venezuela and its presence in the Colombian fauna was called into question (Villarreal-M. and Rodríguez 2004).
Other species of the genus Stygnoplus are known from the Caribbean region, in the Venezuelan Cordillera de La Costa and the Lesser Antilles, with the exception of S. longipalpus (Goodnight & Goodnight, 1942), which occurs in the Guiana Shield. Recently, the only Amazonian species originally described in this genus by Pinto-da-Rocha and Tourinho (2012) were transferred to Yapacana Pinto-da-Rocha, 1997 (Villarreal et al. 2021b). We have decided not to include S. forcipatus in the list at this time due to lack of evidence of its presence in the country.

Diversity
The Colombian opilionofauna of the suborder Laniatores is composed of 173 species, three of which are described in the present work. The 12 families listed in Colombia demonstrate the country's high taxonomic diversity for this group. The most diverse family is Cosmetidae with 67 species (Fig. 13B), followed by Cranaidae with 40 species (Figs 13C, D, 14D). It is worth noting that a significant number of Laniatores species in Colombia are poorly documented, with only vague records for the country or department, and their precise type localities remain unknown. Specifically, there are 16 species of Cosmetidae, five species of Cranaidae, three species of Zalmoxidae, one species of Manaosbiidae, two species of Nomoclastidae, one species Incertae sedis, and one species of Stygnidae with uncertain recorded information. These findings highlight our limited understanding of the diversity and distribution of this group in Colombia. Most records come from a few departments, indicating a sampling bias. Approximately 22% of the species are known from the department of Cundinamarca (38 spp.) near the capital of the country, followed by Tolima and Boyacá with 13 species each. In contrast, all six departments of the Amazon Table 2. Number of Opiliones species by family recorded in WWF terrestrial ecoregions from Colombia. When the locality is unknown or not precise enough, the abbreviation DNA is used, and the record is not assigned to any ecoregion. DNA = does not apply.

WWF Ecoregion
Code ( Table 2). The ecoregion with the highest number of species is the Magdalena Valley montane forests, with a total of 59 species. The ecoregions with the fewest species are Eastern Cordillera real montane forests, Iquitos várzea, Llanos, and Purus várzea, each with one species (Figs 15, 16). Only 21 species are found in more than one ecoregion, with Eucynorta quadripustulata (Fig. 13B) being the most widely recorded species, occurring in five ecoregions. Figure 14. Laniatores species described from Colombia A the pie chart shows authors who described two or more species. For each author, every described species was counted, regardless of whether the authorship of the species is shared with other authors. For example, Usatama infumatus was described by Kury, García & Medrano, 2019, and was counted as a species for each author. Authors with one species are included in the others category B accumulation curve of Laniatores species described from Colombia. Pink: descriptions made between 1833 and 1911; ochre: 1912-1963; green: 1964-2023 C Laniatores species in RUNAP D families of Laniatores species described from Colombia.
The Colombian NSPA lists 1,486 protected zones, of which 60 are NNP and FFS. The remaining areas are designated with some other category of protection (RUNAP 2022). Regarding Opiliones Laniatores, 36 species have been recorded from 29 areas, totaling 47 records, and 11 species have been recorded in seven NNP or FFS. The protection category with the highest number of the recorded species is RDIM, with 16 species, followed by NPFR with ten species, NNP with eight species, FFS with four species, PNR with three species, RFPR with two species, and RNSC with one species.
Similar to other neotropical countries, the taxonomic knowledge of harvestmen in Colombia is primarily based on the contributions of foreign authors. Remarkably Carl F. Roewer described almost half of the species known of the country. Therefore, for a pragmatic approach we divided the history into three periods: the first period prior to Roewer´s contributions, the second period in which most of the descriptions took place, led by Roewer's works but with contributions from other authors, and finally, the third, post-Roewerian or modern period, with contributions from a diverse group of authors, including local authors for the first time. Our goal is not to present a detailed chronology, but rather to provide a general overview of the historical panorama and the individual contribution of the authors to the knowledge of the Colombian fauna. For a detailed chronicle and an in-depth look at the early history of opilionology, we recommend referring to Kury (2010), who presented an excellent compilation and analysis of papers between 1758 and 1804. Figure. 14B shows a timeline

Diversity of Colombian Opiliones
Colombia is recognized as a megadiverse country, renowned for its numerous protected natural areas compared to other neotropical countries. Despite ongoing conservation efforts, there is a need to increase the knowledge on biodiversity in these areas to support their proper management and handling. Unhappily, the information available on Laniatores in these areas is still limited, despite the growing number of arachnologists in the region and the country, and the increasing research focus on local fauna (Fig. 14C).
The central and eastern cordillera are the most populated and accessible areas of Colombia. They are home to the largest universities and biological collections in the country and, consequently, the region with the highest number of descriptions and records of Laniatores (Figs 15, 16). Local researchers have made significant contributions to the study of Laniatores in the last decade, including the review of some genera such as Rhaucus or Eulibitia (García and Kury 2017;Medrano and Kury 2017) and description of some new species (e.g., Kury et al. 2019;García and Ahumada-C. 2022).
In contrast, the peripheral zones, like the Amazon, the Pacific, or the Caribbean, have received less attention from researchers. Nevertheless, recent efforts have been made to strengthen research in these regions. For instance, a school of arachnology at the Universidad del Valle, has emerged to study the Laniatores of Valle del Cauca in the Pacific region. Similarly, in the Caribbean, harvestmen have been the focus of some degree work, and recent collections have yielded specimens that have been added to the country's biological collections, resulting in the descriptions of new species and new records (e.g., Pinzón and Pinto-da-Rocha 2020; Ahumada-C. et al. 2022;Kury et al. 2023).
Therefore, we believe that although the national inventory of Laniatores is far from being complete, further work on the local fauna will increase our knowledge of the group over time. Regarding the Colombian protected areas, the RNSC is the area with the lowest number of records of Laniatores (one species), while RDIM presents the highest number of records (16 species), probably because it is the only protected area in the country conceived as a rational use mode (Molina 2013), with research permits that are easier to access. However, when considering the total number of records for Laniatores across all categories of protected areas, studies are also scarce (Figs 17, 18). The implementation of strategies that facilitate the necessary regulatory procedures to collect specimens in protected natural areas, as well as increased investment in basic science projects, could help to generate a greater understanding of Colombian biodiversity in these areas. A successful case is the study by García and Medrano (2015) in the Reserva Natural Rio Ñambí, the only faunal study on harvestmen conducted in a protected natural area, resulting in the discovery of approximately 28 species new to science, demonstrating the hidden diversity in these areas, and the importance of inventories or taxonomic studies on the protected natural areas.
Continued exploration of ecosystems is imperative to search for Opiliones and achieve reliable taxonomic identifications, which are essential to determine the true distribution of the species. Such data is important in establishing their threat category, according whit the criteria from the International Union for Conservation of Nature (IUCN). This should be a goal for arachnology, given the high endemicity exhibited by these organisms and the significant anthropogenic pressures to which they are exposed.

Taxonomic accounts
Phalangodus is an Andean cranaid genus whose species could be considered examples of short-range endemism, as defined by Harvey (2002). The distribution of Phalangodus andresi sp. nov. is within the area of generic distribution and represents the first case of sympatry in the genus, as it was collected in close proximity to P. anacosmetus, a widely distributed species found in the highlands of the department of Cundinamarca (Villarreal and García 2016).
Phalangodus andresi sp. nov. possesses all generic diagnostic characters presented by Villarreal and García (2016), and can be diagnosed by its smaller size, by having dorsal scutum tuberculated (shared only with P. palpiconus), being distinguished by the type of ornamentation of the femur IV of the males (refer to the diagnosis for further details). Unfortunately, of the type locality for P. palpiconus was erroneously attributed to Chile, which is unlikely due to the distribution pattern of the genus and even the family, as discussed by Hara et al. (2014), making impossible to establish any biogeographic relationship between both species.
Moreover, the family Ampycidae is known to have a little-explored diversity, primarily in Ecuador and Colombia. However, the taxonomic assignment of new species is a challenge for taxonomists due to the paucity of old descriptions and poor generic diagnosis. In addition, many species exhibit similar external morphology, and for more species, genitalia remain unknown, with just some exceptions (e.g., Villarreal and Kury 2012;Kury and Quintero 2014;Tourinho and Mendes 2014;Hara et al. 2017). Despite these challenges, recent studies on some species have shed light on the external morphology and genitalia of the subfamily. Further research is necessary to address the taxonomic uncertainties and increase our knowledge of the diversity of Ampycidae.
as the constant feedback. To Marco R. Hara for the feedback regarding the identity of Phalangodus andresi sp. nov. in relation to P. palpiconus. We would like to express our gratitude to the Vice-Rector's Office for Research of the Universidad del Quindío for their invaluable assistance in reviewing the English writing of our manuscript. To Andrés F. García for the Scanning Electron Microscopy (SEM) photographs used in Fig. 7. The SEM photographs were taken at the Microscopy Platform Rudolf Barth/IOC-Fiocruz. OV thanks Pío Colmenares (AMNH) for coordinating the donation of optical equipment and supplies to MIZA through the Cooperation Agreement between MIZA-AMNH. The following photographers kindly provided the use of their photographs of live harvestmen for Fig. 13: Andrés García, Arthur Anker, Hugo Vides, Julio González /Entomopixel, and Sarah Crews. To Miguel Medrano, Ligia Benavides, and one anonymous referee, thank you for valuable suggestions and comments that greatly improved the present work; to Nathalie Yonow for her editorial revision work. To Rodrigo Baos for his kindness in San Pedro, Buenaventura, Valle del Cauca.

Conflict of interest
The authors have declared that no competing interests exist.

Ethical statement
No ethical statement was reported.

Funding
This study has been partly supported by the Vice-Rector's Office for Research of the

Data availability
All of the data that support the findings of this study are available in the main text or Supplementary Information.