Scatimus strenua Martínez-Revelo, Lopera-Toro, & Medina, 2020

Martínez-Revelo, Diego Esteban, Lopera-Toro, Alejandro & Medina, Claudia A., 2020, Review of Scatimus Erichson (Coleoptera: Scarabaeidae) in Colombia with the description of a new species, Zootaxa 4890 (4), pp. 521-534 : 523-529

Scatimus strenua Martínez-Revelo, Lopera-Toro, & Medina , new species

( Figs. 1 A–B, 2A–B, 3B, D–F, H, 4D, H, L, P, 5–6)

Type material. Holotype. “ Colombia: Antioquia: El Carmen / de Viboral, Finca la Meseta, Bosque montano bajo, / 5° 52’/ 45.05’’N 75° 12’2.33’’W, WGS84 / 1646 m, T.Exc.H. # T10 _24, 4.xii. / 2016, A. Lopera, J. Cárdenas ”— male (IAvH-E-196594). Printed on red label: “ Scatimus strenua Martínez-Revelo, Lopera-Toro & Medina sp. nov. ” Paratypes [7 males, 5 females]. Colombia: Antioquia: El Carmen de Viboral, Finca San José, Bosque montano bajo, Trampa de Excremento Humano , # T4 _48, 5°53’16.68’’N 75°11’37.11’’W, 1198 m , WGS84 , 2.xii.2016, A. Lopera, J. Cárdenas— 2 females (IAvH-E-196589-90) ; # T6 _24, 5°53’16.49’’N 75°11’40.59’’W, 1214 m , 1.xii.2016 — 1 female, (IAvH-E-196652); # T7 _24, 5°53’14.54’’N 75°11’41.36’’W, 1237 m , 1.xii.2016 — 1 male, (IAvH-E-196588); # T7 _48, 2.xii.2016 — 1 male (IAvH-E-196591) ; # T8 _48, 5°53’13.01’’N 75°11’41.63’’W, 1250 m , 2.xii.2016 — 1 male (IAvH-E-196653); # T9 _24, 5°53’13.08’’N 75°11’39.77’’W, 1125 m , 1.xii.2016 — 1 male (IAvH-E-196586); # T9 _48, 2.xii.2016,— 2 males (IAvH-E-196592-93); Finca la Meseta , Bosque montano bajo , # T1 _48, 5°52’58.07’’N 75°11’55.32’’W , WGS84 , 1437 m, 5.xii.2016 — 1 male (IAvH-E-196595) ; # T2 _48, 5°52’56.41’’N 75°11’54.70’’W, 1494 m , 5.xii.2016 — 1 female (IAvH-E-196596); Finca la Samaria , Bosque montano bajo , # T2 _72, 5°53’20.0’’N 75°11’13.2’’W, 1166 m , 2.xii.2016 — 1 female (IAvH-E-196585). Printed on yellow labels: “ Scatimus strenua Paratype Martínez-Revelo, Lopera-Toro & Medina sp. nov. ”

Non-type material examined. Colombia: Antioquia: Amalfi, Vereda Salazar (Bodega Vieja), 6°58’31.9’’N 75°5’2.9’’W, 1310 m, 3.v.2009, Grisales, S. —1 sex undetermined ( MEFLG) .

Diagnosis. Scatimus strenua is close to S. strandi but can be distinguished from this and other species of the S. strandi clade by the following characters: anterior margin of metafemur enlarged and very sharp in S. strenua (411), simply rounded in S. cribosus (410). Median lobe of metaventrites anterior edge arcuate medially in S. strenua (251, Fig. 3E ), distinctly angular medially in S. pacificus , S. furcatus , and S. quadricuspis (250). Body elongate in dorsal view in S. strenua (11, Fig. 1 A–B, E), body oval in S. onorei and S. erinnyos (10, Fig. 1 C–D). Surface of abdominal ventrites with weakly defined microsculpture on segment 3–4 laterally in S. strenua (473, Fig. 3F ), only on segment 3 in S. onorei (474). Lateral surface of mesepimeron lacking transverse carina medially in S. strenua (272, Fig. 3D ), present in S. erinnyos and S. strandi (271, Fig. 3C ). Surface around dorsal portions of eye lacking microsculpture in S. strenua (90), present in S. erinnyos (91) and surface of mesofemur and metafemur with minute puncturesthrough-out in S. strenua (380, Fig. 3H ), with several coarse and umbilicate punctures at apex anteriorly in S. erinnyos (381, Fig. 3G ). Lateral marginal bead of pronotum distinctly reduced medially in S. strenua (160, Fig. 3B ) and sharply defined medially in S. strandi (161, Fig. 3A ).

Description. Holotype male, ( Fig. 1A ). Pinned. Measurements. Body length 5.84 mm, elytral width 3.73 mm. Body elongate, lateral edges parallel on median third in dorsal view (11, Fig. 1 A–B, E). Color. Black, with some brown reflections. Head ( Figs. 2 A–B). Dorsal surface of head glabrous, lacking setiferous punctures (51). Surface of clypeus and gena smooth, lacking wrinkles or transverse rugulae (30), with scattered, fine punctures more pronounced towards the margins. Head with a single frontal carina between the eyes, arcuate medially, with lateral extremities anteriorly directed (63). Anterior edge of clypeus with two moderately developed and reflexed teeth, clypeal median emargination angulate (40), lateral margins arcuate between teeth and gena, disc of clypeus concave, vertical surface of clypeus V-shaped with a small central depression in the lower edge. Clypeogenal suture pronounced, surface of gena adjacent to the eye lacking coarse punctures (70). Smooth eyes completely divided by canthus, dorsal portion of the eye approximately twice as long as wide (80), surface around dorsal portions of eye lacking microsculpture (90). Antennomeres 7–8 with pit on distal surface (101). Pronotum ( Fig. 3B ). Surface smooth. Anterior edge with membranous projection at head insertion (111). Lateral edge of pronotum simple, lacking setiferous punctures (121), with eight coarse setiferous punctures present in lateral fossae and three anteriorly to fossae (131), non-setiferous punctures each with a minute granule medially (141). Lateral declivity of pronotum lacking setiferous punctures (151), lateral marginal bead of pronotum distinctly reduced medially (161), posterior margin of pronotum lacking coarse punctures (171). Elytra. Elytral disc smooth and shiny, with scattered minute punctures (3.5 X). Anterolateral angle of elytron slightly produced (180). Striae distinctly impressed with oval to rounded punctures separated by 3–4 times diameters on disc and lacking setiferous punctures on interstriae. Striae 1–3 with punctures larger, deeper and more pronounced and fused on apical declivity, striae 7 incomplete on posterior third, striae 8 deeply impressed with interstria lacking setiferous punctures (192). Apical declivity of elytron lacking setiferous punctures on interstriae (201). Proepipleural sulcus shallower on anterior half with a distinct depression (fovea) adjacent to the anterior angle (211). Thoracic ventrites ( Fig. 3D, F ). Prosternum behind procoxa lacking setiferous punctures (223), with few rounded punctures aligned with the posterior edge, margin of the punctures incomplete. Anterior region of propleuron strongly excavated and delimited posteriorly by a complete propleural carina, surface of excavated portion with rivose microsculpture and fine setae. Area posterior of propleuron with rounded punctures located on the posterior edge and reaching half the margin around procoxa. Mesoventrite with a moderately deep anteriomedian fossa and with setae anteriorly (231), lacking a median longitudinal carina (240). Mesoepisternal suture well defined, thicker near the procoxa, and thinner towards the lateral edge. Disc of mesoventrite with dense ocellate punctures. Median lobe of metaventrite anterior edge arcuate medially (251), with straight lateral borders. Medial edge of mesocoxal cavity at least almost straight on a short distance (260). Lateral surface of mesepimeron lacking transverse carina medially (272), anteromedial surface punctate (280). Posterior portion of metepisternon rounded (291), posteromedial area of metepisternon flat (301), lateral lobes of metaventrite lacking a posterior transverse row of punctures (310). Legs ( Fig. 3H ). Protibiae tridentate on the outer margin, basal tooth smallest. External teeth of protibiae with additional small indentations at basal angles (351), dorsoapical transverse carina of protibiae emarginate (361). Ventral surface of protibiae with two rows of setae, the outer row with large curved apex setae, and the inner row with smaller straight setae. Ventral surface of profemora lacking setiferous punctures (321), with a row of long setae on the anterior margin and scattered coarse punctures in the posterior margin. Trochanto-femoral pit of prothoracic leg rounded (340). Procoxa with anteromarginal sulcus (331). Mesotibiae with two transverse carinas on the external surface (421). Surface of mesofemur with minute punctures throughout (380). Ventral surface of mesocoxa with coarse setiferous punctures, setae longer than punctures diameter (372). Metatibiae with two transverse carinae on the external surface. Dorsal surface of metatibiae with a longitudinal medial row of straight setae and strong microsculpture on external half only (431), lacking dorsal accessory setiferous punctures (441). Ventral portion of metatibiae broadly arcuate in cross section near apex (451). Anterior edge of metafemur lacking marginal bead (400), with anterior margin enlarged and very sharp (411). Dorsal surface of metafemur with minute punctures throughout. Longitudinal median sulcus of metacoxa sharply defined posteriorly (391). Abdominal ventrites ( Fig 3F ). Median portion of abdominal ventrites unmodified, each suture clearly visible throughout (461). Surface of abdominal ventrites with weakly defined microsculpture laterally on ventrites 3 and 4 (473), abdominal ventrites 7 and 8 feebly grooved along suture (49:1). Pre-pygidium is clearly visible, with a central, longitudinal, deep sulcus. Pygidium. Basal sulcus of pygidium narrowed and shallowly impressed medially (500). Surface of pygidium lacking setiferous punctures (512). Transversal sulcus well defined with the superior border thickened towards the middle. Aedeagus ( Fig. 4D, H ). Parameres approximately conical (521). Ventral surface of paramere with strong microsculpture (531), parameres apex not projecting ventrally (540). Dorsal portion of paramere sclerotized throughout (550). Paramere hook feebly developed (561). Endophallus ( Figs. 4L, P ). Internal sac with four apical endophallites, the basal semicircular (Bsc), frontolateral peripheral (FLP) endophallites and lamella copulatrix (LC) are absent. The superior right peripheral endophallite (SRP) is semicircular, with a broader and rounded extremity, narrow and curved in its middle part, and ending in a handle-shaped projection with bilobed apex. The additional ( AS) sclerite is short, with a forked apex. The subaxial (SA) and axial (A) endophallites are superposed. The SA endophallite is elongate, one extremity with a truncated apex and the other one with a forked apex. The A endophallite is elongate the same size as SA, one extremity with an acute apex and the other one rounded.

Female. ( Fig. 1B ). Body length 6.63 mm, elytral width 4.18 mm. Similar to the male except in: clypeus with four teeth (11), middle ones larger, pronotum with one to seven large rounded punctures anterior to and five to ten in the lateral fossae abdominal ventrite 7 parallel sided (481).

Variation. Male: body length 5.61–6.27 mm, elytral width 3.77–4.35 mm. Pronotum with minimum two to maximum eight large rounded punctures anterior to and five to ten in the lateral fossae. Female: body length 5.11– 6.63 mm; Elytral width 3.24–4.18 mm.

Etymology. The species epithet “ strenua ” refers to a Roman goddess of the new year, purification, and well-being, and should be treated as a noun in apposition. The name is dedicated to the courageous women from this area of Colombia, that have overcome the armed conflict that affected their region in the 1980s and 1990s.

Ecology and distribution. The landscape inhabited by S. strenua is part of the “Rio Melcocho” watershed. Vegetation cover is mostly forest classified by Espinel (2011) as Very Humid pre-Montane forest with influence of sub-Andean and Andean forests (van der Hammen & Rangel 1997). Scattered pastures and plantations are located on the few flat areas of these mountains. The tree canopy average height is 25 m with emergent trees of up to 35 m. Although the canopy is mostly continuous with abundant epiphytes, most timber producing trees have been already harvested, and the remaining forest can be considered as a mature secondary forest. The soils are well drained and covered by a leaf litter layer 10–15 cm thick. Biogeographically, these forests belong to the Magdalena Valley mountain forests in the Tumbes-Chocó-Magdalena province, a key biological hotspot. The new species is part of a dung beetle ( Scarabaeinae ) ensemble of approximately 45 species including other endemic beetles such as Dichotomius andresi Sarmiento & Amat and Cryptocanthon parvus Howden , and moist forest beetles such as Sylvicanthon aequinoctialis (Harold) , Scybalocanthon moniliatus (Bates) , and Sulcophanaeus noctis (Bates) , frequently collected in the Magdalena Valley.

On the phylogenetic placement of Scatimus strenua within the genus Scatimus . The cladistic analysis resulted in a single parsimonious cladogram with length: 122 steps, CI = 0.680, RI = 0.811. The topology of the strict consensus cladogram ( Fig. 5 ) is similar to the one obtained by Génier and Kohlmann (2003). The three species groups were recovered, the S. ovatus clade was recovered as monophyletic with the following non-ambiguous synapomorphies: mesoventrites with a deep anteromedian fossa and with setae medially (232), dorsal portion of paramere widely membranous at juncture medially (551), hook of paramere strongly developed (562).

The S. cucullatus clade also was recovered as monophyletic, but with ambiguous synapomorphies: body in dorsal view elongate, lateral edges parallel on median third (11), head with a single straight frontal carina (60), and anterolateral angle of elytron subquadrate (181).

Scatimus strenua clustered within the S. strandi clade, supported by a single non-ambiguous synapomorphy: head with a single frontal arcuate carina with lateral extremities anteriorly directed (63), and forming a monophyletic unit with S. onorei + S. erinnyos + ( S. strandi + S. strenua ), supported by the following two ambiguous synapomorphies: non setiferous punctures of pronotal lateral declivity each with a minute granule medially (141), and posteromedial area of metepisternon flat or concave (301).

Updated species checklist. Two species of the S. ovatus clade, S. ovatus and S. fernandezi were already confirmed for Colombia by Medina et al. (2001). Escobar (2000) first reported S. strandi in Colombia but neither he nor Medina et al. (2001) included a locality, possibly causing Génier & Kohlmann (2003) to miss the presence of the species in Colombia. Recently, the presence of S. strandi was confirmed in the department of Meta (Cárdenas et al. 2020) and we here report three additional localities for the species in the departments of Caquetá and Nariño. Scatimus fernandezi previously was recorded in Casanare ( Medina et al. 2001; Génier & Kohlmann 2003) and we here add new departmental records from Meta and Putumayo. Scatimus ovatus was reported in Medina et al. (2001) without a precise locality; Génier and Kohlmann (2003) registered it in Magdalena; Solís et al. (2011) mentioned “confer” Scatimus ovatus in Atlántico; González-Alvarado & Medina (2015) reported the species in Bolívar, Magdalena, Sucre, Tolima, and Santander; and Mendivil et al. 2020 from Caldas. Antioquia, Boyacá, and La Guajira are new departmental records ( Table 1 ).

A fifth species could be added to the Colombian list when expeditions are performed on the border with Panama. Scatimus erinnyos is registered in Cerro Pirre and the Estación Ambiental Cana (7°45.32’N, 77°41.07’W), Panama , approximately 8 km from the department of Chocó in Colombia . It is possible that S. erinnyos is distributed throughout the continuous Chocó-Darién moist forest, that includes the departments of Antioquia, Cauca, Chocó , Nariño, and Valle del Cauca in Colombia .

Figures

FIGURE 1. Species of Scatimus from Colombia.A, Scatimus strenua holotype łmale (Antioquia, 5°52’45.05’’N 75°12’2.33’’W); B, Scatimus strenua, paratype female (Antioquia, 5°53’16.68’’N 75°11’37.11’’W); C, Scatimus fernandezi male (Casanare, 5°16’20.9”N 72°24’2.6”W); D, Scatimus ovatus male (Tolima, 5°8’58.3’’N 74°47’6.1’’W); E, Scatimus strandi male (Nariño, 0°30’N 77°13’W). Scale bars: 2 mm.

FIGURE 2. Scatimus heads in dorsal view. A, Scatimus strenua male (Antioquia, 5°52’45.05’’N 75°12’2.33’’W); B, Scatimus strenua female (Antioquia, 5°53’16.68’’N 75°11’37.11’’W); C, Scatimus erinnyos male (Costa Rica, Parque Nacional Guanacaste); D, Scatimus strandi male (Nariño, 0°30’N 77°13’W); E, Scatimus fernandezi (Casanare, 5°16’20.9”N 72°24’2.6”W); F, Scatimus ovatus (Tolima, 05°08’58.3’’N 74°47’06.1’’W). Scale bars = 0.5 mm.

FIGURE 3. Scatimus pronotum, mesepimeron, metasternum, abdomen, and metafemur. A, Scatimus strandi pronotum (lateral view); B, Scatimus strenua pronotum (lateral view); C, Scatimus strandi mesepimeron (ventral view); D, Scatimus strenua mesepimeron (ventral view); E, Scatimus strenua metasternum (ventral view); F, Scatimus strenua abdomen; G, Scatimus erinnyos metafemur (ventral view); H, Scatimus strenua metafemur (ventral view). Scale bar = 0.5 mm.

FIGURE 4. Male genitalia of Scatimus from Colombia. A, Scatimus fernandezi aedeagus (lateral view, scale bar = 0.5 mm); B, Scatimus ovatus aedeagus (lateral view, scale bar = 0.5 mm); C, Scatimus strandi aedeagus (lateral view, scale bar = 0.5 mm); D, Scatimus strenua aedeagus (lateral view, scale bar = 0.5 mm); E, Scatimus fernandezi parameres (ventral view, scale bar = 0.25 mm); F, Scatimus ovatus parameres (ventral view, scale bar = 0.25 mm); G, Scatimus strandi parameres (ventral view, scale bar = 0.25 mm); H, Scatimus strenua parameres (ventral view, scale bar = 0.25 mm); I, Scatimus fernandezi superior right peripheral sclerite (SRP, scale = 0.25 mm); J, Scatimus ovatus superior right peripheral sclerite (SRP, scale = 0.25 mm); K, Scatimus strandi superior right peripheral sclerite (SRP, scale = 0.25 mm); L, Scatimus strenua superior right peripheral sclerite (SRP, scale = 0.25 mm); M, Scatimus fernandezi additional (AS), subaxial (SA), and axial (A) sclerites superposed, scale bar = 0.25 mm; N, Scatimus ovatus additional (AS), subaxial (SA), and axial (A) sclerites superposed, scale bar = 0.25 mm; O, Scatimus strandi additional (AS), subaxial (SA), and axial (A) sclerites superposed, scale bar = 0.25 mm; P, Scatimus strenua additional (AS), subaxial (SA), and axial (A) sclerites superposed, scale bar = 0.25 mm.

FIGURE 5. Results of the phylogenetic analyses. A, The single most parsimonious tree from TNT, with character evolution of Scatimus; all characters states are treated as unordered and equally weighted. Characters were mapped on branches using unambiguous character changes in Winclada (black circles, non homoplasious changes; white circles, homoplasious changes); numbers above the circles indicate characters, and number below circles indicate their states. B), Strict consensus of the five most parsimonious trees (length = 122 steps; consistency index = 0.680; retention index = 0.811) obtained with heuristic search. Clade support is indicated on branches (Bremer support values/bootstrap proportions).