Research Article |
Corresponding author: Catalina Romero-Ortiz ( icromeroo@unal.edu.co ) Academic editor: Jana Christophoryová
© 2023 Catalina Romero-Ortiz, Carlos E. Sarmiento, Mark S. Harvey.
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:
Romero-Ortiz C, Sarmiento CE, Harvey MS (2023) A new genus and five new species of pseudoscorpions (Arachnida, Pseudoscorpiones, Withiidae) from Colombia. ZooKeys 1184: 301-326. https://doi.org/10.3897/zookeys.1184.106698
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The pseudoscorpion family Withiidae is widely distributed around the world, with most of its diversity in tropical areas. Five new species and a new genus from Colombia are described: Cystowithius florezi sp. nov., Parawithius bromelicola sp. nov., Oligowithius achagua sp. nov., and the genus Paciwithius gen. nov. with two species Paciwithius valduparensis sp. nov. and Paciwithius chimbilacus sp. nov. A reassessment of the subgenus Dolichowithius (Oligowithius) Beier, 1936 allows the elevation to a full generic level, and the transfer of the only known species to Oligowithius, forming the new combination Oligowithius abnormis (Beier, 1936), comb. nov.
Arachnids, bromeliads, Colombia, diversity, taxonomy
With a surface area of more than 1.1 million square kilometers, Colombia is considered a hot-spot of biodiversity with nearly 2,000 species of birds (
The pseudoscorpion family Withiidae Chamberlin, 1931 comprises more than 170 species arranged in 37 genera. The family is divided into two subfamilies, Paragoniochernetinae and Withiinae, with the latter divided into four tribes, Cacodemoniini, Juxtacheliferini, Protocheliferini, and Withiini. The status of these groups is highly uncertain, with only the Cacodemoniini seemingly supported by a strong synapomorphy in the male genitalia (
In this paper, we describe a new genus, Paciwithius gen. nov. with two new species, describe one new species of Cystowithius, register a new record of the species Cystowithius ankeri Garcia & Romero, 2021 and propose an identification key for the species of this genus, describe a new species of the genus Parawithius, and provide a reassessment of the subgenus Dolichowithius (Oligowithius) Beier, 1936.
The specimens examined for this study are lodged in the Arachnological collection at the
Instituto de Ciencias Naturales – Universidad Nacional de Colombia (
Measurements were taken at the highest magnification possible using an ocular graticule, taken to the nearest 0.005 mm. After study, the specimens were rinsed in water and returned to vials with the other body parts submerged in 75% ethanol. Dissected portions were placed in small capillary tubes sealed with cotton on both ends. The type specimens are deposited in the
Arachnology collection at the Instituto de Ciencias Naturales-Universidad Nacional de Colombia (
Terminology and measurements largely follow
The following abbreviations were used for the trichobothria of the movable finger b = basal; sb = sub-basal; st = sub-terminal; t = terminal. For the trichobothria of the fixed finger: eb = exterior basal; esb = exterior sub-basal; est = exterior sub-terminal; et = exterior terminal; ib = interior basal; isb = interior sub-basal; ist = interior sub-terminal; it = interior terminal. For the cheliceral setae: b = basal seta; es = exterior seta; is = interior seta; ls = laminal seta. For the abdominal setae: gls = glandular setae.
Paciwithius chimbilacus sp. nov.
Paciwithius can be distinguished from all other withiids by the absence of tergal keels, absence of eyespots, absence of a tactile seta on tarsus IV, the trichobothrium it is midway between isb and et, the presence of a patch of glandular setae on sternite VIII with an elongated shape, and the first three to six tergites are entire. The male genitalia have dorsal apodemes extending over its entire length, lateral apodemes extending over most of the ejaculatory canal atrium, ejaculatory canal curved dorsally in lateral view and lateral rods straight, extending beyond the lateral apodemes in dorsal view. This genus was mentioned as “nr. Victorwithius” in
The genus name refers to the Latin for peace, pax. Colombia has had a long history of violence and since the peace agreement in 2016, much effort has been put into making this country a peaceful land, including the Truth Commission (Comisión de la Verdad). Thanks to all the efforts for peace, we do believe that the change is inevitable. The name is masculine.
This new genus is placed within the Cacodemoniini as it has the triangular elongation made by the fusion of the dorsal and the lateral apodemes in the male genitalia that are characteristic of that tribe (
Holotype. Colombia • ♂; Meta, San Martín, Vda. San Francisco, Hacienda La María; [3°39'55.5"N, 73°39'29.7"W]; 400 m; on bat guano;
Paratype. Colombia • 1 ♂; same data as for the holotype;
Males of Paciwithius chimbilacus sp. nov. can be distinguished from P. valduparensis sp. nov. by a slender pedipalpal femur and patella (femur: 4.10–4.35× longer than broad in P. chimbilacus and 4.00 in P. valduparensis; patella: 3.33–3.43× longer than broad in P. chimbilacus and 2.27× in P. valduparensis), and a stouter chela (3.16–3.33× longer than broad in P. chimbilacus and 3.52× in P. valduparensis).
Adults. Color: carapace brownish, darker than the tergites. Tergites yellow-brown, heavily granulated. Legs yellowish, proximal segments darker than the distal ones. Pedipalps reddish brown, heavily granulated; chela and fingers reddish (Fig.
Dimensions (mm) (L/W): male: holotype: body length 1.93. Pedipalps: trochanter 0.34/0.20, femur 0.70/0.16, patella 0.63/0.18, chela (with pedicel) 0.94/0.26, chela (without pedicel) 0.88, hand (without pedicel) length 0.52, movable finger length 0.44. Chelicera 0.19, movable finger length 0.14. Carapace 0.66/0.52 (width at medial area). Leg I: femur 0.13/0.13, patella 0.27/0.11, tibia 0.29/0.08, tarsus 0.28/0.06. Leg IV: femur + patella 0.55/0.16, tibia 0.49/0.10, tarsus 0.37/0.06. Male: paratype: body length 1.78. Pedipalps: trochanter 0.36/0.19, femur 0.69/0.17, patella 0.64/0.19, chela (with pedicel) 0.92/0.27, chela (without pedicel) 0.86, hand (without pedicel) length 0.46, movable finger length 0.42.
Carapace
(Fig.
Paciwithius chimbilacus sp. nov. (male holotype,
Chelicera
(Fig.
Pedipalp
(Fig.
Coxal region : coxal chaetotaxy: 20: 23: 24: 35; maxilla with 38 setae including two apical setae and one very small internal, sub-oral setae; median maxillary lyrifissure medial in position, posterior lyrifissure not visible.
Legs
(Fig.
Abdomen
(Fig.
Genitalia
: see
This species epithet is derived from the common name of bats in the Meta region “chimbilaco”, due to the bat guano where the specimens were found.
Holotype. Colombia • ♂; Cesar, Valledupar, Ecoparque Los Besotes, campamento base; [10°34'30.0"N, 73°16'19.8"W]; 600 m; 17 Jul. 2015; CarbioTeam leg.; manual capture;
Males of Paciwithius valduparensis sp. nov. can be separated from P. chimbilacus sp. nov. by their stouter pedipalpal femur and patella (femur: 4.10–4.35× longer than broad in P. chimbilacus and 4.00× in P. valduparensis; patella: 3.33–3.43× longer than broad in P. chimbilacus and 2.27× in P. valduparensis), and a slender chela (3.16–3.33× longer than broad in P. chimbilacus and 3.52× in P. valduparensis).
Adults. Color: yellowish brown, darker in carapace and tergites; carapace metazone without paired pale spots; pedipalps brownish, somewhat paler than body, fingers reddish; legs yellow-brown, uniform color; all specimen heavily granulated.
Dimensions (mm): male: holotype: body length 1.84. Pedipalps: trochanter 0.28/0.18, femur 0.61/0.15, patella 0.60/0.26, chela (with pedicel) 0.94/0.26, chela (without pedicel) 0.90, hand (without pedicel) length 0.44, movable finger length 0.47. Chelicera 0.21, movable finger length 0.15. Carapace 0.66/0.52 (width at medial area). Leg I: femur 0.14/0.11, patella 0.24/0.10, tibia 0.27/0.07, tarsus 0.29/0.05. Leg IV: femur + patella 0.50/0.14, tibia 0.40/0.09, tarsus 0.35/0.05.
Carapace
(Fig.
Paciwithius valduparensis sp. nov. (male holotype,
Chelicera
(Fig.
Pedipalp
(Fig.
Coxal region : coxal chaetotaxy: 10: 14: 15: 19; maxilla with 32 setae including two apical setae and one very small internal, sub-oral seta; median maxillary lyrifissure medial in position, posterior lyrifissure not visible.
Legs
(Fig.
Abdomen
(Fig.
Genitalia
: see
This species is named after the city in which it was found. The Valledupar demonym is valduparensis. This is considered the place where the vallenato music was born. The specific epithet is an adjective.
Cystowithius
Cystowithius smithersi Harvey, 2004, by original designation.
See
The genus Cystowithius is endemic to Central and South America (
Holotype. Colombia • ♂; Tolima, Juntas, Reserva Natural Ibanasca; [4°33'22.0"N, 75°19'17.2"W]; 1700 m; 12 Feb. 2007; C. Cortes leg.; Pinus plantation; on low vegetation, manual capture;
Cystowithius florezi can be separated from the other Cystowithius species as follows. Cystowithius florezi sp. nov. is very similar to C. colombicus Harvey, 2004 and C. ecuadoricus Harvey, 2004, so we provide a detailed comparison for each one in Table
Comparative characters between three species of Cystowithius. Bold font denotes ratios.
Species/ Characters | C. colombicus Harvey, 2004 | C. ecuadoricus Harvey, 2004 | C. florezi sp. nov. |
---|---|---|---|
Body length | 2.00–2.11 | 2.16–2.29 | 1.94 |
Carapace length/width | 0.74/0.52 | 0.75–0.80/0.62 | 0.67/0.51 |
1.43 | 1.20–1.28 | 1.31 | |
Pedipalpal femur length/width | 0.87–0.97/0.17–0.20 | 0.80–0.82/ 0.17–0.18 | 0.60/0.17 |
4.78–5.25 | 4.47–4.74 | 3.60 | |
Pedipalp patella length/width | 0.77–0.88/0.19–0.21 | 0.67–0.69/0.20–0.21 | 0.61/0.18 |
3.88–4.46 | 3.18–3.47 | 3.30 | |
Chela with pedicel length/width | 1.12–1.26/0.28–0.33 | 1.09–1.14/0.28–0.30 | 1.02/0.28 |
3.80–4.04 | 3.65–4.01 | 3.64 | |
Movable finger ratio (length/hand width) | 0.49–0.61 | 0.50–0.56 | 0.52 |
0.90–1.24 | 0.86–1.11 | ||
Hand length/width | 0.50–0.60 | 0.50–0.58 | 0.49 |
Femur/patella IV | 0.67/0.17 | 0.60/0.21 | 0.57/0.18 |
2.93 | |||
Glandular setae | sternites VII–IX with patches of gls arranged ca. 20: 27: 13 respectively | sternites VI–IX with patches of gls, arranged 6: 42: 10: 8 (lectotype) respectively | sternite VIII with patches of gls, arranged 40 in a small circle |
Sternal pockets on anterior margins of sternites | VI–VII | V–VIII | VI–VIII |
Sense spots on chelal fingers | without sense spots on movable finger | without sense spots on movable finger but with small sense spot slightly distal to st that contains three small nubbins | without sense spots on movable finger but with small spot slightly distal to st that contains two small nubbins |
Color
: with sclerotized portions, generally yellow-brown; carapace and pedipalps darker; carapace metazone with paired pale spots; legs darker at the edges (Fig.
Dimensions (mm): male: holotype: body length 1.94. Pedipalps: trochanter 0.36/0.18, femur 0.60/0.17, patella 0.61/0.18, chela (with pedicel) 1.02/0.28, chela (without pedicel) 0.96, hand (without pedicel) length 0.49, movable finger length 0.52. Chelicera 0.19, movable finger length 0.14. Carapace 0.67/0.51 (width at medial area); eye diameter 0.06. Leg I: femur 0.14/0.14, patella 0.29/0.14, tibia 0.20/0.10, tarsus 0.25/0.06. Leg IV: femur + patella 0.57/0.18, tibia 0.43/0.18, tarsus 0.34/0.07, TS 0.74.
Carapace
(Fig.
Cystowithius florezi sp. nov. (male holotype,
Chelicera
(Fig.
Pedipalp
(Fig.
Coxal region : coxal chaetotaxy: 8: 8: 8: 12, with multiple small lyrifissures; maxilla with two apical setae, one very small internal, sub-oral seta and 12 setae; median maxillary lyrifissure medial in position, posterior lyrifissure present.
Legs
(Fig.
Abdomen
(Fig.
Genitalia
: see
This species is dedicated to Professor Eduardo Florez, considered the father of Arachnology in Colombia. He has also been the curator of the Arachnological collection of the Instituto de Ciencias Naturales, where all the material used in this study is lodged.
Cystowithius ankeri Garcia & Romero-Ortiz, 2021: 2–5, figs 1–12, 15.
Colombia • 3 ♂, 3 ♀; Cundinamarca, San Antonio del Tequendama, Parque Natural Chicaque; 04°36'51.3"N, 74°18'55.2"W; 2600 m; 1 Jun. 2009; F. Helbig leg.; on canopy log at 25 m height, manual capture;
Although these specimens were collected 320 km from the type locality in Caldas Department, we refer them to the species C. ankeri. They have similar pedipalp proportions; however, their patches of glandular setae are different. The holotype of C. ankeri has two small patches in sternite VIII, and those examined here have one long patch of setae (Fig.
1 | Chelal hand smooth; setae on chelal hand only barely denticulate; tactile setae of tarsus IV situated closer to its mid-length (TS = 0.59–0.61 mm) | C. chamberlini Harvey, 2004 |
– | Chelal hand evenly granulate; setae on chelal hand distinctly denticulate; tactile seta of tarsus IV situated subdistally (TS = 0.68–0.79 mm) | 2 |
2 | Movable chelal finger > 0.70 mm long; males with sternal invaginations on sternites VI–VIII | C. ankeri García & Romero-Ortiz, 2021 |
– | Movable chelal finger < 0.60 mm long; males with sternal invaginations on sternites V–VIII or VI–VII | 3 |
3 | Pedipalps longer and more slender, i.e., chela (with pedicel) longer than 1.35 mm | C. smithersi Harvey, 2004 |
– | Pedipalps shorter and more robust, i.e., chela (with pedicel) shorter than 1.20 mm | 4 |
4 | Femur of the pedipalp stout (3.96 times longer than broad), males with sternal invaginations on anterior margins of sternites VI–VIII, patches of glandular setae on sternite VIII in a small circle | C. florezi sp. nov. |
– | Femur of the pedipalp slender (> 4.47 times longer than broad), males with sternal invaginations on anterior margins of sternites VI–VIII, patches of glandular setae on sternite VIII in a small circle | 5 |
5 | Setae on tergite XI short and strongly clavate; chelal hand without long, strongly denticulate setae | C. colombicus Harvey, 2004 |
– | Setae on tergite XI long and only slightly clavate; chelal hand with long, strongly denticulate setae | C. ecuadoricus (Beier, 1959) |
Parawithius
Chamberlin, 1931a: 292;
Chelifer nobilis With, 1908, by original designation.
See
Holotype. Colombia • ♂; Cundinamarca, Cogua, Embalse del Neusa Tausa, Llano Grande; [5°11'33.9"N, 73°53'53.8"W]; 2900 m; 7 Mar. 2004; AL Leon leg.; under tree bark;
Paratypes. Colombia • 1 ♂; Cundinamarca, same data as for the holotype. • 1 ♀, 1 ♂; Sesquilé, Camino al Cerro de Las Tres Viejas; 5°02'17.0"N, 73°47'13.0"W; 2740 m; 8 Sep. 2019; C. Romero-Ortiz, F. Garcia, J.J. Lagos, A. Carvajal, D. Mayorga-Ch leg.; on bromeliad;
Other material. Colombia • 1 ♀; Santander, Málaga, Vda. Buenavista, km 7 vía Bucaramanga; 6°42'23.7"N, 72°44'52.6"W; 2620 m; 1 Jan. 2020; C. Romero-Ortiz, J.J. Lagos leg.; on bromeliad under “Loqueto” tree Escallonia pendula (Ruiz & Pav.) Pers.;
Parawithius bromelicola sp. nov. can be separated from P. nobilis (With, 1908) by the stouter pedipalpal segments (i.e. patella 3.36–3.44× longer than broad compared with 3.24–3.30× longer than broad, and the chela without pedicel 3.39–3.47× longer than broad, compared to 3.58–3.85× longer than broad); the extension of the strongly clavate setae on the dorsal surface of fixed chelal finger (i.e., distal to it and est compared to proximal to it and est); from P. iunctus Beier, 1932 and P. pseudorufus Beier, 1932 by the presence of pale spots on the carapace metazone, in addition to the size of chelal fingers compared to the palpal hand (i.e. fingers shorter to the hand in P. bromelicola sp. nov., P. nobilis, and P. iunctus and longer in P. pseudorufus).
Adults. Color: yellowish brown, carapace darker than body, carapace metazone with paired pale spots; pedipalps reddish brown, uniform in color, very granulated; tergites yellow-brown; big leg segments darker at posterior margin (Fig.
Dimensions (mm): male: holotype (followed by male paratypes): body length 2.46 (2.58, 2.46). Pedipalps: trochanter 0.38/0.21 (0.37/0.23, 0.38/0.22), femur 0.83/0.19 (0.84/0.19, 0.77/0.18), patella 0.74/0.22 (0.75/0.22, 0.71/0.21), chela (with pedicel) 1.18/0.30 (1.24/0.31, 1.12/0.30), chela (without pedicel) 1.14 (1.18, 1.06), hand (without pedicel) length 0.59 (0.57, 0.53), movable finger length 0.56 (0.60, 0.53). Chelicera 0.21, movable finger length 0.17. Carapace 0.82/0.58 (0.85/0.72, 0.82/0.74); eyespot diameter 0.07. Leg I: femur 0.16/0.16, patella 0.33/0.14, tibia 0.35/0.10, tarsus 0.32/0.06. Leg IV: femur + patella 0.66/0.19, tibia 0.51/0.12, tarsus 0.37/0.07, TS 0.73.
Female: paratype (followed by female other material): body length 2.68 (2.70). Pedipalps: trochanter 0.41/0.23 (0.36/0.21), femur 0.78/0.21 (0.73/0.18), patella 0.74/0.23 (0.67/0.19), chela (with pedicel) 1.24/0.35 (1.14/0.30), chela (without pedicel) 1.16 (1.08), hand (without pedicel) length 0.59 (0.57), movable finger length 0.60 (0.49). Carapace 0.88/0.68 (0.74/0.64) (width at medial area).
Carapace
(Fig.
Parawithius bromelicola sp. Nov. (male holotype,
Oligowithius achagua sp. nov. (male holotype,
Chelicera
(Fig.
Pedipalp
(Fig.
Coxal region : coxal chaetotaxy: ♂, 10: 10: 5: 7; maxilla with 23 setae including three apical setae (one tactile setae) and one very small internal, sub-oral seta; median maxillary lyrifissure medial in position, posterior lyrifissure present.
Legs
(Fig.
Abdomen
: all tergites divided with a medial suture, with a row of spots in the middle of each hemitergite; sternites entire. Tergal chaetotaxy: ♂, 8: 9: 9: 10: 13: 13: 16: 16: 18: 12: 10 (including two tactile setae): 2; all setae foliate. Sternal chaetotaxy: ♂, 15: (2) 11 (2): (2) 13 (2): 16: 14: 16 + 21 gls: 13 + 86 gls: 11 + 2 gls: 10 (including two tactile setae): 8 (including two tactile setae and some dentate): 2; sternites VII and VIII of ♂ with patches of glandular setae (Fig.
Glandular setae patches of the new species A Paciwithius chimbilacus sp. nov. (holotype,
Genitalia
: see
This species is named after bromeliad plants, due to its close association with them. Most of the specimens were collected by sifting bromeliads on white sheets.
Dolichowithius (Oligowithius) Beier, 1936: 447.
Dolichowithius (Oligowithius) abnormis Beier, 1936, by monotypy.
The genus Oligowithius can be distinguished from other Neotropical withiid genera by the presence of a patch of glandular setae on sternites VII and VIII either in one or two small circular patches in the middle of the sternites, the distal position of the tactile setae on tarsus IV, the presence of two non-corneate eyes, and the trichobothria it and isb located far apart. Moreover, the male genitalia are characterized by a pair of lateral apodemes that are not merged with the dorsal apodemes (Fig.
Dolichowithius (Oligowithius) abnormis Beier, 1936: 446–447, fig. 4.
With the recognition of Oligowithius as a distinct genus, D. (O.) abnormis is here transferred to the genus Oligowithius.
Holotype. Colombia • ♂; Meta, Puerto Gaitán, Carimagua; 160 m; 22 Apr. 2012; D. Martinez leg.; estuary on the eastern plains;
Paratypes. Colombia • 1 ♂ and 1 ♀; Meta, same data as the holotype.
Oligowithius achagua sp. nov. differs from O. abnormis by its smaller size (1.7 mm vs 2.18 mm in O. achagua), more slender patella in O. achagua (3.05× longer than broad) than in O. abnormis (2.74×), and a stouter chela in O. abnormis (3.70× longer than broad) than in O. achagua (4.05× longer than broad).
Adults. Color: yellowish brown; carapace and pedipalp reddish brown; legs yellowish, paler than the abdomen, darker in the edges; carapace metazone without paired pale spots (Fig.
Dimensions (mm): male: holotype (followed by male paratype): body length 2.18 (2.10). Pedipalps: trochanter 0.33/0.16 (0.27/0.15), femur 0.58/0.15 (0.54/0.15), patella 0.54/0.18 (0.50/0.16), chela (with pedicel) 0.94/0.23 (0.88/0.22), chela (without pedicel) 0.88 (0.84), hand (without pedicel) length 0.48 (0.43), movable finger length 0.45 (0.43). Chelicera 0.19, movable finger length 0.16. Carapace 0.74/0.58 (0.68/0.56) (width at medial area); eyespots diameter 0.08. Leg I: femur 0.14/0.14, patella 0.26/0.14, tibia 0.26/0.08, tarsus 0.26/0.06. Leg IV: femur + patella 0.47/0.16, tibia 0.34/0.11, tarsus 0.33/0.06, TS 0.76.
Female: paratype: body length 2.36. Pedipalps: trochanter 0.28/0.16, femur 0.54/0.15, patella 0.52/0.18, chela (with pedicel) 0.94/0.25, chela (without pedicel) 0.88, hand (without pedicel) length 0.48, movable finger length 0.44. Carapace 0.73/0.58.
Carapace
(Fig.
Chelicera
(Fig.
Pedipalp
(Fig.
Coxal region : coxal chaetotaxy: ♂, 6: 7: 9: 18; maxilla with 19 setae including two apical setae and one very small internal, sub-oral seta; median maxillary lyrifissure medial-anterior in position, posterior lyrifissure present.
Legs
(Fig.
Abdomen
(Fig.
Genitalia
: simple structure with most of the components extremely reduced to chitinized lateral apodemes that do not merge with the dorsal apodemes. The level of sclerotization of the ejaculatory channel is weak; however, it is projected from the lateral apodemes, which allows us to classify Oligowithius as a Cacodemoniini (Fig.
This species is named after the indigenous people, original inhabitants of the Meta, Vichada and the Venezuelan Llanos, the Achaguas. The name should be treated as a noun in apposition.
In Withiidae, the presence of patches of abdominal glandular setae, especially in males, have been used as a diagnostic character since
Male genitalia need to be addressed when defining withiid genera. As shown by
Finally, we are aware that a fully resolved phylogeny is needed to support the new clades and the assessment of the old ones; however, as for the new genus Paciwithius and for the new genus rank of Oligowithius, we presented a unique and exclusive combination of morphological characters that allow us to conclude and support the taxonomic changes presented in this work.
We are grateful for the fellowship “Doctorados Nacionales 727-2015” of Minciencias which supported the PhD of CRO. We thank to Eduardo Flórez, curator of the Arachnological Collection of the Instituto de Ciencias Naturales of the Universidad Nacional de Colombia. We also thank Edgar Linares, professor of the same institution, for loan of the necessary equipment for descriptions and drawings. Fabian García kindly made the distribution map for which we are very grateful. Finally, CRO is indebted to Juan José Lagos Oviedo for his continuous support with field trips, time management, and personal balance, including many other significant learnings.
The authors have declared that no competing interests exist.
No ethical statement was reported.
Call 727-2015 Doctorados Nacionales. Minciencias, Colombia.
CRO planned the study, studied the specimens, made the descriptions, draw the figures, and wrote and review the first draft. MSH examined the specimens, took SEM pictures, and reviewed the manuscript. CES helped to plan the study, wrote and review the manuscript.
Catalina Romero-Ortiz https://orcid.org/0000-0001-8939-7814
Carlos E. Sarmiento https://orcid.org/0000-0003-4012-8108
Mark S. Harvey https://orcid.org/0000-0003-1482-0109
All of the data that support the findings of this study are available in the main text.