Corresponding author: Larry Jiménez-Ferbans (
Academic editor: A. Frolov
Employing data from literature, examination of specimens in collections, and a field trip, a list of the species of
Empleando datos de literatura, el examen de especímenes en colecciones y recolectados en campo, elaboramos una lista de las especies de Passalidae de Bolivia. Registramos un total de 38 especies, incluidos los nuevos registros de
Jiménez-Ferbans L, Reyes-Castillo P, Schuster JC (2019) The Passalidae (Coleoptera, Scarabaeoidea) from Bolivia, with the descriptions of three new species. ZooKeys 882: 51–85.
Pedro Reyes-Castillo conducted a field trip to Santa Cruz in February 2010 and the material collected is deposited in the collection of the Instituto de Ecología in Xalapa (
For every species in the list, we included the entomological collection where the specimens from Bolivia are deposited, the authors that have recorded the species, the material examined (labels cited verbatim and separated by slashes), and comments. The classification adopted and the terminology employed for the head is that proposed by
A total of 22 species has been recorded from Bolivia in the literature; meanwhile we found 25 species in the reviewed collections, including the specimens of
Holotype: female, pinned, BOLIVIA: COCHABAMBA, Prov. Carrasco, Yungas. ii.1971. alt. 3200 m. A. Martínez col. // Bosque húmedo de montaña de
Named after the country, Bolivia.
The anterior border of the labrum can be straight or slightly concave. The longitudinal sulcus on the area between frontal ridges can be weak or marked. Medial basal mentum can be fully pubescent or only laterally so.
Holotype: female, pinned, BOLIVIA: Yungas, Incachaca, 2800 m, xii.1960, Zischka leg. //
Among the brachypterous species of
Holotype: female, pinned, BOLIVIA: Yungas del Palmar, 15.iii.1958, 2000 m M. Zlsekka // “
This species is named in honor of Dr. Enio Cano from Guatemala, a passionate scholar of
Five punctures on the anterior half (paratype), punctations restricted to the lateral fossae (11 on right and 82 on the left).
The size of
Since the fauna of
1 | Clypeus hidden below frons, with anterior angles below mediofrontal tubercles (Fig. |
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– | Clypeus (frontoclypeus) exposed dorsally, with anterior angles in front of border of frons (Fig. |
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2 | Maxilla with lacinia unidentate or bidentate in apical third (Fig. |
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– | Maxilla with lacinia bidentate in apical third (Fig. |
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3 | Maxilla with lacinia unidentate in apical third (Fig. |
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– | Maxilla with lacinia bidentate in apical third (Fig. |
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4 | Dorsal mandibular tooth with a concave expansion (in dorsal view). Mesosternum smooth, without punctures over mesosternal scar. Body length 16.0–19.5 mm |
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– | Dorsal mandibular tooth thin, without a concave expansion. Mesosternum with punctures over mesosternal scar and beyond. Body length 14.0–16.0 mm |
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5 | Mesosternal scar oval, weakly defined, shiny. Metasternal fossae and epipleura glabrous. Body length 18.0–19.0 mm |
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– | Mesosternal scar elongate, well-defined, and rugose. Metasternal fossae and epipleura pubescent |
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6 | First lamella of antennal club reduced. Body length 22.7–23.1 mm (Fig. |
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– | First lamella of antennal club not reduced, almost equal in width to second lamella. Body length 18.5–19.5 mm |
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7 | Anterior border of frons straight or almost straight, without secondary mediofrontal tubercles (Fig. |
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– | Anterior border of frons with one or two secondary mediofrontal tubercles (Figs |
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8 | Antennal club with 5 lamellae, first two reduced (half width of third lamella). Body length 22.0–25.0 mm |
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– | Antennal club with three lamellae (Fig. |
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9 | Wings reduced (brachypterous) (Fig. |
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– | Wings fully developed (macropterous) |
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10 | Lateroposterior tubercles larger that central tubercle (Fig. |
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– | Lateroposterior tubercles smaller that central tubercle (Figs |
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11 | Internal tubercles joined to mediofrontal tubercles by a weak ridge. Frontal area, between frontal ridges, not divided by a longitudinal sulcus (Figs |
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– | Internal tubercles not joined to mediofrontal tubercles by a ridge (Fig. |
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12 | Anterior frontal border with strong median indentation, insinuating secondary mediofrontal tubercles (Fig. |
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– | Anterior frontal edge straight, without median indentation. Body length 32.0 mm |
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13 | Apex of central tubercle slightly free (the very tip not detached to the frontal ridgeds and frontal area). Body length 31.0–33.0 mm |
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– | Apex of central tubercle not free, fused with frontal ridges (Fig. |
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14 | Humeri with sparse pubescence at base. Body shorter (25.1–28.0 mm) |
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– | Humeri glabrous. Body longer (42.2–44.3 mm) (Fig. |
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15 | Anterior border of frons with one secondary mediofrontal tubercle. Central tubercle with apex not free. Hypostomal process with a matt groove over apex |
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– | Anterior border of frons with two secondary mediofrontal tubercles; if border straight, then central tubercle with apex distinctly free (“Petrejus” group). Hypostomal process without a matt groove over apex (subgenus |
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16 | Anterior border of frons with two secondary mediofrontal tubercles joined at bases |
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– | Anterior border of frons with or without mediofrontal tubercles, when present secondary mediofrontal tubercles separated |
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17 | Secondary mediofrontal tubercles large and fused with each other almost totally. Lateropostfrontal area glabrous. Body length 24.3–27.0 mm |
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– | Secondary mediofrontal tubercles only contiguous at base. Lateropostfrontal area pubescent. Body length 31.1–33.0 mm |
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18 | Central tubercle with apex very free, reaching or surpassing anterior border of frons. Secondary mediofrontal tubercles absent or rudimentary (“Petrejus” group) |
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– | Central tubercle with apex not free or slightly free (Fig. |
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19 | Central tubercle surpassing widely anterior margin of head, fused to median portion of head almost to anterior margin. Body length 24.0 mm |
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– | Central tubercle not fused to median portion of head |
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20 | Central tubercle concave at apex. Body longer (40.0–51.0 mm) |
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– | Central tubercle acute, not concave at apex. Body shorter (23.0–30.0 mm) |
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21 | Central tubercle strongly sulcate at base. Humeri pubescent. Body length 30.0 mm |
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– | Central tubercle not sulcate at base. Humeri glabrous. Body length 23.0–30.0 mm |
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22 | Habitus opaque. Body length 39.5 mm |
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– | Habitus shiny |
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23 | Mesosternal fossae glabrous or with only 1–3 setae ( |
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– | Mesosternal fossae densely pubescent |
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24 | Antennal club with four lamellae, fourth one very reduced and tomentose. Body length 27.1–34.0 mm |
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– | Antennal club with three lamellae |
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25 | Central tubercle very free, reaching anterior border of head. Pronotum pubescent on lateral fossae. Body length 36.0–45.1 mm |
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– | Central tubercle slightly free, not reaching anterior border of head. Lateral fossa of pronotum glabrous |
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26 | Last abdominal sternite with incomplete groove. Body longer (44.4–52.8 mm) (Fig. |
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– | Last abdominal sternite with medially complete groove. Body shorter (29.1–42.0 mm) (Fig. |
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27 | Central tubercle with apex very free, reaching anterior cephalic border. Body length 33.0–38.0 mm |
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_ | Central tubercle with apex not free or barely free. Body length 34.2–39.1 mm |
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28 | Frontoclypeal suture present |
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– | Frontoclypeal suture absent |
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29 | Antennal club with three lamellae. Body length 18.2–23.1 mm |
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– | Antennal club with four or five lamellae. Body length 17.0–21.0 mm |
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30 | Anterior labral border deeply concave, with an excavation behind concavity (dorsal depression sensu |
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– | Anterior labral border straight or slightly concave or convex, without an excavation behind border ( |
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31 | Brachypterous. Body length 34.0–45.0 mm |
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– | Macropterous (subgenus |
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32 | Mesosternum glabrous (not including anterior angles, which can have some scarce short setae) |
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– | Mesosternum with dense pubescence |
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33 | Central tubercle with apex free. Body length 36.0–40.0 mm |
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– | Central tubercle with apex not free |
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34 | Lateropostfrontal areas pubescent (rarely glabrous). Metasternum with pubescence beyond anterior border (mesocoxal cavity) and lateral fossa, reaching lateromedial metasternum. Body length 37.0–49.0 mm |
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– | Lateropostfrontal areas glabrous. Metasternum with pubescence restricted to anterior border (mesocoxal cavity) and lateral fossa |
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35 | Postfrontal groove (occipital sulcus sensu |
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– | Postfrontal groove present. Superior spurs of mesotibiae and metatibiae straight or almost straight. Body length 39.0–46.0 mm |
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36 | Lateropostfrontal area glabrous. Body length 33.0–41.0 mm |
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– | Lateropostfrontal area pubescent |
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37 | Lateropostfrontal area with 2–15 long setae. Central tubercle high, in lateral view higher than internal tubercles. Body length 30.0–37.0 mm |
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– | Lateropostfrontal area with 2–10 short setae. Central tubercle almost at same level of internal tubercles in lateral view. Body length 28.0–30.0 mm |
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Dado que la fauna de
1 | Clípeo oculto debajo de la frente, con ángulos anteriores debajo de los tubérculos mediofrontales (Fig. |
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– | Clípeo (clípeo-frente) expuesto dorsalmente, con ángulos anteriores por delante del borde frontal (Fig. |
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2 | Maxila con lacinia uni o bidentada en el tercio apical (Fig. |
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– | Maxilla con lacinia bidentada en el tercio apical (Fig. |
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3 | Maxilla con lacinia bidentada en el tercio apical (Fig. |
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– | Maxilla con lacinia unidentada en el tercio apical (Fig. |
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4 | Diente dorsal mandibular con una expansión cóncava (en vista dorsal). Mesosternón liso, sin puntos sobre la cicatriz mesosternal. Longitud total 16.0–19.5 mm |
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– | Diente dorsal mandibular delgado, sin expansión cóncava. Mesosternón con puntos sobre la cicatriz mesosternal y más allá. Longitud total 14.0–16.0 mm |
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5 | Cicatriz mesosternal oval, poco marcada y brillante. Foseta metasternal y epipleura glabras. Longitud total 18.0–19.0 mm |
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– | Cicatriz mesosternal alargada, bien definida y opaca. Foseta metasternal y epipleura pubescentes |
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6 | Primer artejo de la maza antenal reducido. Longitud total 22.7–23.1 mm (Fig. |
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– | Primer artejo de la maza antenal no reducido, de largo similar al segundo. Longitud total 18.5–19.5 mm |
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7 | Borde frontal anterior recto o casi recto, sin tubérculos mediofrontales secundarios. Tubérculo central corto, con ápice no libre (fusionado a la frente y quillas frontales) (Fig. |
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– | Borde frontal anterior con uno o dos tubérculos mediofrontales secundarios (Figs |
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8 | Maza antenal con cinco lamelas, las dos primeras reducidas. Longitud total 22.0–25.0 mm |
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– | Maza antenal con tres lamelas (Fig. |
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9 | Alas reducidas (braquíptero) (Fig. |
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– | Alas desarrolladas (macróptero) |
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10 | Tubérculos lateroposteriores de mayor tamaño que el tubérculo central (Fig. |
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– | Tubérculos lateroposteriores más pequeños que el tubérculo central (Figs |
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11 | Tubérculos internos unidos a tubérculos mediofrontales por una quilla débil. Área frontal, entre quillas frontales, no dividida longitudinalmente por un surco (Figs |
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– | Tubérculos internos no unidos a tubérculos mediofrontales por una quilla (Fig. |
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12 | Borde frontal anterior con fuerte hendidura media, insinuando dientes mediofrontales secundarios (Fig. |
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– | Borde frontal anterior sin hendidura media. Longitud total 32.0 mm |
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13 | Ápice del tubérculo central ligeramente libre, con solo una pequeña porción despegada de la frente. Longitud total 31.0–33.0 mm |
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– | Ápice del tubérculo central no libre, unido a la frente (Fig. |
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14 | Humeri con pubescencia escasa en la base. Talla pequeña (25.1–28.0 mm) |
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– | Humeri glabros. Talla grande (42.2–44.3 mm) (Fig. |
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15 | Borde frontal anterior con un tubérculo mediofrontal secundario. Proceso hipostomal con un surco mate sobre el ápice |
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– | Borde frontal anterior con dos tubérculos mediofrontales secundarios; si el el borde es recto, sin tubérculos, entonces el tubérculo central es muy libre (grupo "Petrejus"). Proceso hipostomal sin surco sobre el ápice (subgénero |
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16 | Borde frontal anterior con dos tubérculos mediofrontales secundarios, contiguos en su base |
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– | Borde frontal anterior sin tubérculos mediofrontales secundarios o con dos tubérculos separados en sus bases |
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17 | Tubérculos mediofrontales secundarios grandes, fusionados entre si en casi toda su extensión. Áreas lateroposfrontales glabras. Longitud total 24.3–27.0 mm |
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– | Tubérculos mediofrontales secundarios pequeños, solo contiguos en su base. Áreas lateroposfrontales pubescentes. Longitud total 31.1–33.0 mm |
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18 | Tubérculo central con ápice muy libre, alcanzando o sobrepasando el borde frontal anterior. Tubérculos mediofrontales ausentes o rudimentarios (grupo “Petrejus”) |
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– | Tubérculo central con ápice no libre o ligeramente libre (Fig. |
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19 | Tubérculo central sobrepasando ampliamente el margen de la frente, fusionado a la parte media de la cabeza, casi hasta el borde anterior. Longitud total 24.0 mm |
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– | Tubérculo central no fusionado a la parte media de la cabeza |
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20 | Tubérculo central con concavidad en el ápice. Talla grande (40.0–51.0 mm) |
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– | Tubérculo central sin concavidad en el ápice. Talla mediana (23.0–30.0 mm) |
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21 | Tubérculo central con surco marcado en la base. Humeri pubescentes. Longitud total 30.0 mm |
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– | Tubérculo central sin surco en la base. Humeri glabros. Longitud total 23.0–30.0 mm |
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22 | Habitus opaco. Longitud total 39.5 mm |
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– | Habitus brillante |
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23 | Foseta mesosternal glabra o con solo 1–3 sedas ( |
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– | Foseta mesosternal densamente pubescente |
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24 | Maza antenal con cuatro lamelas, la cuarta muy reducida y tomentosa…..27.1–34.0 mm |
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– | Maza antenal con tres lamelas |
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25 | Tubérculo central muy libre, alcanzando el borde anterior de la cabeza. Foseta lateral del pronoto pubescente. Longitud total 36.0–45.1 mm |
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– | Tubérculo central solo ligeramente libre, nunca alcanzando el borde anterior de la cabeza. Foseta lateral del pronoto glabra |
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26 | Surco marginal sobre último esternito abdominal incompleto. Talla grande (44.4–52.8 mm) (Fig. |
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– | Surco marginal sobre último esternito abdominal completo. Talla mediana a grande (29.1–42.0 mm) (Fig. |
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27 | Ápice del tubérculo central muy libre, alcanzando el borde frontal anterior. Longitud total 33.0–38.0 mm |
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– | Ápice del tubérculo central no libre o apenas ligeramente libre, no alcanzando el borde frontal anterior. Longitud total 34.2–39.1 mm |
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28 | Sutura frontoclipeal presente |
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– | Sutura frontoclipeal ausente (Fig. |
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29 | Maza antenal con tres lamelas. Longitud total 18.2–23.1 mm |
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– | Maza antenal con cuatro o cinco lamelas. Longitud total 17.0–21.0 mm |
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30 | Borde anterior del labro profundamente cóncavo, con una excavación por detrás de la concavidad ("dorsal depression" |
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– | Borde anterior del labro recto o ligeramente cóncavo o convexo, sin excavación por detrás del borde ( |
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31 | Braquíptero. Longitud total 34.0–45.0 mm |
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– | Macróptero (subgénero |
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32 | Mesosternón glabro (no incluyendo el ángulo anterior, que puede tener sedas cortas y dispersas) |
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– | Mesosternón con pubescencia abundante |
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33 | Tubérculo central con ápice libre. Longitud total 36.0–40.0 mm |
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– | Tubérculo central con ápice no libre |
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34 | Áreas lateroposfrontales pubescentes (raramente glabras). Metasternón con pubescencia más allá del borde anterior (cavidad metacoxal) y foseta lateral, alcanzando el área lateromedial del metasternón. Longitud total 37.0–49.0 mm |
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– | Áreas lateroposfrontales glabras. Metasternón con pubescencia restringida al borde anterior (cavidad metacoxal) y foseta lateral |
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35 | Surco posfrontal (occipital |
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– | Surco posfrontal presente. Espolones superiores de meso y metatibias rectos o casi rectos. Longitud total 39–46 mm |
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36 | Áreas lateroposfrontales glabras. Longitud total 33.0–41.0 mm |
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– | Áreas lateroposfrontales pubescentes |
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37 | Áreas lateroposfrontales con sedas largas (2–15 sedas). Tubérculo central alto, en vista lateral mucho más elevado que tubérculos internos. Longitud total 30.0–37.0 mm |
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– | Áreas lateroposfrontales con sedas cortas (2–10 sedas). Tubérculo central bajo, en vista lateral casi al mismo nivel que tubérculos internos. Longitud total 28.0–30.0 mm |
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Head and pronotum:
Bolivia has a total area of 1,098,581 km2 and its territory includes a high variety of ecosystems. The country is divided in 12 ecoregions (Ibisch et al. 2003), of which, the Southwest Amazonia, Cerrado, Chiquitania, and Yungas seem to be suitable for
The number of species known from Bolivia is small in comparison with other tropical countries of the New World. For example, Mexico, Guatemala, Colombia, and Brazil have more than 80 species recorded for each country (
Without doubt, the number of species of Bolivia is underestimated due to the lack of a systematic exploration of this country. Thus, more surveys are needed, especially in ecosystems such as montane forest and tropical rain forest, which normally harbor many species. Some departments with a domain of tropical rain forest have not been sampled for
From the total of 38 species listed above, we have studied material for 23 species. For the other 15 species, some authors have recorded specimens of all of them. However, three species can be discussed. The record of
Finally, the record of
We thank the curators of collections: Germán Amat (