﻿Passalidae (Coleoptera, Scarabaeoidea) from the Caribbean coast of Colombia: synopsis, key, and new species description

﻿Abstract Bess beetles (Passalidae) are a subsocial family of Coleoptera with approximately 1000 known species of saproxylophagous diet and pantropical distribution, with few extratropical species. Because of their high levels of endemism (especially in mountains), feeding habits, and complex subsociability; Passalidae is considered an excellent biological subject for taxonomic, biogeographical, and evolutionary studies. Colombia is the richest country with more than 118 recorded species of Passalidae, most of the species being related to humid and mountain areas. Colombia’s Caribbean region constitutes the northern portion of the country, extending for more than 130,000 km2 and includes four of the eight biogeographical provinces of Colombia. Since the 2000s this region has been the subject of systematic surveys for Passalidae; as a result, 18 passalid species have been recorded to date. After new explorations and review of entomological collections, the knowledge of the passalid fauna for the region is updated, recording 28 species (8 new records, 2 new species) for which are provided species diagnoses, photographs, and a taxonomic key. The dry plain, characteristic of the lowlands, is dominated by widely distributed species such as Passaluspunctiger and Passalusinterstitialis, while the mountainous systems provide species of more restricted distributions, some of them endemic to the Colombian Caribbean.


Introduction
Beetles of the family Passalidae are saproxylophagous diet, playing an important role in nutrient recycling . Adults establish multigenerational colonies inside rotting logs, on which they also feed. Although passalids are not eusocial, they exhibit intricate social relationships that include sound communication, in which up to 15 different signals have been detected that vary according to the context (e.g., stress, courtship). Approximately 1000 species are known worldwide, distributed mainly in tropical-humid zones (Schuster 1978;Boucher 2006), most of which exhibit small ranges (i.e., high degree of endemism). Recent studies suggest that the family Passalidae originated in Pangaea, more than 200 million years ago (Beza-Beza et al. 2021); however, the family has a markedly pantropical distribution, with few extratropical species.
The previously mentioned characteristics make Passalidae an excellent biological subject for ecological, biogeographical, taxonomic, and evolutionary studies. Consequently, in Colombia, Passalidae is one of the faunistically bestknown families of Coleoptera, with an estimated 118 species distributed in the country, making it the country with the highest number of recorded Passalidae species in the world (Jiménez-Ferbans et al. 2018a). Maybe due to affinity with humid ecosystems and historical process, the biogeographic regions with the highest levels of species richness are the Chocó and Amazon provinces, with 41 and 24 species respectively (Amat-García and Reyes -Castillo 2007;Jiménez-Ferbans et al. 2018b). However, many regions have not been studied comprehensively and further exploration is likely to increase the known number of species, especially in mountainous areas, such as the Andes and Sierra Nevada de Santa Marta (SNSM), given the high levels of endemism typified by montane passalid species (Beza-Beza et al. 2021).
The Colombian Caribbean is constituted, in geopolitical terms, by seven departments or states (La Guajira, Cesar, Magdalena, Atlántico, Bolívar, Sucre and Córdoba), covering more than 130,000 km 2 . It includes four of the eight main biogeographic provinces of Colombia ( Fig. 1) and presents a high geomorphological heterogeneity that includes: savanna, tropical dry forest, and high Andean forest  Hernández-Camacho et al. (1992), and the localities with records from Passalidae. ecosystems, among others. Climatologically, there is a gradient of conditions in the Caribbean, starting in the northern zone (La Guajira and part of Magdalena) with arid, semi-arid, and semi-dry climates that decrease towards the south, until diverging into semi-humid, humid, and very humid climates in the departments of Bolivar, Cordoba, and Cesar (Carvajal-Cogollo and Rangel-Ch 2012).
The Caribbean region of Colombia is the most comprehensively know for passalids, based in amount of works published in the last two decades (e.g., Jiménez-Ferbans and Amat-García 2009;Jiménez-Ferbans et al. 2012, 2022Taboada-Verona and Murillo-Ramos 2020). Jiménez-Ferbans andAmat-García (2009, 2010), and  represented the first works that systematically studied the passalids of the region, registering 18 species and proposing a key and synopsis for their identification. Then, Jiménez-Ferbans et al. (2012, 2022 described new species for the Sierra Nevada de Santa Marta, increasing the number of species to 21. Additionally, recent explorations have allowed the discovery of species not registered for the region and increasing the number of sites with records of passalids. This work updates the knowledge of the family Passalidae in the Colombian Caribbean region, providing diagnoses, a taxonomic key, and the description of new species. A refined database of the revised material is also provided, with the aim of promoting the study of the family, facilitating the identification of species and the recognition of areas with exploration gaps.

Materials and methods
We studied and digitized 5400 specimens deposited in Colección Entomológica Universidad del Magdalena (CBUMAG-ENT), Colombia (GBIF dataset: https:// www.gbif.org/dataset/54F7AF87-E0A3-4816-AE9F-ED77C45F2455). We also included 380 records from Museo de Zoología de la Universidad de Sucre, Colombia (GBIF dataset https://www.gbif.org/dataset/6F1BED71-743E-407B-8862-E7E2E1CED896), and conducted a literature review. The total records, including those from literature, are included in a Darwin core formatted file (Suppl. material 1). GBIF data sets went through a quality process to guarantee its usability in research and decision-making processes, this was supported by the GBIF-BID "Data mobilization for key entomological groups across the Caribbean Region of Colombia". For this, the datasets were verified in OpenRefine 3.7.2 (http://www.openrefine.org) and validated in GBIF Data Validator and the geographic validation service in QGIS provided by SiB Colombia (https:// biodiversidad.co/formacion/laboratorios/QGIS). Taxonomic validation was performed according to the GBIF taxonomic tree.
The classification used is that of Beza-Beza et al. (2020) and Jiménez-Ferbans et al. (2023). While the terminologies of morphology are those proposed by Boucher (2006) for the head, except that we retain the term "frontal fossae" of Reyes-Castillo (1970), and the terminology of Reyes-Castillo (1970) for the rest of the body. For the taxonomic determinations, we employed the keys in Kuwert (1891), Marshall (2000), Boucher (2006), Gillogly (2005), Jiménez-Ferbans et al. (2018b); original descriptions and comparison with determined specimens deposited in collections. Total body length (based on the material examined) was measured from the apex of the mandible to the tip of the elytra with a digital caliper, when possible, we measured at least 10 specimens for each species.
For each species we examined, we give a diagnosis based on external morphological characters and photographs of relevant body areas. The photographs were taken using a Canon EOS Rebel SL3 camera. Then, they were stacked in layers by the software Helicon Focus v. 8.0.1 to generate a single image of combined focus. This image was edited for light and contrast correction in Adobe Photoshop, and the final combined figures were made using the same software, following the guide proposed by Bevilaqua (2020). Suppl. material 2 shows the label information for the photographed specimens.

Results
The passalid fauna of the Colombian Caribbean Coast comprises 28 species of the genera Passalus (10 spp), Rhodocanthopus (2 spp), Spasalus (2 spp), Paxilus (1 spp), Veturius (4 spp), Popilius (3 spp), Heliscus (1 spp), Odontotaenius (1 spp), and Verres (2 spp) ( Table 1). The biogeographical province of Chocó-Magdalena is the richest one, despite being the one with fewer locations with records, followed by the SNSM, which has the largest number of sampled localities ( Table 2). As might be expected given the passalids preference for moist environments, the dry plain, characteristic of the lowlands of northern Colombia, has the least number of species, all of them being widely distributed.

Tribe Passalini
The species of this tribe are recognized by having the clypeus hidden under the frons, with anterior angles located under the external tubercles (Reyes -Castillo 1970) and frontoclypeus absent (Boucher 2006). Fig. 2 Diagnosis. 39.6-46.5 mm total length. Body robust. Anterior border of the frons with two prominent secondary mediofrontal tubercles. Mediofrontal tubercles large, located on base of each laterofrontal tubercle. Central tubercle with apex free. Lateroposterior tubercles distinct. Eyes large. Antennal club tri-lamellate, with lamellae long. Lacinia with apex bidentate. Mediobasal area of mentum protruding and glabrous, sometimes with scarce setae on the posterior border. Marginal pronotal groove occupying 2/3 of the pronotum anterior border. Prosternellum rhomboidal, truncate. Mesosternum pubescent, with inconspicuous and elongated scars. Metasternum pubescent anterolaterally and in lateral groove; disc smooth, delimited posteriorly to middle by punctations. Humeri pubescent, epipleura pubescent in basal 2/3. Last abdominal sternite with complete marginal groove. Anterior ventral border of the profemur with well-developed groove. Meso-and metatibia with one or two small spines.

Passalus (Passalus) coniferus Eschscholtz, 1829
Comments. A South American species, with distribution in the Antilles (Reyes- Castillo 1973). It is a species of highly variable size, Reyes-Castillo and Table 1. Species of Passalidae from the Colombian Caribbean Coast. The distribution indicates the department on the Caribbean Coast, followed by the general distribution of the species. Altitude data were taken from the material examined and ranges cited by other authors are given in parentheses.  Amat- García (2003) notes that the specimens from the Sierra Nevada de Santa Marta are larger than those from the rest of the country, with the apex of the horn free and very long.   -Castillo 1973). This species sometimes is confused with P. punctiger, but is distinguished by its larger size and incomplete marginal groove in last abdominal sternite (complete in P. punctiger). Fig. 4 Diagnosis. 23.6-29.9 mm total length. Body flattened. Anterior border of the frons with two prominent secondary mediofrontal tubercles. Mediofrontal tubercles large, located on base of each laterofrontal tubercle. Central tubercle with apex not free. Lateroposterior tubercles distinct. Eyes large. Antennal club tetra-lamellate, fourth lamella reduced. Lacinia with apex bidentate. Mediobasal area of mentum glabrous and slightly protruding. Marginal pronotal groove occupying 2/3 of the pronotum anterior border. Prosternellum rhomboidal, truncate. Mesosternum glabrous, with distinct elongated scars. Metasternum pubescent anterolaterally and in lateral groove; disc smooth and delimited posteriorly to middle by punctations. Humeri pubescent, epipleura pubescent in basal third. Last abdominal sternite with complete marginal groove. Anterior ventral border of the profemur with distinct groove. Mesotibia lacking spine or with single small spine.

Passalus (Passalus) punctiger Lepeletier & Serville, 1825
Comments. Distributed from Mexico to Argentina. This the most common species for the lowlands of the Colombian Caribbean. Fig. 6 Diagnosis. 28.7-34.2 mm total length. Body slightly flattened. Anterior border of frons with two prominent secondary mediofrontal tubercles, almost completely fused. Mediofrontal and laterofrontal fused, large. Central tubercle with apex very free, reaching or surpassing anterior frons border. Lateroposterior tubercles small, distinct. Eyes large. Antennal club tri-lamellate, with lamellae long. Lacinia with apex bidentate. Mediobasal area of mentum protruding and pubescent. Marginal pronotal groove occupying 2/3 of the pronotum anterior border. Prosternellum rhomboidal, truncate. Mesosternum glabrous, with distinct and elongate scars. Metasternum pubescent anterolaterally, lateral groove glabrous; disc smooth and delimited by punctations excluding the anterior part. Humeri pubescent, epipleura with some setae basally. Last abdominal sternite with complete marginal groove. Anterior ventral border of the profemur with conspicuous groove. Meso-and metatibia with small spines or unarmed.

Passalus (Passalus) serankuai Jiménez-Ferbans, Reyes-Castillo & Amat-García, 2014
Comments. Described from SNSM, this species seems to be endemic to this biogeographical province. Diagnosis. Hemibrachypterous. Frons wide, anterior frontal edge straight, without middle indentation or secondary mediofrontal tubercles. Central tubercle wide at the base, without a sulcus in the posterior part, apex free, almost reaching the anterior frons border. Medial basal mentum impunctate and glabrous. Prosternellum rhomboidal and opaque in the area between procoxae, without longitudinal groove. Anterolateral part of metasternum and lateral fossa pubescent, pubescence reaching the posterior region of the lateral fossa. Metasternal disc without punctures, delimited by numerous punctures posteriorly. Humeri pubescent and epipleura glabrous.
Head (Figs 7A,D,8A): labrum with anterior border slightly concave, evenly covered by setae. Clypeus hidden under the frons, anterior angles developed under the mediofrontal + laterofrontal tubercles and slightly smaller than these. Frons wide, anterior frontal edge straight, without middle indentation and secondary mediofrontal tubercles. Mediofrontal + laterofrontal tubercles projected forward, larger than internal tubercles. Internal tubercles small, joined to mediofrontal + laterofrontal tubercles by an inconspicuous ridge, placed at mid distance between the mediofrontal tubercles and the central tubercle base. Posterofrontal ridges "V" shaped. Area between the frontal ridges heavily punctuated without a median sulcus and cephalic mamelon (sensu Jiménez-Ferbans and Reyes-Castillo 2014). Mesofrontal structure of the "marginatus" type (Reyes-Castillo 1970), with central tubercle wide at the base, without a sulcus in the posterior part, apex free, almost reaching anterior frons border. Lateroposterior tubercles large, parallels to central tubercle. Lateropostfrontal areas glabrous, shiny, and impunctate. Eyes reduced, not extending past ocular canthi (dorsal view) and with canthus covering almost 1/2 of the eye in lateral view. Canthus glabrous. Postorbital pits shallow. Postfrontal groove semicircular and complete. Hypostomal process slightly separated from the mentum, glabrous and reaching the superior part of the middle zone of the mentum. Medial basal mentum protruding ventrally, impunctate and glabrous. Mentum with rounded lateral fossae, shallow and pubescent laterally. Antennal club tri-lamellate. Dorsal tooth straight on dorsal view and slightly sinuous on lateral view. Internal tooth of the left mandible bidentate, simple on the right mandible. Mandibular fossae short, not reaching the base of the mobile tooth. Maxilla with lacinia bidentate at the apex. Ligula tridentate, with middle tooth longer than the lateral teeth. Middle palpomere of the labial palp 1.3× wider and with almost the same length as the distal palpomere.
Thorax (Figs 7,8): Pronotum rounded, same width as elytra, with punctuations extending outside the lateral fossae and marginal groove. Marginal groove wide, occupying ¾ of the anterior margin of the pronotum. Longitudinal sulcus conspicuous. Lateral fossae distinct. Pre-epimeron shiny and heavily pubescent. Prosternellum rhomboidal and opaque in the area between procoxae. Mesosternum with erased mesosternal scars, indicated by an opaque area, impunctate and glabrous. Posterior corner of the mesepisternum and mesepimere glabrous and shiny. Anterolateral part of the metasternum and lateral fossa pubescent, pubescence reaching the posterior region of the lateral fossa. Metasternal disc without punctures, delimited by numerous punctures posteriorly. Posterior metasternal lateral fossa of the same width as epipleura.
Abdomen (   Head (Figs 9A, D, 10A): labrum with anterior border concave, covered by setae less dense in the middle region. Clypeus hidden under the frons, anterior angles under the mediofrontal + laterofrontal tubercles and smaller than these. Frons wide, anterior frontal edge straight, with middle indentation and secondary mediofrontal tubercles rudimentary. Mediofrontal + laterofrontal tubercles projected forward, larger than internal tubercles. Internal tubercles small, rudimentary, joined to mediofrontal + laterofrontal tubercles by an inconspicuous ridge, placed at mid distance between the mediofrontal tubercles and the central tubercle base. Posterofrontal ridges V-shaped, inconspicuous. Area between the frontal ridges 3 punctures with a median sulcus and without cephalic mamelon (sensu Jiménez-Ferbans and Reyes-Castillo 2014). Mesofrontal structure of the "marginatus" type (Reyes-Castillo 1970), with central tubercle wide at the base, with a sulcus in the posterior part, apex free, reaching or surpassing anterior frons border. Lateroposterior tubercles large, parallel to central tubercle. Lateropostfrontal areas glabrous, shiny, and impunctate. Eyes reduced, not extending past ocular canthi (dorsal view) and with canthus covering almost 1/2 of the eye in lateral view. Canthus glabrous. Postorbital pits shallow. Postfrontal groove semicircular and complete. Hypostomal process slightly separated  from mentum, glabrous and reaching the superior part of the middle zone of the mentum. Medial basal mentum protruding ventrally, with punctures and setae on posterior border. Mentum with rounded lateral fossae, shallow and pubescent laterally. Antennal club tri-lamellate. Dorsal tooth straight on dorsal view and slightly sinuous on lateral view. Internal tooth bidentate on left and right mandible. Mandibular fossa short, not reaching the base of the mobile tooth. Maxilla with lacinia bidentate at the apex. Ligula tridentate, with middle tooth longer than the lateral teeth. Middle palpomere of the labial palp 1.3× wider and with almost the same length as the distal palpomere.
Mesotibiae with a small spine on the outer margin. Metatibiae unarmed.
Aedeagus (Fig. 9C, D): Basal piece (ventral view) fully fused with parameres and with deep V-shape cleft. Median lobe globose, little sclerotized on the ventral surface, length is 1.1× the length of the basal piece and parameres, measured at the median ventral line. Lateral projections of the parameres short and apex rounded on lateral view.
Variation: In some paratypes, the area between the frontal ridges has 0-4 punctures, and the metasternal disc with 0-10 punctures.
Etymology. Named after Mr. Víctor Flórez Carrillo, for his dedication to entomological exploration in the Serranía del Perijá. Fig. 11 Diagnosis. 31.6-34.2 mm total length. Body robust. Anterior border of the frons with small middle indentation, without secondary mediofrontal tubercles. Mediofrontal and laterofrontal fused, midsize. Central tubercle with apex not free. Lateroposterior tubercles slightly distinct and rounded. Eyes reduced, with canthus covering 1/2 of the eye in lateral view. Antennal club tri-lamellate, with lamellae long. Lacinia with apex bidentate. Mediobasal area of mentum protruding and heavily punctate and pubescent. Marginal groove wide, occupying 3/4 of the anterior margin of the pronotum. Prosternellum rhomboidal, truncate. Mesosternum without mesosternal scars, indicated only by an opaque area, impunctate and glabrous. Metasternum pubescent anterolaterally and in lateral groove; disc smooth and delimited by numerous punctations medially and posteriorly. Humeri and epipleura glabrous. Last abdominal sternite with complete marginal groove. Anterior ventral border of the profemur with thin groove. Mesotibiae with small spines on the outer margin. Metatibiae unarmed.

Passalus (Pertinax) gaboi Jiménez-Ferbans & Reyes-Castillo, 2022
Comments. Passalus gaboi had been the last species described from the Colombian Caribbean. It seems to be endemic to the Sierra Nevada de Santa Marta. Fig. 12 Diagnosis. 22.9-25.5 mm total length. Body robust. Anterior border of the frons straight, with small and irregular tubercles, without secondary mediofrontal tubercles. Mediofrontal and laterofrontal fused, small, projected forward. Central tubercle small, with apex not free. Lateroposterior tubercles small, slightly distinct. Eyes large. Antennal club tri-lamellate, with lamellae short. Lacinia with apex bidentate. Mediobasal area of mentum protruding and glabrous. Marginal groove widened, occupying 2/3 of the anterior margin of the pronotum. Prosternellum rhomboidal, truncate. Mesosternum glabrous; mesosternal scars absent. Metasternum glabrous anterolaterally and in lateral groove; disc smooth and delimited by punctations posteriorly. Humeri and epipleura glabrous. Anterior ventral border of the profemur with thin groove. Meso-and metatibiae with small spines or unarmed.

Passalus (Pertinax) punctatostriatus Percheron, 1835
Comments. Distributed from Mexico to central and northern Colombia. Amat-García et al. (2004) cited this species from the Amazon region, but without citing specimens. We doubt its presence in southern Colombia. P. punctatostriatus has been collected in shady coffee plantations in Serranía del Perijá and in Sierra Nevada de Santa Marta (Jiménez-Ferbans and Amat-García 2009).   Fig. 14 Diagnosis. 20.1-23.2 mm total length. Body subcylindrical. Anterior border of the frons with strong middle indentation, suggesting secondary mediofrontal tubercles. Mediofrontal and laterofrontal fused, large. Central tubercle with apex not free. Lateroposterior tubercles large, conspicuous, and conical. Eyes large. Antennal club tri-lamellate, with lamellae long. Lacinia with apex bidentate. Mediobasal area of mentum protruding and glabrous. Marginal groove narrow, occupying 2/3 of the anterior margin of the pronotum. Prosternellum rhomboidal, acute. Mesosternum glabrous; mesosternal scars distinct and elongated. Metasternum glabrous anterolaterally and in lateral groove; disc smooth and delimited by punctations posteriorly to laterally. Humeri pubescent and epipleura pubescent in basal third. Anterior ventral border of the profemur with conspicuous groove. Meso-and metatibiae with small spines or unarmed.

Passalus (Pertinax) rugosus Gravely, 1918
Comments. Jiménez-Ferbans et al. (2018b) considered this species as endemic to the western Andes of Colombia. However, the present records extend its distribution to the eastern Andes. Fig. 15 Diagnosis. 19.5-22.4 mm total length. Body subcylindrical. Anterior border of the frons with strong middle indentation, without secondary mediofrontal tubercles. Mediofrontal and laterofrontal fused, large. Central tubercle with apex not free. Lateroposterior tubercles small, slightly distinct. Eyes large. Antennal club tri-lamellate, with lamellae long. Lacinia with apex bidentate. Mediobasal area of mentum protruding and glabrous. Marginal groove widened, occupying 2/3 of the anterior margin of the pronotum. Prosternellum rhomboidal, truncate. Mesosternum glabrous, except for a few short setae on the anterior margin; mesosternal scars inconspicuous and elongated. Metasternum glabrous anterolaterally and in lateral groove; disc smooth and delimited by punctations posteriorly to laterally. Humeri with long setae at its base, epipleura glabrous. Anterior ventral border of the profemur with conspicuous groove. Meso-and metatibiae with small spines or unarmed.

Passalus (Pertinax) unimagdalenae Jiménez-Ferbans, Reyes-Castillo & Amat-García, 2012
Comments. This is the most common species above 1500 m a.s.l. in Sierra Nevada de Santa Marta. P. unimagdalenae is similar to P. punctatostriatus, from which it differs by having internal tubercles large and humeri with long setae at the very base. Fig. 16 Diagnosis. 17.5-18.8 mm total length. Anterior border of the frons straight, sometimes with a small central notch, without secondary mediofrontal tubercles. Mediofrontal tubercles large. Internal tubercules smaller than mediofrontal tubercles,   located between mediofrontal tubercles and central tubercle. Central tubercle with apex not free. Lateroposterior tubercles distinct. Anterior cephalic angles reach the border of the frons. Frontal fossae glabrous. Eyes large. Antennal club with five lamellae, with the basal lamella no more than half as long as the apical four. Lacinia with apex unidentate. Mediobasal area of mentum flat and glabrous. Prosternellum pentagonal. Mesosternum glabrous, with conspicuous oval scars. Metasternum pubescent anterolaterally and glabrous in lateral groove; disc smooth, completely delimited by punctures. Humeri pubescent and epipleura pubescent in basal 1/4. Anterior ventral border of the profemur without groove or inconspicuous.

Paxillus leachi MacLeay, 1819
Comments. This species is distributed throughout the American continent, from Mexico to Argentina. Fig. 17 Diagnosis. 16.3-19.8 mm total length. Body subcylindrical. Anterior border of the frons with strong middle indentation, without secondary mediofrontal tubercles. Mediofrontal and laterofrontal fused, large. Central tubercle with apex not free. Lateroposterior tubercles conspicuous and conical. Eyes reduced. Antennal club tri-lamellate, lamellae short. Lacinia with apex bidentate. Mediobasal area of mentum protruding and glabrous. Marginal groove widened, occupying 2/3 of the anterior margin of the pronotum. Prosternellum rhomboidal, acute. Mesosternum glabrous; mesosternal scars oval and distinct. Metasternum glabrous anterolaterally and in lateral groove; disc smooth and delimited by punctations posteriorly to laterally. Humeri and epipleura glabrous. Anterior ventral border of the profemur with conspicuous groove. Mesoand metatibiae with strong spines.

Rhodocanthopus maillei (Percheron, 1841)
Comments. Species very similar to R. maillei, from which it differs by its larger size, reduce punctation on the pronotum, which do not reach the mid zone and the metasternal disc not abundantly punctate. Fig. 18 Diagnosis. 14.0-16.5 mm total length. Body flattened. Anterior border of frons with strong middle indentation, on the sides of which are two insinuated tubercles. Mediofrontal and laterofrontal fused, large. Central tubercle with apex not free. Lateroposterior tubercles distinct and transverse. Eyes reduced. Antennal club tri-lamellate, lamellae short. Lacinia with apex bidentate. Mediobasal area of mentum protruding and glabrous. Marginal groove widened, occupying 1/2 of the anterior margin of the pronotum. Prosternellum rhomboidal, truncate. Mesosternum glabrous; mesosternal scars elongated and conspicuous. Metasternum glabrous anterolaterally and in lateral groove; disc smooth and delimited by punctations posteriorly to laterally. Humeri glabrous; epipleura with scattered setae in basal area. Anterior ventral border of the profemur with conspicuous groove. Meso-and metatibiae with strong spines.   (Jiménez-Ferbans et al. 2016), this species seems to be endemic to the Chocó-Magdalena province. Fig. 19 Diagnosis. 17.6-18.2 mm total length. Body robust. Mediofrontal and laterofrontal fused, large. Central tubercle with apex not free. Eyes large. Antennal with five lamellae. Lacinia with apex unidentate. Mediobasal area of mentum protruding, with setae along almost its entire extension. Marginal groove widened, occupying 2/3 of the anterior margin of the pronotum. Anterior border of mentum almost straight. Prosternellum rhomboidal. Anterior ventral border of the profemur with conspicuous groove.

Spasalus crenatus (MacLeay, 1819)
Comments. Distributed widely in South America, this species has been recorded also in Lesser Antilles and Puerto Rico (Jiménez-Ferbans et al. 2015).

Spasalus paulinae Amat-García & Fonseca, 1998
Diagnosis. 15.75-16.30 mm total length. Body robust. Mediofrontal and laterofrontal fused, large. Central tubercle with apex not free. Eyes large. Antennal with five lamellae. Lacinia with apex unidentate. Mediobasal area of mentum protruding, without setae in the posterior border. Marginal groove widened, occupying 2/3 of the anterior margin of the pronotum. Anterior border of mentum almost straight. Prosternellum rhomboidal. Anterior ventral border of the profemur with conspicuous groove.
Comments. Known only from material type from Sierra Nevada de Santa Marta. The differences between S. crenatus and S. paulinae are subtle, they may be synonymous.

Tribe Proculini
The species of this tribe are recognized by having a frontoclypeus, below which the clypeus is hidden (Boucher 2006). Fig. 20 Diagnosis. 28.7-32.5 mm total length. Anterior border of the labrum straight or slightly concave. Frontoclypeus straight, not swollen in the middle. Frontal-clypeal suture present and strong. Internal tubercles conspicuous, joined to central tubercle by Y-shaped ridges. Central tubercle small, with apex not free; lateroposterior tubercles distinct and transverse, with a superior groove extending over the total length of the tubercles. Frontal fossae pubescent posteriorly. Postfrontal groove complete, slightly erased in the middle. Mediobasal area of the mentum glabrous. Marginal groove over anterior border  of the pronotum not expanded. Mesosternum glabrous. Metasternum pubescent anterolaterally and in lateral groove; disc not delimited by punctations. Mesotibiae and metatibiae with a small spine or unarmed. Humeri and epipleura glabrous; anterior vertical face of elytra pubescent. Fig. 21 Diagnosis. 28.8 mm total length. Anterior border of the labrum straight or slightly concave. Frontoclypeus swollen in the middle. Frontal-clypeal suture present and strong. Internal tubercles small, not joined to the central tubercle. Central tubercle large, with apex free; lateroposterior tubercles absent. Frontal fossae glabrous. Postfrontal groove complete. Mediobasal area of mentum glabrous. Marginal groove over anterior border of pronotum not expanded. Mesosternum glabrous. Metasternum with scarce pubescence anterolaterally and in lateral groove; disc delimited by punctations posteriorly. Mesotibiae and metatibiae with a small spine. Humeri and epipleura glabrous; anterior vertical face of elytra pubescent. Fig. 22 Diagnosis. 23.1-25.0 mm total length. Anterior border of the labrum slightly concave. Frontoclypeus straight or slightly concave. Frontal-clypeal suture present and strong. Internal tubercles conspicuous, joined to central tubercle by Y-shaped ridges. Central tubercle small, with apex not free and posterior to level of lateroposterior tubercles which are conspicuous and transverse. Frontal fossae pubescent. Postfrontal groove complete. Mediobasal area of the mentum glabrous. Marginal groove over anterior border of the pronotum slightly expanded. Mesosternum glabrous. Metasternum with scarce pubescence anterolaterally and in lateral groove; disc not delimited by punctations. Meso-and metatibiae with small spines. Humeri with scarce setae; epipleura glabrous; anterior vertical face of elytra pubescent.

Popilius erotylus Reyes-Castillo & Castillo, 1992
Comments. Popilius erotylus was described from Panama. The specimens studied here show some setal variation in comparison with the type material; they could be a new species very closely related to P. erotylus. Fig. 23 Diagnosis. 21.4-23.5 mm total length. Anterior border of the labrum slightly concave. Frontoclypeus straight or slightly expanded forward. Frontal-clypeal suture present and strong. Internal tubercles small, joined to central tubercle by Y-shaped ridges. Central tubercle small, with apex not free and almost even with lateroposterior tubercles which are distinct and transverse. Frontal fossae pubescent. Postfrontal groove complete. Mediobasal area of the mentum glabrous. Marginal groove over anterior border of the pronotum not   expanded. Mesosternum glabrous. Metasternum pubescent anterolaterally and in lateral groove, pubescence extending beyond lateral groove; disc not delimited by punctations. Meso-and metatibiae with a small spine. Humeri and epipleura glabrous or with scarce setae basally; anterior vertical face of elytra pubescent.

Popilius gibbosus (Burmeister, 1847)
Comments. Common species from the north of the Eastern Cordillera (Serranía del Perijá and Sierra de Venezuela) (Gillogly 2005). It is found in the intermediate and high areas of the Andes, where it is endemic. Fig. 24 Diagnosis. 20.6-24.4 mm total length. Anterior border of the labrum slightly concave. Frontoclypeus straight or slightly concave. Frontal-clypeal suture present and strong. Internal tubercles conspicuous, joined to central tubercle by Y-shaped ridges. Central tubercle small, with apex not free and even with lateroposterior tubercles which are conspicuous and transverse. Frontal fossae glabrous or rarely pubescent. Postfrontal groove complete. Mediobasal area of the mentum glabrous. Marginal groove over anterior border of the pronotum slightly expanded. Mesosternum glabrous. Metasternum with scarce pubescence anterolaterally and in lateral groove; disc not delimited by punctations. Meso-and metatibiae with small spines. Humeri with scarce setae; epipleura glabrous; anterior vertical face of elytra pubescent.

Popilius marginatus (Percheron, 1835)
Comments. P. marginatus has been recorded from Argentina, Bolivia, Brazil, Colombia, French Guiana, Peru and Suriname (Gillogly 2005;Jiménez-Ferbans et al. 2013). Some specimens from SNSM have some differences with the characteristics described by Gillogly (2005) and could be a new species. Fig. 25 Diagnosis. 32.9 mm total length. Anterior border of the labrum deeply concave, with an excavation in labrum behind concavity of margin. Frontoclypeus slightly curved, with a central notch. Frontal-clypeal suture absent. Internal tubercles large, with free apex projecting forward, not joined to central tubercle. Central tubercle short, with apex free projected forward; lateroposterior tubercles conspicuous and transverse. Frontal fossae glabrous. Postfrontal groove complete. Mediobasal area of the mentum glabrous. Marginal groove over anterior border of the pronotum expanded. Mesosternum glabrous. Metasternum pubescent anterolaterally and in lateral groove, pubescence extending beyond lateral groove; disc delimited by punctations posteriorly. Meso-and metatibiae unarmed. Humeri with scarce setae; epipleura glabrous; anterior vertical face of elytra glabrous.

Verres corticicola Kaup, 1871
Comments. Marshall (2000) cited this species from Mexico to Costa Rica, indicating that it is common to collect. In the Colombian Caribbean it seems to be relegated to the humid forests of the biogeographical province of Chocó-Magdalena.   Fig. 26 Diagnosis. 34.4-38.5 mm total length. Anterior border of the labrum deeply concave, with an excavation in labrum behind concavity of margin. Frontoclypeus slightly curved, with a central notch. Frontal-clypeal suture absent. Internal tubercles small, blunt, not joined to central tubercle. Central tubercle with apex slightly free, oblique; lateroposterior tubercles distinct and transverse. Frontal fossae glabrous. Postfrontal groove complete. Mediobasal area of the mentum glabrous. Marginal groove over anterior border of the pronotum not expanded. Mesosternum glabrous. Metasternum pubescent anterolaterally and in lateral groove, pubescence extending beyond lateral groove; disc delimited by punctations posteriorly. Meso-and metatibiae unarmed. Humeri with scarce setae; epipleura glabrous; anterior vertical face of elytra glabrous.

Verres hageni Kaup, 1871
Comments. Jiménez-Ferbans and Amat-García (2009) cited V. hageni from the Caribbean coast of Colombia, but without citing specimens. Consequently, specimens examined in this work are the first from the region. Fig. 27 Diagnosis. 28.7-33.6 mm total length. Anterior border of the labrum slightly concave. Frontoclypeus straight. Frontal-clypeal suture absent. Internal tubercles small, joined to central tubercle by V-shaped ridges. Central tubercle short, with apex not free; lateroposterior tubercles large, conspicuous, and transverse. Frontal fossae pubescent. Postfrontal groove incomplete, interrupted behind central tubercle. Mediobasal area of the mentum pubescent. Marginal groove over anterior border of the pronotum not expanded. Mesosternum with two rows of setae running longitudinally parallel. Metasternum pubescent anterolaterally and in lateral groove; disc not delimited by punctations. Meso-and metatibiae unarmed. Humeri and epipleura glabrous; anterior vertical face of elytra pubescent. Fig. 28 Diagnosis. 40.6-47.9 mm total length. Anterior border of the labrum concave. Frontoclypeus straight. Frontal-clypeal suture absent. Internal tubercles prominent, joined to central tubercle by V-shaped ridges. Central tubercle large, with apex not free; lateroposterior tubercles absent. Frontal fossae glabrous. Postfrontal groove complete. Mediobasal area of the mentum glabrous. Marginal groove over anterior border of the pronotum not expanded. Mesosternum with scarce setae. Metasternum pubescent anterolaterally and in lateral groove; disc not delimited by punctations. Meso-and metatibiae unarmed. Humeri with scarce setae basally; epipleura glabrous; anterior vertical face of elytra sparsely pubescent.   Comments. This species seems to be common in the middle elevations of Sierra Nevada de Santa Marta. It has been collected in both forest areas and in shady coffee plantations; in the latter within logs of Erythrina sp. (Leguminosaeae), in which there were also colonies of Leptogenys sp. (Formicidae). Veturius impressus is the only Proculini endemic to the Caribbean region of Colombia. Fig. 29 Diagnosis. 47.5-50.4 mm total length. Anterior border of the labrum straight or slightly concave. Frontoclypeus almost straight. Frontal-clypeal suture absent. Internal tubercles small, joined to central tubercle by V-shaped ridges. Central tubercle small, with apex slightly free, upward projected; lateroposterior tubercles large, conspicuous, and transverse. Frontal fossae glabrous. Postfrontal groove complete. Mediobasal area of the mentum pubescent. Marginal groove over anterior border of the pronotum expanded. Mesosternum glabrous. Metasternum pubescent anterolaterally and in lateral groove; disc not delimited by punctations. Meso-and metatibiae unarmed. Humeri and epipleura glabrous; anterior vertical face of elytra pubescent. Fig. 30 Diagnosis. 41.4-46.5 mm total length. Anterior border of the labrum straight or slightly concave. Frontoclypeus almost straight. Frontal-clypeal suture absent. Internal tubercles small, joined to central tubercle by V-shaped ridges. Central tubercle small, with apex not free; lateroposterior tubercles large, distinct, and transverse. Frontal fossae with some setae. Postfrontal groove complete. Mediobasal area of the mentum glabrous. Marginal groove over anterior border of the pronotum expanded. Mesosternum glabrous. Metasternum pubescent anterolaterally and in lateral groove; disc not delimited by punctations. Mesoand metatibiae with a small spine. Humeri and epipleura glabrous; anterior vertical face of elytra pubescent.

Discussion
With 28 species, the Colombian Caribbean hosts an important richness of passalid species, which also exhibits a unique composition. The dry plain, characteristic of the lowlands of northern Colombia, is dominated by widely distributed species such as Passalus punctiger and P. interstitialis, and to a lesser extent by P. interruptus. Meanwhile, the mountainous systems