Research Article |
Corresponding author: Ranulfo González-Obando ( ranulfo.gonzalez@correounivalle.edu.co ) Academic editor: Kazunori Yoshizawa
© 2018 Ranulfo González-Obando, Alfonso N. García Aldrete, Nancy Carrejo-Gironza, Julian Mendivil.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
González-Obando R, Aldrete ANG, Carrejo-Gironza N, Mendivil J (2018) A new genus of Ptiloneuridae, its position within the family, and descriptions of five species (Psocodea, ‘Psocoptera’). ZooKeys 780: 11-34. https://doi.org/10.3897/zookeys.780.26753
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Upon examination of ptiloneurid specimens recently collected in forests of natural areas and Natural National Parks in Colombia, ten males and seven females were found that represent five species of an undescribed ptiloneurid genus. It differs from the other genera in the family by characters of the wings, hypandrium, phallosome, epiproct, female gonapophyses, and sternum IX. An identification key to the males of the genera of Ptiloneuridae, and a key to the species of the new genus are presented. A test on the validity and monophyly of the new genus, and its position within the family was also conducted.
biodiversity, Colombia, neotropics, taxonomy
Ptiloneuridae, a family of Psocodea (Psocomorpha: Epipsocetae), presently includes the genera Brasineura Silva-Neto & García Aldrete, Euplocania Enderlein, Loneura Navás, Loneuroides García Aldrete, Omilneura García Aldrete, Perucania New & Thornton, Ptiloneura Enderlein, Ptiloneuropsis Roesler, Timnewia García Aldrete, Triplocania Roesler, and Willreevesia García Aldrete.
Within the framework of the project “Revisión Taxonómica y Endemismo de los Psócidos (Insecta: Psocodea: ‘Psocoptera’) de Areas Protegidas de Colombia”, 17 specimens of Ptiloneuridae, corresponding to five species that could not be assigned to any of the known ptiloneurid genera were found. These specimens were collected in localities of Valle del Cauca, Meremberg Nature Reserve and La Candelaria (Huila), Planes de San Rafael Nature Reserve (Risaralda), National Natural Park Tamá (Norte de Santander), and Chicaque Nature Reserve (Cundinamarca) (Figure
Seventeen specimens were available for study, ten males and seven females. Nine specimens were dissected in 80% ethanol, and their parts (head, right legs and wings, and genitals), were mounted in Canada balsam, following standard procedures. Before dissecting, whole specimens were placed in 80% ethanol and observed at 50× to record color. Standard measurements were taken with a filar micrometer, and are given in mm. Abbreviations of parts measured are as follows: FW and HW: right fore- and hind- wing lengths, F, T, t1, t2 and t3: lengths of femur, tibia and tarsomeres 1, 2 and 3 of right hind leg, f1-fn: lengths of flagellomeres 1-n, Mx4: length of fourth segment of right maxillary palp, IO: minimum distance between compound eyes, D and d: antero-posterior and transverse diameter, respectively, of right compound eye, all in dorsal view of head. PO: d/D, H: head length, MxW: maximum head width, Ratio head length (H)/(D), L/W: forewing length/forewing width, lp/wp: pterostigma length/pterostigma width, al/ah: areola postica length/tall, l/w: hindwing length/hindwing width. The specimens studied are deposited in the Entomological Museum, Universidad del Valle (MUSENUV), Santiago de Cali, Colombia.
To test the validity and the monophyly of the new genus, and to establish its position within the family, a cladistic analysis was performed, using the free Software TNT 1.1 (
As in
Data matrix of morphological characters used for the cladistic analysis of Ptiloneuridae and outgroup (S. caatinga). For characters 1–19 see Silva-Neto et al. (2016).
Character | |||||||||||||||||||||||
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 | |
S. caatinga | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | - | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
B. diamantina | 1 | 1 | 2 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 |
C. candelaria | 0 | 3 | 2 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 2 | 1 | 1 |
C. chicaque | 0 | 3 | 2 | 0 | 0 | 0 | 0 | 0 | - | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 2 | 1 | 1 |
C. occidentalis | 0 | 2 | 1 | 0 | 0 | 0 | 0 | 0 | - | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 2 | 1 | 1 |
C. pilosa | 0 | 2 | 2 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 2 | 1 | 1 |
E. cerata | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 |
L. colombianus | 0 | 3 | 2 | 1 | 0 | 0 | 0 | 1 | 2 | 2 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 2 | 0 | 0 |
L. jinotegaensis | 0 | 3 | 3 | 0 | 0 | 0 | 0 | 0 | 1 | 2 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 |
O. circumvittata | 0 | 0 | 3 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 |
P. bidorsalis | 0 | 4 | 5 | 0 | 0 | 0 | 0 | 0 | 2 | 2 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 |
P. diamantina | 0 | 3 | 2 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 |
P. longiareola | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 3 | 3 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
T. cervantesi | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 |
T. magnifica | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 2 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 |
T. greeni | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 2 | 2 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
W. dominica | 0 | 0 | 2 | 0 | 0 | 0 | 0 | 0 | 1 | 4 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 1 | 0 | 0 |
The 19 characters used by
20. Costal margin, between base and nodus: (0) straight (1) convex (Figures
21. Shape of proximal area of pterostigma (at around 1/3 of its length): (0) not petiolated (Figure
22. Anterior region of hypandrium: (0) without ringed, short digitiform process, (1) with vestigial or short ringed digitiform process (Figures
23. Oval cuticular depression on central-median area of clunium: (0) absent, (1) present (Figures
Colocania occidentalis González, García Aldrete & Mendivil, sp. n.
The generic name is a compound word, formed with the root “colo” from Colombia, and the suffix “cania”, common in Ptiloneuridae, as in Euplocania, Perucania, Triplocania. It refers to its endemicity in Colombia.
Forewings with costal margin, between base and nodus, strongly to gently convex, pterostigma elongate, petiolate proximally at around 1/3 of its length, Rs curved strongly towards pterostigma; anterior area of the hypandrium with a short ringed digitiform process. Phallosome most commonly with two separate stems proximally usually projected to the hypandrium (Figures
1 | Hindwing M one-branched | 2 |
– | Hindwing M two to five-branched | 7 |
2 | Forewing 2A joining wing margin; no crossveins between 2A and wing margin | 3 |
– | Forewing 2A joining 1A; one crossvein between 2A and wing margin; two crossveins between 1A and wing margin | Timnewia García Aldrete |
3 | Forewing areola postica high, with apex rounded | 4 |
– | Forewing areola postica low, very long | Perucania New & Thornton |
4 | Labral sclerites incomplete, not reaching anterior margin of labrum | 5 |
– | Labral sclerites complete, almost reaching anterior margin of labrum | Willreevesia García Aldrete |
5 | Forewing M three-branched, occasionally M3 forked | Triplocania Roesler |
– | Forewing M more than three-branched | 6 |
6 | Forewing M four to five-branched | Euplocania Enderlein |
– | Forewing M six-branched | Omilneura García Aldrete |
7 | Forewing areola postica free, high, with apex rounded | 8 |
– | Forewing areola postica high, rigidly triangular, joined to M by a crossvein | Ptiloneuropsis Roesler |
8 | Forewing 2A without one crossvein to wing margin pterostigma long, smooth | 9 |
– | Forewing 2A with one crossvein to wing margin, pterostigma long, distinctly spurred | Loneuroides García Aldrete |
9 | Forewing with costal margin, between the base and the nodus strongly to gently convex, pterostigma proximally petiolate at around 1/3 of its length, Rs strongly curved towards pterostigma; hypandrium strongly convex, of one piece; anterior area of the hypandrium with a short ringed digitiform process, sometimes visible only by the presence of concentric rings | Colocania gen. n. |
– | Forewing with costal margin between the base and the nodus almost straight, pterostigma proximally little petiolate; Rs slightly curved towards pterostigma or almost straight; hypandrium simple or with three or more well-defined sclerites | 10 |
10 | Forewing M four to seven-branched; hindwing M two to five-branched | 11 |
– | Forewing M eight-branched; hindwing M five-branched; hypandrium a broad sclerite projected posteriorly to form a wide, three lobed sclerite, with a dense field of setae on each postero-lateral corner, and a dense field of setae on each side of the central projection | Ptiloneura Enderlein |
11 | Hypandrium simple, of one sclerite; hindwing M two to four-branched; phallosome with external parameres distally forked or rounded, enclosing a membrane with numerous pores | Brasineura Silva Neto & García Aldrete |
– | Hypandrium simple or formed by a central sclerite and one or two smaller ones on each side; hindwing M two to five-branched; phallosome with external parameres without a membranous region with numerous pores | Loneura Navás |
From 88 most parsimonious cladograms, obtained for each K value analyzed (K = 1–9), 17 consensus topologies were retained (L = 75–76, CI = 48–49, RI = 61–62). In all of them the clade formed by the species of Colocania was maintained, although between K = 5–9 the relations of this with the rest of the genera is unstable. This however, remains constant between K = 1–4 (L = 76, CI = 48, RI = 61), which includes the value calculated as appropriate (K = 2) through the use TNT script (setk.run) on the data set. In this way, the strict consensus topology obtained in the cladistic analyses with different optimality criteria of parsimony, showed that Colocania is a monophyletic group, supported by high symmetric resampling (88%) and Bootstrap (82%) (Figure
COLOMBIA. Huila. Belén, La Candelaria, 2128 m. 02°13'36.0"N, 76°07'27.4"W.
Holotype male. 18.IV.2015. On tree trunk. R. González. Deposited in Entomological Museum, Universidad del Valle (MUSENUV, slide code 29033).
Forewings hyaline, without marginal bands as in C. chicaque sp. n., and C. occidentalis sp. n., differing from them by wing characters, phallosome, and details of hypandrium and epiproct. Unlike the above species, the pterostigma has large pigmented bands proximally and distally (Figure
Colocania candelaria sp. n. Male. 1 Forewing 2 Hindwing 3 Front view of head 4 Hypandrium 5 Left paraproct and epiproct 6 Phallosome. Abbreviations: (aes) anterior endophallic sclerites, (ep) external parameres, (les), lateral endophallic sclerite, (ms) mesal sclerite, (msp) mesal sclerite processes, (st) side struts. Scale bars in mm.
Color (in 80% ethanol). Body pale brown, with pigmented dark brown areas as indicated below. Compound eyes black, ocelli hyaline, with ochre centripetal crescents. Vertex with three dark brown areas, a central one and two lateral ones between the compound eyes. Front with brown area between ocellar group and epistomal sulcus, as illustrated (Figure
As in diagnosis, plus the following: Head elongate: H/MxW: 1.52; small compound eyes, H/d: 4.24; H/D: 3.1, IO/MxW: 0.80. Upper ends of compound eyes almost reaching the level of the vertex. Outer cusp of lacinial tip broad, with six denticles. Mx4/Mx2: 1.13. Forewing (Figure
FW: 6250, HW: 4075, F: 1470, T: 2560, t1: 900, Mx4: 360, ctt1: 26, f1: 1230, f2: 1240, f3: 1000, f4: 760, f5: 480, f6: 410, f7: 320, f8: 295, f9: 210, f10: 200, IO: 580, D: 360, d: 260, IO/d: 2.23, PO: 0.72.
The specific epithet refers to the town of La Candelaria (La Plata, Huila) where the holotype was collected.
COLOMBIA. Cundinamarca. Soacha, Chicaque Natural Reserve, Tirolesa, 2240 m. 04°36'40.38"N, 74°18'41.7"W.
Holotype male. 24–28.II.2014. Malaise trap. D. Forero. Deposited at the Pontífica Universidad Javeriana Museum (MPUJ). Bogotá, Colombia.
Forewings hyaline, without marginal bands as in C. candelaria sp. n., and C. occidentalis sp. n., differing from them by characters of the forewing, phallosome and details of hypandrium and epiproct. Unlike C. candelaria the pterostigma has only a distal pigmented band, sometimes with a median, little defined, small pigmented area (Figure
(in 80% ethanol). Body pale brown, with pigmented dark brown areas as indicated below. Compound eyes black, ocelli hyaline, with ochre centripetal crescents. Vertex with brown areas, central and lateral, between compound eyes. Front brown as illustrated (Figure
As in diagnosis, plus the following: Head elongate: H/MxW: 1.48, H/D: 3.0; H/d: 4.0; IO/MxW: 0.74. Vertex slightly emarginated; upper ends of compound eyes almost reaching the level of the vertex. Outer cusp of lacinial tip broad, with eight denticles. Mx4/Mx2: 1.09. Forewing (Figure
FW: 6700, HW: 4500, F: 1610, T: 2760, t1: 1240, t2: 100, t3: 152, Mx4: 380, ctt1: 42, f1: 1480, f2: 1380, IO: 600, D: 400, d: 300, IO/d: 2, PO: 0.75.
The specific epithet refers to the Chicaque Natural Reserve, where the holotype was collected.
COLOMBIA. Norte de Santander. National Natural Park Tamá. Orocue Station, 2433 m. 07°25'27.6"N, 72°26'31.8"W.
Holotype female. 2.VII.2016. On wood beam with mosses and fungi. R. González, N. Carrejo. Deposited in Entomological Museum, Universidad del Valle (MUSENUV, slide code 29034), Santiago de Cali, Colombia.
Related to C. pilosa sp. n. in having the forewing with similarly pigmented pattern and shape of the areola postica. It differs from the female of C. pilosa by having the hindwing M widely forked, by the shape of sternum IX, and by the shape and pilosity of the epiproct, without distally curved setae.
(in 80% ethanol). Body pale brown, with pigmented dark brown areas as indicated below. Compound eyes black, ocelli hyaline, with ochre centripetal crescents. Vertex and front with brown spots, central and lateral between the compound eyes (Figure
As in diagnosis, plus the following: Head elongate (Figure
FW: 6075, HW: 3975, F: 1460, T: 2700, t1: 1170, t2: 105, t3: 160, Mx4: 375, ctt1: 46, f1: 1320, f2: 1330, f3: 1030, IO: 620, D: 370, d: 273, IO/d: 2.27, PO: 0.74.
The specific epithet refers to the Colombian Department Norte de Santander, where the holotype was collected.
COLOMBIA. Valle del Cauca. Santiago de Cali, El Saladito, San Antonio, 2142m. 03°29'23.5"N, 76°37'39.4"W.
Holotype male. 27.I.2012. On tree trunks covered with lichens and mosses. R. González. Deposited in Entomological Museum, Universidad del Valle (MUSENUV, slide code 29035), Santiago de Cali, Colombia. Paratypes: 2 males, same data as the holotype, J. Mendivil & R. González. 1 male, 2 females, Valle del Cauca, Santiago de Cali, Los Andes-Charco Azul, 1687 m. 03°25'21.7"N, 76°37'0.1"W, 23.I.2013. R. González (Female: MUSENUV, slide code 29036). 2 males, 2 females, Los Andes-Quebrada Honda, 1900 m. 03°26'01.8"N, 76°38'40.3"W, 23.I.2013. R. González. 1 male, 1 female, Risaralda, Santuario, Planes de San Rafael, 2092 m. 05°07'13.9"N, 76°00'04.5"W, R. González (male: MUSENUV, slide code 29037). All paratypes on tree trunks covered with lichens and mosses.
Forewings hyaline, without marginal pigmented band as in C. candelaria sp. n., and C. chicaque sp. n., differing from them by having the forewing M four-branched (Figure
Color (in 80% ethanol). Body cream to pale brown, with pigmented dark brown areas as indicated below. Compound eyes black, ocelli hyaline, with ochre centripetal crescents. Vertex with dark brown spots, central and lateral between compound eyes. Front with brown spots between ocellar group and epistomal sulcus, as illustrated (Figure
As in diagnosis, plus the following: Head elongate (Figure
FW: 4375, HW: 2987, F: 1310, T: 2050, t1: 990, t2: 96, t3: 135, Mx4: 320, ctt1: 38, f1: 1000, f2: 990, f3: 770, f4: 510, f5: 355, f6: 300, f7: 240, f8: 200, f9: 176, f10: 150, f11: 180, IO: 535, D: 320, d: 235, IO/d: 2.28, PO: 0.73.
Color (in 80% ethanol). Body, head, legs, epiproct, paraprocts, and wings as in the male, plus the following: Subgenital plate with pigmented area V-shaped, arms wider proximally. Clunium, epiproct and paraprots brown. Gonapophyses dark brown. Sternum IX cream yellowish, darker on the edges.
As in diagnosis, plus the following: Head elongate (Figure
FW: 4350, HW: 2925, F: 1300, T: 2050, t1: 940, t2: 104, t3: 124, Mx4: 330, ctt1: 35, f1: 1130, f2: 1100, f3: 820, f4: 580, f5: 365, f6: 290, f7: 210, IO: 570, D: 312, d: 230, IO/d: 2.48, PO: 0.74.
The specific name refers to the distribution of the species in localities of the western Andean Cordillera in Colombia.
COLOMBIA. Huila. La Plata, Belén, Meremberg Nature Reserve, 2352 m. 02°13'06.6"N, 76°07'01.1"W.
Holotype male. 20.I.2015. On tree trunks covered with lichens and mosses. R. González. Deposited in Entomological Museum, Universidad del Valle (MUSENUV, slide code 29038), Santiago de Cali, Colombia. Paratype female, 28.X.2016. Same data as the holotype (MUSENUV, slide code 29039).
It is related to C. norsantanderina sp. n., described above (see diagnosis of the latter). Both species seem to have a sister-group relationship, to be confirmed when the male of the latter be found. The ninth sterna and gonapophyses in both species are quite similar, but the epiprocts are distinct. The male presents side struts with arms independent, proximally paralell, with processes directed outwards, unique among the species of the genus, distally curved outwards and not fused to the external parameres (Figure
Color (in 80% ethanol). Head pattern (Figure
As in diagnosis, plus the following: Head elongate (Figure
FW: 6475, HW: 4025, F: 1450, T: 2570, t1: 1180, t2: 120, t3: 160, Mx4: 325, ctt1: 36, f1: 1150, f2: 1120, f3: 900, f4: 690, f5: 410, f6: 360, f7: 260, f8: 230, f9: 200, f10: 173, f11: 200, IO: 610, D: 370, d: 270, IO/d: 2.26, PO: 0.73.
Color (in 80% ethanol). Body, head, legs, wings, epiproct and paraprocts as in the male, plus the following: pigmented area of subgenital plate U-shaped (Figure
As in diagnosis, plus the following: Head elongate (Figure
FW: 5550, HW: 3775, F: 1440, T: 2530, t1: 1110, t2: 114, t3: 140, Mx4: 380, ctt1: 40, f1: 1090, f2: 1115, f3: 880, IO: 620, D: 350, d: 250, IO/d: 2.48, PO: 0.71.
The specific epithet refers to the densely pilose female epiproct.
1 | Females | 2 |
– | Males | 4 |
2 | Forewing with a marginal pigmented band brown to yellowish from R4+5 to A1, M five branched (Figures |
3 |
– | Forewing hyaline, without marginal pigmented band; with apical dark spot in pterostigma (Figure |
C. occidentalis sp. n. |
3 | Epiproct and paraprocts strongly setose, macrosetae strongly curved distally (Figure |
C. pilosa sp. n. |
– | Epiproct and paraprocts not as above, macrosetae not strongly curved distally (Figure |
C. norsantanderina sp. n. |
4 | Forewing with a marginal pigmented band, brown to yellowish from R4+5 to A1, M five branched (Figure |
C. pilosa sp. n. |
– | Forewing hyaline, without marginal pigmented band as above, pterostigma with proximal and distal, or only distal pigmented bands, M four-five branched (Figures |
5 |
5 | Forewing M four branched (Figure |
C. occidentalis sp. n. |
– | Forewing M five branched, M5 bifurcate (Figures |
6 |
6 | Pterostigma with large proximal and distal pigmented bands (Figure |
C. candelaria sp. n. |
– | Pterostigma with only distal pigmented band (Figure |
C. chicaque sp. n. |
Colocania is, so far, endemic to Colombia, its species have been collected between 1687 and 2450 meters of altitude, in Andean and Subandean areas (see Rangel and Aguilar 1995) of Central and East-western mountain ranges (Figure
On basis of the forewing pigmentation pattern, two species groups are recognized, group A, with the forewings having a marginal pigmented band, from R4+5 to distal end of Cu2, including C. norsantanderina sp. n. and C. pilosa sp. n., and group B, with forewings hyaline, including C. candelaria sp. n., C. chicaque sp. n. and C. occidentalis sp. n. However, C. candelaria and C. chicaque are related only on the basis of one homoplasic character, the forewings with a ratio of crossvein Rs-M and the portion of vein M before it of 1:2 (char. 12) (Figure
Strict consensus cladogram for 13 genera of Ptiloneuridae under implied weights for K = 2 (L = 76; IC = 48; IR = 61). Full circles (black, red, or green) indicate unique character changes, black circles (unambiguous), color circles (ambiguous); empty circles indicate parallelisms or reversals. ACCTRAN (green circles) and DELTRAN (red circles) optimization. Character number above and character states below each circle. Bootstrap (bold numbers on the branches) and Symmetric resampling (bold numbers under the branches) values>50.
We wish to thank Colciencias and Universidad del Valle, Vicerectoría de Investigaciones, Santiago de Cali, Colombia, for two grants to the authors (Projects Col – CT 550-2014 and CI 7874) to study Psocoptera. We thank Posgrado de Biología, Facultad de Ciencias Naturales y Exactas of Universidad del Valle for sponsoring this publication. We thank Dimitri Forero for the loan of the holotype of C. chicaque. We also thank Svanhild Büch and Rubén Luna of the Meremberg Nature Reserve, Ovidio Ledesma, from Planes de San Rafael (Risaralda), Libardo Suárez Fonseca and Orlando Benítez of the National Natural Park Tamá. We thank Oscar Saenz (Universidad del Valle, Santiago de Cali, Colombia) for the phylogenetic analysis. RGO, NSC and JMN thank Departamento de Biología, Facultad de Ciencias Naturales y Exactas, Universidad del Valle, Santiago de Cali, Colombia, for research support. ANGA thanks Instituto de Biología, Universidad Nacional Autónoma de México, for continuous research support.