Taeniolinum neusicus, Tulande-M & Prado & Triana, 2018

Tulande-M, Esteban, Prado, César Camilo & Triana, Hernán Darío, 2018, The first Taeniolinum from the Andes Mountains and Colombia (Chilopoda: Geophilomorpha), Zootaxa 4532 (1), pp. 113-124 : 114-121

Taeniolinum neusicus sp. n.

Type material: Holotype ♂, ( MPUJ _ ENT0049000 ): Colombia: Cundinamarca: Tausa: Parque Forestal Embalse del Neusa. Reference forest , point B-152: (5°9' 0.887"N / 73°56'21.454"W, WGS84 , 3100 m) 14 december 2015, Esteban Tulande-M. coll.

Paratypes 5 ♀, 1 specimen per voucher, ( MPUJ _ ENT0049082 ) and ( MPUJ _ ENT0049083 ): same data as holotype, 25 February 2015; ( MPUJ _ ENT0049064 ) : same data as holotype, except: point B-52 (5°9' 4.144"N / 73°56'25.350"W, WGS84 , 3100 m); ( MPUJ _ ENT0049016 ) : same as holotype except: point B-242 (5°8'56.330"N / 73°56'24.052"W, WGS84 , 3100 m), 26 July 2014; ( MPUJ _ ENT0049048 ) : same data as holotype except: point B221 (5°8' 57.633"N / 73°56' 27.299"W, WGS84 , 3100 m) 26 July 2014 . 5 ♂: ( MPUJ _ ENT0049090 ): same data as holotype except: point B-65 (5°9' 3.493"N / 73°56' 27.947"W, WGS84 , 3100 m); ( MPUJ _ ENT0049001 ): same as holotype, 12 April 2016; ( MPUJ _ ENT0049004 ) : same data as holotype except: Guanquica: point G-253 (5° 11' 26.078"N / 73° 56'9.731"W, WGS84 , 3050 m) 30 October 2014; ( MPUJ _ ENT0049018 ) and ( MPUJ _ ENT0049019 ): same as holotype except: Point B-140 (5°9' 0.889"N / 73°56' 29.247"W, WGS84 , 3100 m); 18 August 2015 .

Other material examined: same data as holotype, 2 specimens ( MPUJ _ ENT0048997 ), 1 ♂, 1 ♀ ( MPUJ _ ENT0049005 ), 2 specimens ( MPUJ _ ENT0049036 ), 1 specimen ( MPUJ _ ENT0049060 ), 1 ♀ ( MPUJ _ ENT0049063 ), 1 ♂ ( MPUJ _ ENT0049074 ), 1 specimen ( MPUJ _ ENT0049075 ), 1 specimen ( MPUJ _ ENT0049076 ). Colombia: Cundinamarca: Tausa: Parque Forestal Embalse del Neusa. Reference forest (5°8'0.8"N / 73°56'21"W to 5°9'4.7"N / 73°56'21"W, WGS84 ) Between July 2014 and April 2016: 1 ♀, 2 ♂ ( MPUJ _ ENT0049002 ), 1 ♂ ( MPUJ _ ENT0049003 ), 6 specimens ( MPUJ _ ENT0049006 ), 1 specimen ( MPUJ _ ENT0049007 ), 1 specimen ( MPUJ _ ENT0049009 ), 4 specimens ( MPUJ _ ENT0049010 ), 1 ♂ ( MPUJ _ ENT0049013 ), 1 ♂ ( MPUJ _ ENT0049015 ), 1 ♀ ( MPUJ _ ENT0049020 ), 2 ♀ ( MPUJ _ ENT0049022 ), 3 ♀ ( MPUJ _ ENT0049023 ), 2 specimens ( MPUJ _ ENT0049025 ), 4 specimens ( MPUJ _ ENT0049027 ), 1 specimen ( MPUJ _ ENT0049029 ), 2 specimens ( MPUJ _ ENT0049030 ), 1 ♀ ( MPUJ _ ENT0049034 ), 1 specimen ( MPUJ _ ENT0049035 ), 2 specimens ( MPUJ _ ENT0049039 ), 2 specimens ( MPUJ _ ENT0049041 ), 1 specimen ( MPUJ _ ENT0049042 ), 1 specimen ( MPUJ _ ENT0049043 ), 1 ♀ ( MPUJ _ ENT0049045 ), 1 specimen ( MPUJ _ ENT0049046 ), 1 specimen ( MPUJ _ ENT0049049 ), 5 specimens ( MPUJ _ ENT0049054 ), 5 specimens ( MPUJ _ ENT0049055 ), 2 specimens ( MPUJ _ ENT0049056 ), 1 specimen ( MPUJ _ ENT0049057 ), 1 specimen ( MPUJ _ ENT0049058 ), 1 ♀ ( MPUJ _ ENT0049059 ), 1 ♀ ( MPUJ _ ENT0049061 ), 1 ♂ ( MPUJ _ ENT0049062 ), 3 ♂ ( MPUJ _ ENT0049065 ), 1 ♀, 2 ♂ ( MPUJ _ ENT0049066 ), 1 ♂ ( MPUJ _ ENT0049067 ), 1 specimen ( MPUJ _ ENT0049068 ), 1 ♀ ( MPUJ _ ENT0049069 ), 1 ♂ ( MPUJ _ ENT0049070 ), 1 ♂ ( MPUJ _ ENT0049071 ), 1 ♂ ( MPUJ _ ENT0049072 ), 1 ♀ ( MPUJ _ ENT0049073 ), 1 ♀ ( MPUJ _ ENT0049077 ), 1 ♀ ( MPUJ _ ENT0049078 ), 1 ♂ ( MPUJ _ ENT0049079 ), 1 ♀ ( MPUJ _ ENT0049080 ), 1 ♀ ( MPUJ _ ENT0049081 ), 1 ♂ ( MPUJ _ ENT0049087 ), 1 ♀ ( MPUJ _ ENT0049088 ), 1 ♂ ( MPUJ _ ENT0049089 ), 1 specimen ( MPUJ _ ENT0049091 ), 1 specimen ( MPUJ _ ENT0049092 ), 1 ♀ ( MPUJ _ ENT0049014 ), 1 specimen ( MPUJ _ ENT0049084 ), 1 ♀ ( MPUJ _ ENT0049085 ), 1 ♂ ( MPUJ _ ENT0049086 ). Colombia: Cundinamarca: Tausa: Parque Forestal Embalse del Neusa : Laureles : formerly P. patula plantations (0.5 years after cut) (5°9'26"N / 73°56'25"W to 5°9'37"N / 73°56'31"W, WGS84 , 3100 m): 1 ♂ ( MPUJ _ ENT0048998 ), 2 ♂ ( MPUJ _ ENT0048999 ), 1 specimen ( MPUJ _ ENT0049008 ), 1 specimen ( MPUJ _ ENT0049017 ), 1 ♀ ( MPUJ _ ENT0049021 ), 1 ♂ ( MPUJ _ ENT0049024 ), 1 specimen ( MPUJ _ ENT0049028 ), 1 specimen ( MPUJ _ ENT0049031 ), 1 specimen ( MPUJ _ ENT0049033 ), 1 specimen ( MPUJ _ ENT0049050 ), 1 specimen ( MPUJ _ ENT0049051 ). Colombia: Cundinamarca: Tausa: Parque Forestal Embalse del Neusa Chapinero: formerly P. patula plantations (2.5 years after cut) (5°10'19"N / 73°57'8"W to 5°10'28"N / 73°57'17"W, WGS84 , 3000 m): 1 ♂ ( MPUJ _ ENT0049011 ), 1 specimen ( MPUJ _ ENT0049026 ), 4 specimens ( MPUJ _ ENT0049032 ), 2 specimens ( MPUJ _ ENT0049037 ), 2 specimens ( MPUJ _ ENT0049038 ), 1 specimen ( MPUJ _ ENT0049040 ), 1 specimen ( MPUJ _ENT:0049047), 1 specimen ( MPUJ _ ENT0049053 ). Colombia: Cundinamarca: Tausa: Parque Forestal Embalse del Neusa: Guanquica: formerly P. patula plantations (5 years after cut) (5°11'26"N / 73°56'8"W to 5°11'35"N / 73°56'19"W, WGS84 , 3000 m), 1 ♀ ( MPUJ _ ENT0049044 ), 1 specimen ( MPUJ _ ENT0049052 ); Colombia: Cundinamarca: Bogota Eastern hills: Quebrada La Vieja : (4°38'52.53"N / 74°02'45.23"W, WGS84 , 2700 m.) Cesar Camilo Prado. coll., 1 ♀ (CAUD-216.GEO-0036), 1 ♂ (CAUD-216.GEO-0037), 1 ♂ (CAUD-216.GEO-0038), 1 ♂ (CAUD- 216.GEO-0039), 1 ♂ (CAUD-216.GEO-0040), 1 ♂ (CAUD-216.GEO-0041), 1 ♀ (CAUD-216.GEO-0042), 1 ♀ (CAUD-216.GEO-0044), 1 ♂ (CAUD-216.GEO-0045) .

Diagnosis: A species of Taeniolinum with diminutive serrations at the internal border of the tarsungula, with a maximum body length of 24 mm, with 45 to 49 leg-bearing segments. T. neusicus sp. n. differs from T. interger ( Chamberlin, 1926) by the following set of characters (the ones from the latter in parentheses): ventral pores present generally from the second leg-bearing segment to the penultimate, when present from the first leg-bearing

segment never exceeding more than 3 pores (from the first segment to the penultimate, first with 9 pores), 14 to 18 claviform setae over the external border of the XIV antennomere (a.a.) and 3 over the internal border, (12 claviform setae over the external border of a.a. XIV and 1 over the internal).

T. neusicus sp. n. differs from T. arborum Pereira, Minelli & Barbieri, 1994 by the following set of characters (the ones from the latter in parentheses): internal border of a.a. XIV with 3 and external with 14-18 claviform sensilla (5 over internal and 5 over external border of a.a. XIV), with specialized setae type a and b on d. side of a.a. II (with a specialized type c setae on d. side of a.a. II), tarsungula with serrations over the external border (smooth without serrations).

T. neusicus sp. n. differs from T. guadeloupense Demange & Pereira, 1985 by the following set of characters (the ones from the latter in parentheses): tarsungula with serrations over the external border (smooth without serrations), 3 claviform setae over the internal border of the XIV antennomere (a.a.) (5-6 claviform setae over the internal border of a.a. XIV), with 1 type a, b and c specialized setae on d. side of a.a. V (with 1 type b and 3 type c setae on d. side of a.a. V).

Description of holotype: Male with 45 leg-bearing segments, body length 23 mm, maximum body width 0.65 mm, maximum width of cephalic plate 0.41mm, length of cephalic plate 0.41 mm maximum width of forcipular coxosternite: 0.46 mm. Color of cephalic plate and antenna orange, trunk yellow with some paramedian green patches over the paramedian pre and metatergites, these colors getting lost over time in ethanol.

Antennae: 2.11 times as long as cephalic plate, distally attenuate, ratio of width of a.a. I /width of a.a. XIV 1.8:1, the first a.a. almost two times wider than the last, from I to XIII a.a. goes progressively wider than long. Proportion of the length/width of the a.a. XIV 1.1: 1. ( Figure 1.C )

Ventral surface of a.a. I-IV with setae of various lengths and relatively few in number over a.a. I-VI, becoming progressively shorter and more numerous from a.a. VII - XIV. Dorsal chaetotaxy: Setae less numerous and longer than on ventral side, a.a. XIV with 18 claviform setae on the external margin of apical half and three on the internal ( Figure 1.C ); distal end of this a.a. with 9 hyaline specialized setae with an acute tip. Ventral and dorsal surface of a.a. II, V, IX and XIII with a series of very small specialized setae. Ventral setae located at the apical laterointernal, and middle areas, represented by two different types a and b (according to the classification proposed by Pereira et al. 1995). Type a setae very thin and acute, not split apically ( Figure 1 .C-a); type b setae bigger and with a trifid tip ( Figure 1 .C-b). Dorsal setae restricted to the apical lateroexternal and middle area of aforementioned a.a., represented by type a and b setae similar to the ones on the ventral side, bearing also a type c seta, similar to the type a but much longer, ocher and with a blunt tip. Number and distribution of these specialized setae over each a.a. as in Table 1.

Cephalic plate: Slightly wider than long (length/width ratio 0.85:1), anterior border edges rounded, lateral borders slightly rounded, posterior border almost straight with corners rounded. Form and chaetotaxy as in Figure 1.A .

Clypeus: With 5+5 postantennal setae and 1+1 setae at the mid clypeal area ( Figure 1.B ).

Labrum: mid piece well-developed and pigmented, with 16 unpigmented sharp triangular teeth, And with 1+1 more sharp and slightly sclerotized teeth at their sides (lateral pieces) ( Figure 1.B ).

Mandible: Dentate lamella not subdivided into blocks, 8 teeth in both mandibles, pectinate lamella with 21 or 22 hyaline teeth: with blunt tip and yellowish coloration ( Figure 2. E ).

First maxillae: Coxosternite without obvious lappets, devoid of setae at the dorsal or ventral surfaces, telopodite with lappets and ventrally bearing a 1+1 setae arrangement. Coxal projections subtriangular, rounded tip, provided with 1+1 long setae glabrous over the dorsal surface ( Figure 2.A ).

Second maxillae: Coxosternum divided by a shallow sulcus, with 2+2 short setae over the anterior border and 1+1 over the posterior one. Apical claw of telopodite well developed and bipectinate, ventral edge with 14 teeth, dorsal edge with 8 ( Figure 2.A ).

Forcipular segment: Telopodites not reaching clypeal anterior border, forcipular tergite rectangular. Coxosternite without chitinous lines, anterior border without denticles, forcipular coxosternite width/length ratio 1.64: 1. All articles of telopodites without denticles, ratio length of forcipular tarsungula/ length of trochanterprefemur 1.3: 1; internal border of the tarsungula with 21 diminutive serrations going from the base to the median zone. Calyx of the venom gland located between the base of the tarsungula and the tibia, very similar to the one of T. integer (see Pereira et al. 2000, fig. 18).

Metasternites of leg-bearing segments I to penultimate: Ventral pore-fields present in an uninterrupted series from sternite 2 to penultimate, first 25 sternites with pore-fields grouped elliptically, the rest of pore-fields grouped in one row, all pore-fields located very close to the posterior border ( Figure 2.D ) number of ventral pores of some metasternites: II (4); III (7); IV (9); V (14); VI (15); VII (16); VIII(18); IX (24); X (24); XI (21); XII (28); XIII (27); XIV (26); XV(27); XVI (24); XVII (20); XVIII (23); XIX (20); XX (18); XXI (17); XXII (18); XXIII (17); XXIV (17); XXV (13); XXVI (15); XXVII (14); XXVIII (13); XXIX (15); XXX (10); XXXI (11); XXXII (10); XXXIII (8); XXXIV (8); XXXV(11); XXXVI (11); XXXVII (10); XXXVIII (6); XXXIX (9); XL (6); XLI (8); XLII (5); XLIII (3); XLIV (2), Figure 2.D .

Ultimate leg-bearing segment: Intercalar pleurites present at both sides of ultimate pretergite. Ultimate presternite divided along the sagittal plane, width/length of tergite 1.34:1 width/length of sternite 1.16:1, coxopleura very slightly prominent at their distal end. Two single (“homogeneous”) coxal organs on each coxopleuron, opening on the membrane between coxopleuron and sternite. Posterior coxal organs bigger than the anterior ones. ( Figure 2.C )

Postpedal segments: Genital segments with 15 long setae and gonopods with 6+6 setae, pleuras apparently present ( Figure 2.C ).

Description of female paratype (MPUJ_ENT 0049064): Female with 47 leg-bearing segments, body length 23mm, maximum body width 0.77mm, differs with holotype in labrum, and number of ventral pores, and ultimate leg-bearing segment size, the rest of taxonomical characters as in the holotype.

Labrum: Mid-piece well-developed and pigmented, with 15 sharp triangular teeth without pigmentation. Lateral pieces with 2+2 slightly sclerotized teeth, longer and sharper than those at the mid-piece.

Metasternites of leg-bearing segments 1 to penultimate: Ventral pore-fields grouped as in holotype, in an uninterrupted series from sternite 2 to penultimate. Number of ventral pores of some metasternites: II (8); III (10); IV (11); V (16); VI (11); VII (17); VIII(18); IX (22); X (27); XI (24); XII (23); XIII (27); XIV (27); XV(26); XVI (29); XVII (26); XVIII (20); XIX (22); XX (18); XXI (20); XXII (19); XXIII (17); XXIV (13); XXV (13); XXVI (13); XXVII (14); XXVIII (12); XXIX (13); XXX (13); XXXI (10); XXXII (20); XXXIV (11); XXXV(13); XXXVI (10); XXXVII (10); XXXVIII (9); XXXIX (11); XL (18); XLI (13); XLII (8); XLIII (9); XLIV (5); XLV (6); XLVI (3).

Ultimate leg-bearing segment: Presternite divided by its sagittal plane, width/length of tergite: 1.14: 1; width/ length of sternite: 1.17: 1, Posterior coxal organs bigger than the anterior ones.

Postpedal segments: Genital segment with 6 long setae and gonopods with 2+2 setae, pleura apparently present.

Variations in other adults: Body length of males and females ranging between 20 mm to a maximum of 24 mm.

Leg-bearing segments: 45, 47 and 49 in males; 43, 45, 46 and 47 in females,

Labrum: Different number of teeth at the mid-piece: 14, 16, 17, 18, 21 and 23 were observed, also some of these teeth are not triangular and sharp as in holotype, instead they got a rounded tip. The number of teeth at the lateral-pieces varied as well, different combinations: 1+2, 2+2 and 3+3 being the 2+2 form the most common, and only one individual does not have teeth at the lateral pieces at all.

Clypeus: Different arrangements of postantennal setae: 2+2, 3+3, 4+4, 5+5 and 6+8. Setae at the median clypeal area as in the holotype for all individuals.

Metasternite of leg-bearing segments 1 to penultimate: Ventral pore-fields of some males present in an uninterrupted series from sternite 1 to penultimate, females as in holotype. Number of ventral pores of some metasternites: I (2), (3); II (4), (6), (7), (8), (10), (13), (14), (15); V (14), (16), (18), (21), (27), (28), (33); X (22), (24), (27), (34), (38), (46); XV (40), (52), (60), (65); XX (18), (21), (44), (51), (35), (59); XXV (14), (16), (27), (42), (40), (47); XXX (27), (13), (19), (34), (30); penultimate: (4), (2), (1).

Subadults: Body length from 7 mm up to a maximum of 19 mm.

Leg-bearing segments: two female individuals with 43 leg-bearing segments, the rest ranging between 45 to 47.

Clypeus: The distribution of postantennal setae varies in different arrangements: 2+2, 2+3, 3+3, 4+4, 4+5, 5+4, 5+5, 5+6, 5+7, 6+4, 6+6 and 7+6. Setae at the median piece of clypeus as in the holotype, with the exception of two males which had a prelabral setae

Labrum: different number of teeth at the mid-piece was observed for subadults: 12, 14, 15, 16, 17, 18, 19, 20, 21 and 23. The lateral pieces have also different arrangements: 1+1, 1+2, 2+2, 3+3, 4+3, 4+4 y 5+6.

Metasternites of leg-bearing segments 1 to penultimate: Ventral pores of the subadults present from the first leg-bearing segment to the penultimate or present from the second to the penultimate. Number of pores in some of the leg-bearing segments: I (2), (3); II (2), (3), (6), (8), (9), (15), (22); V (6), (10), (13), (15), (16), (17), (21), (27), (33); X (12), (17), (23), (29), (32), (43); XV (12), (22), (25), (43); XX (5), (9), (12), (18), (22), (44); XXV (6), (19), (20), (25), (36); XXX (13), (17), (21), (25), (23); Penultimate (4), (2), (3), (1).

Etymology: The specific epithet is a latinized adjective, in masculine form, derived from the name of the type locality, Parque Forestal Embalse del Neusa.

Type locality: Colombia: Eastern Andes Mountains: Cundinamarca: Parque Forestal Embalse del Neusa and Bogotá Eastern hills Quebrada La Vieja .

Habitat and ecological considerations. This is the first record of the genus Taeniolinum in Colombia and the Andes Mountains, the specimens were collected at two tropical high montane forest patchs ( Figure 3 ) within an altitudinal range of 2700 to 3200 meters above sea level, which is a new distribution range for Taeniolinum , as well as the maximum altitude at which it the genus has been recorded.

The individuals collected at Parque Forestal Embalse del Neusa, correlated positively with high values of Aluminum (Al = 5 - 28 cmol / kg) and sodium (Na = 0.02-0.22 cmol / kg) also showed a preference for acid soils (pH <3.5), and temperatures between 9 and 11 C°, the number of individuals observed decreased when the values of pH (> 3.5) and temperature (> 11 C°) increased. The thickness of the litter layer (horizon O) also influenced the number of individuals observed, having preference for the thicker layers typical of tropical high montane forests. At the forest relict, a total of 22 individuals / m 2 was estimated, the litter was composed mainly of leaves of Weinnmania tomentosa L.F, Chusquea scandens , Clusia multiflora Kunth, and Myrcine guianesis.

T. neusicus sp. n. may feed mainly of enchytraenid worms, collembolans, and dipteran larvae/pupa, because these groups were abundant in the monoliths in which T. neusicus sp. n. was found ( Figure 4 ).

T.neusicus sp. n. was also found in areas that were formerly Pinus patula plantations, in 2009 the environmental authority of the park: Corporación Autónoma Regional de Cundinamarca decided to carry out the removal of these plantations. The new species was found at three of these areas with different ages post clearcutting: 0, 2.5 and 5 years after cut. The areas that were formerly P. patula plantations are meant to be subjects of ecological restoration process to recover native montane forest, and taking into account that the number of individuals of this species varies depending on the age after felling, we see promising the implementation of T. neusicus sp. n. as a model for the monitoring and design of future restoration and conservation programs.

Morphologic intraspecific variation analysis. The width of the cephalic plate (Ce.W), was the variable best explained by the other variables (83%) followed by the length of the antenna (Ant.L = 75%) and length of the ultimate tergite (Ult.L = 75%), the number of legs and the number of labral teeth were the variables that were less explained R2 = 11% and 35% respectively ( Table 2).

The width of the last tergite showed the greatest number of significant correlations (7) followed by the width of the cephalic plate and length of the antennae (6), body length and length of the last tergite, each with 4 correlations. No significant difference was found between males and females for each of the variables ( Figure 5 ).

Figures

FIGURE 1. Taeniolinum neusicus sp. n., holotype (MPUJ_ENT0049000). A) Cephalic plate, B) Clipeus (Clip) and labrum (lab), C) Rigth Antennae (dorsal) showing the location of specialized setae (type a, type b and type c).

FIGURE 2. Taeniolinum neusicus sp. n., holotype (MPUJ_ENT0049000). A) First and second maxilae, B) Forcipular segment, C) ultimate leg-bearing segment; (pps) post pedal segment, (cp) coxal pores, D) Metasternites of leg-bearing segments, E) mandible; (dl) dentate lamella, (pl) pectinate lamella.

FIGURE 3. Habitat of Taeniolinum neusicus sp. n. tropical montane forest at 3000 m a.s.l, located inside Parque Forestal Embalse del Neusa, Colombia, Cundinamarca. (Photo: Laura Quintero).

FIGURE 4. Specimen of Taeniolinum neusicus sp. n. found inside a dipteran pupa, probably while brooding. (Photo: Esteban Tulande-M).

FIGURE 5. Correlation plot of morphological characters of T. neusicus sp. n; Long = Body lenght, Leg.p = Number of leg pairs, Ce.L= Cephalic plate lenght, Ce.W = Cephalic plate width, Ant.L = antennae length, Lab.med = median labral teeth, Lab.lat = lateral labral teeth Lab.T = Total labral teeth, Clip= clipeal setae, Ult.L= ultimate tergite length, Ult.W = Ultimate terguite width. Color indicating direction of correlation, size of the circle indicating strenght of correlation.