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 6, 24, 36). Clunium with oval cuticular depression on central-median area. Females with sternum IX oval to subtrapeziform (Figures 18, 30, 42), with a generally globose or pear-shaped area in the middle.

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 1); phallosome with mesal endophallic sclerite widened, apically narrow and with two elongated teeth, one of them curved outwards (Figure 6).

Figures 1–6. 

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 3). Postclypeus brown, with diagonal dark brown stripes. Genae, anteclypeus, labrum, mandibles, maxillae, and labium brown to pale brown. Antennae brown, scape pale brown, flagellomeres distally cream. Maxillary palps pale brown, Mx4 dark brown distally. Tergal lobes of meso- and metathorax brown. Thoracic pleura pale brown. Mesopleura with dark brown spots. Legs: coxae, trochanter, and femora pale brown, tibiae and tarsi brown. Wings hyaline, forewing pterostigma with a large dark brown band proximally and distally. Abdomen cream, with subcuticular brown spots; clunium, hypandrium, and phallosome dark brown; epiproct and paraprocts brown.

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 1): L/W: 2.72. Pterostigma: lp/wp: 5.97, areola postica tall, triangular; al/ah: 1.27, R₄₊₅ almost straight, M five-branched, M₅ distally forked. Hindwings (Figure 2): l/w: 3.08. M four branched. Hypandrium (Figure 4), with three pigmented areas, two antero-lateral, curved, backwards and a central, posterior one, wide and narrow, setose as illustrated; phallosome with side struts independent, with two separate basal stems anteriorly wide, narrowing distally and curved outwards, basally articulated to a mesal process that projects to the hypandrium; external parameres laminar, dilated distally, apex rounded, bearing pores (Figure 6); anterior pair of endophallic sclerites oval, antero-lateral pair curved as illustrated, rounded distally and overlapping with the basal part of the external parameres (Figure 6). Paraprocts (Figure 5) broad, elliptic, with distal setal field as illustrated, sensory fields oval, with 34 trichobothria on basal rosettes. Epiproct (Figure 5) broadly trapeziform, with a group of three mesal macrosetae and a setal field distally on each side; posterior border with a field of microsetae and a row of four-five setae.

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 7); phallosome with narrow mesal endophallic sclerite, with distal processes curved as illustrated. Unlike C. occidentalis, the forewing has M five-branched, with M5 distally forked (Figure 7).

Figures 7–12. 

Colocania chicaque sp. n. Male. 7 Forewing 8 Hindwing 9 Front view of head 10 Hypandrium 11 Left paraproct and epiproct 12 Phallosome. Scale bars in mm.

(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 9). Postclypeus brown, with diagonal dark brown bands. Genae, anteclypeus, labrum, mandibles, maxillae and labium pale brown. Maxillary palps pale brown, Mx4 distally dark brown. Tergal lobes of meso- and metathorax brown. Thoracic pleura creamy to pale brown. Legs pale brown. Wings hyaline, forewing pterostigma as in diagnosis, veins brown. Abdomen pale brown, with subcuticular spots brown; clunium brown, clearer dorsally, with tubercle brown, hypandrium, phallosome, epiproct, and paraprocts brown.

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 7): L/W: 2.73. Pterostigma elongate: lp/wp: 5.88, areola postica tall, triangular: al/ah: 1.24, asymmetric; hindwings (Figure 8): l/w: 3.10, M four-branched. Hypandrium (Figure 10), setose as illustrated. Phallosome with side struts V-shaped, distally with arms curved outward; external parameres laminar, dilated distally, apex rounded, bearing pores (Figure 12); anterior pair of endophallic sclerites laminar-oval, with anterior margin serrate; antero-lateral pairs boomerang-shaped, curved inward proximally and with a series of laminar teeth, distally without teeth and reaching near the basal part of the external parameres (Figure 12). Paraprocts (Figure 11) robust, almost elliptic, with distal setal field as illustrated, sensory fields oval, with 26 trichobothria on basal rosettes. Epiproct (Figure 11) trapeziform, with a group of three mesal macrosetae and a setal field distally.

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 15). Postclypeus pale brown, with diagonal bands dark brown. Genae, anteclypeus, mandibles, maxillae, and labium pale brown, labrum brown. Antennae brown, scape and pedicel pale brown, flagellomeres cream apically. Maxillary palps pale brown, Mx4 brown distally. Tergal lobes of meso- and metathorax brown. Thoracic pleura creamy to pale brown with brown spots. Legs: coxae dark brown, fore- coxae pale brown, trochanters cream to pale brown, femora pale brown, with brown spots, tarsi 1 brown apically, tarsi 2–3 brown. Forewing with marginal brown band from apex of R4+5 to A1; M five branched; pterostigma with a dark brown band proximally and distally. Abdomen pale brown, with subcuticular brown spots; clunium brown, paler dorsally, with tubercle brown, subgenital plate with V-shaped pigmented area (Figure 16); gonapophyses, paraprocts, and epiproct dark brown.

As in diagnosis, plus the following: Head elongate (Figure 15): H/MxW: 1.41; H/D: 3.0, H/d: 4.0: IO/MxW: 0.78. Outer cusp of lacinial tip broad, with seven denticles. Mx4/Mx2: 1.14. Forewing (Fig. 13), L/W: 2.83. Pterostigma elongate: lp/wp: 5.73, areola postica triangular, al/ah: 1.35, M five branched. Hindwings (Figure 14): l/w: 2.89, M two branched. Subgenital plate (Figure 16) broad, slightly pointed posteriorly, setose, with macrosetae posteriorly. Gonapophyses (Figure 18): v1 elongate, pilose, distally acuminate; v2+3 pilose distally, with a row of 10 setae on v3; distal process acuminate, with microsetae on surface. Paraprocts (Figure 17) triangular, rounded apically, distal setal field with abundant setae as illustrated, sensory fields with 35 trichobothria on basal rosettes. Epiproct (Figure 17) triangular, rounded apically, with abundant setae and macrosetae distally as illustrated.

Figures 13–18. 

Colocania norsantanderina sp. n. Female. 13 Forewing 14 Hindwing 15 Front view of head 16 Subgenital plate 17 Epiproct and right paraproct 18 Sternum IX and left gonapophyses. Scale bars in mm.

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 19), and the hindwing M three-branched (Figure 20), and by details of the phallosome and hypandrium.

Figures 19–24. 

Colocania occidentalis sp. n. Male. 19 Forewing 20 Hindwing 21 Front view of head 22 Phallosome 23 Epiproct and right paraproct 24 Hypandrium. Scale bars in mm.

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 21), with pale cream band parallel to the antennal sockets. Postclypeus brown, with diagonal pale brown bands. Anteclypeus and labrum pale brown. Genae brown. Postgena pale cream. Antennae brown, flagellomeres cream apically. Maxillary palps pale brown, Mx4 dark brown distally. Tergal lobes of meso- and metathorax dark brown; thoracic pleura dark brown with pale spots. Legs pale brown, coxae with brown spots basally, femora with gray-brown ring basally and gray-brown spots widely distributed; tibiae with gray-brown spot, tarsi 1 brown apically; tarsi 2–3 brown. Wings hyaline, forewing pterostigma with one distal large dark brown band, some with additional proximal spot (not illustrated). Abdomen pale cream, with subcuticular bands brown; clunium and hypandrium brown; clunium with pale central area, hypandrium anteriorly cream; phallosome brown, with sclerites dark brown; epiproct and paraprocts brown.

As in diagnosis, plus the following: Head elongate (Figure 21): H/MxW: 1.44; H/D: 3.1, H/d: 4.19; IO/MxW: 0.78. Outer cusp of lacinial tip broad, with nine denticles. Mx4/Mx2: 1.10. Forewings (Figure 19): L/W: 2.69. Pterostigma elongate: lp/wp: 6.91, areola postica tall, triangular, with apex rounded: al/ah: 1.52; M four-branched. Hindwings (Figure 20): l/w: 3.02; M three branched. Hypandrium with three pigmented, setose areas, the two anterior widely separated and weakly connected to the posterior area, the latter narrow and extended laterally as a boomerang of little curvature (Figure 22). Phallosome (Figure 24) with laminar external parameres, wide, with apically rounded lobe bearing pores, curved inwards; mesal sclerite process with curved outwards apical teeth as illustrated. Side struts V-shaped, basally with an attached complex structure that projects over the hypandrium, distally curved outwards. Anterior endophallic sclerites laminar and oval, with serrate anterior margin, lateral sclerites curved as illustrated. Paraprocts (Figure 23) almost elliptic, with a distal setal field; sensory fields with 24 trichobothria on basal rosettes. Epiproct (Figure 23) semi-oval, rounded posteriorly, setal field with setae and macrosetae, posterior margin with small setae as illustrated.

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 27): H/MxW: 1.46; H/D: 3.3, H/d: 4.43; IO/MxW: 0.81. Outer cusp of lacinial tip broad, with eight denticles. Mx4/Mx2: 1.14. Wings (Figures 25 and 26) as in the male, L/W: 2.72. Pterostigma: lp/wp: 5.08, areola postica: al/ah: 1.23. Hindwings (Figure 26): l/w: 3.00. Subgenital plate (Figure 28) broad, posteriorly rounded, setose, with apical macrosetae as illustrated. Gonapophyses (Figure 30): v1 elongate, acuminate, distally with microsetae; v2+3 with short anterior heel, with four setae on v3; distal process sinuous, acuminate, bearing microsetae. Sternum IX broad, convex anteriorly, with a median concavity posteriorly, and a central, rounded mesal area as illustrated (Figure 30). Paraprocts (Figure 29) triangular, distal setal field with abundant setae and macrosetae as illustrated, sensory fields with 22–24 trichobothria on basal rosettes. Epiproct (Figure 29) triangular, with rounded apex, with abundant macrosetae distally, particularly on posterior margin as illustrated.

Figures 25–30. 

Colocania occidentalis sp. n. Female. 25 Forewing 26 Hindwing 27 Front view of head 28 Subgenital plate 29 Epiproct and right paraproct 30 Sternum IX and left gonapophyses. Scale bars in mm.

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 36).

Figures 31–36. 

Colocania pilosa sp. n. Male. 31 Forewing 32 Hindwing 33 Front view of head 34 Hypandrium 35 Left paraproct and epiproct 36 Phallosome. Scale bars in mm.

Color (in 80% ethanol). Head pattern (Figure 33). Compound eyes black, ocelli hyaline, with ochre centripetal crescents. Front and postclypeus pale brown, anteclypeus and labrum dark brown centrally, with sides pale brown. Genae pale brown with brown spots. Antennae brown, scape, pedicel and flagellomeres 1–3 pale brown, cream distally. Maxillary palps creamy, Mx4 distally brown. Tergal lobes of meso- and metathorax pale brown, with brown spots. Thoracic pleura creamy to pale brown, with brown spots. Legs pale brown, tarsi 2–3 dark brown, fore- coxae cream, middle- and hind- coxae brown, darker proximally, femora with brown spots. Forewing with marginal brown band from apex of R4+5 to A1; M five-branched; pterostigma with a proximal and a distal brown band. Hindwings hyaline, veins brown. Abdomen pale brown to cream, with subcuticular brown spots; clunium, epiproct, and paraprocts brown, epiproct with large, pale central area. Sclerites of hypandrium and phallosome dark brown.

As in diagnosis, plus the following: Head elongate (Figure 33): H/MxW: 1.37; compound eyes small, H/d: 3.94, H/D: 2.9, IO/MxW: 0.78. Outer cusp of lacinial tip broad, with eight denticles. Mx4/Mx2: 0.98. Forewings (Figure 31): L/W: 2.85. Pterostigma elongate: lp/wp: 5.56, areola postica tall, triangular, with rounded apex: al/ah: 1.40. M five-branched. Hindwings (Figure 32): l/w: 2.88. M three-branched. Hypandrium broadly trapeziform, setose, pigmented as illustrated in Figure 34. Phallosome (Figure 36), mesal endophallic sclerites elongate, shaped as an inverted V, basally separated, wide apically (mesal sclerite process) and overlapping on the external parameres and with short processes of rounded apices, with ante-apically lateral mesal sclerite process as illustrated; anterior endophallic sclerites oval and laminar, antero-lateral sclerites curved, boomerang-shaped as illustrated. Paraprocts (Figure 35) almost elliptic, with abundant setal field distally; sensory fields with 30 trichobothria on basal rosettes. Epiproct (Figure 35) semioval, rounded posteriorly, bearing macrosetae; lateral and mesal macrosetae as illustrated.

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 40); gonapophyses and paraprocts dark brown, epiproct brown.

As in diagnosis, plus the following: Head elongate (Figure 39): H/MxW: 1.43; H/D: 3.2, H/d: 4.48; IO/MxW: 0.79. Outer cusp of lacinial tip broad, with seven denticles. Mx4/Mx2: 1.15. Wings (Figures 37 and 38) as in the male, L/W: 2.78. Pterostigma: lp/wp: 5.13, areola postica: al/ah: 1.31. Hindwings (Figure 38): l/w: 3.02. Subgenital plate (Figure 40) broad, slightly pointed posteriorly, setose. Gonapophyses (Figure 42): v1 elongate, broad, acuminate, distally bearing microsetae; v2+3 pilose, with short proximal heel, 4 setae on v3; distal process sinuous, acuminate, with microsetae on surface distally. Paraprocts (Figure 41), sensory fields with 34 trichobothria on basal rosettes. Epiproct (Figure 41) broad, scutiform, widely rounded distally, setal field with abundant setae as illustrated, macrosetae and abundant setae distally curved, as illustrated.

Figures 37–42. 

Colocania pilosa sp. n. Female 37 Forewing 38 Hindwing 39 Front view of head 40 Subgenital plate 41 Epiproct and right paraproct 42 Sternum IX and left gonapophyses. Scale bars in mm.

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.

Comments on the species of Colocania gen. 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 48). The genus might also be found in Venezuela, as C. norsantanderina was found in a montane area near the Merida mountain range in Venezuela.

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 47).

Figures 43–46. 

Pterostigma (43, 44) and central area of clunium showing oval cuticular depression (45, 46) 43 Perucania longiareola 44 Ptiloneura bidorsalis 45 Colocania candelaria (male) 46 C. norsantanderina (female).

Figure 47. 

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.

Figure 48. 

Distribution of the species of Colocania.