Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

A new type of nest in the genus Quindina (Opiliones: Nomoclastidae)

with hypotheses about its origin and a new species for Colombia

Héctor Lancheros*1; https://orcid.org/0000-0002-0632-5151

Daniel Arias Cuellar1; https://orcid.org/0000-0001-8486-6319

Ricardo Pinto-da-Rocha2; https://orcid.org/0000-0002-3959-2205

1. Programa de Biología, Universidad El Bosque, Bogotá, Colombia; lancheroshector@unbosque.edu.co

(*Correspondence); dsarias@unbosque.edu.co

2. Departamento de Zoologia/Instituto de Biociências, Universidade de São Paulo, São Paulo, Brazil; ricrocha@usp.br

Received 06-VIII-2024. Corrected 24-IX-2024. Accepted 27-IX-2024.

ABSTRACT

Introduction: Parental care is a phenomenon that has been recorded for several clades of the order Opiliones,

but nest building has only been reported in the Neotropical genus Quindina Roewer, 1914 (Nomoclastidae).

Objective: To describe a new type of nest used in parental care by a new species of Quindina and determine its

placement in the phylogenetic tree of the genus, to propose hypotheses about the origin of the two types of nests

reported so far.

Methods: Specimens and nests were collected in La Calera, Cundinamarca, Colombia. Observations of parental

care were made both in the field and in terrariums. A new species was described based on somatic characters

and penis structure. A phylogenetic analysis was carried out, consensus methods with equal weights, implicit

weighting, and successive weighting were used.

Results: We describe a new case of parental care with a new type of disc-shaped nest, which we call “pendular

disc nest”. The nest is built by males of Quindina pendula sp. nov.; this species can be recognized externally by

the absence of white spots, the presence of a mushroom-shaped set of patches in the middle part of the prosoma,

and the presence of a dark yellow background around most of the tubercles. Results of the phylogenetic analy-

sis showed Q. pendula sp. nov. is sister to all Quindina species except Quindina marginata. Based on the tree,

obtained by successive weighting, seven phylogenetic hypotheses, related to the possible origin of the two types

of nests recorded in the genus Quindina, were proposed.

Conclusions: This study presents a new case of egg guarding and the building of nests in arthropods, which is

not very common in nature.

Key words: Nomoclastinae; egg guarding; paternal care; pendular disc nest; open mud nest; Andean Forest.

RESUMEN

Un nuevo tipo de nido en el género Quindina (Opiliones: Nomoclastidae)

con hipótesis sobre su origen y una nueva especie para Colombia

Introducción: El cuidado parental es un fenómeno que se ha registrado para varios clados del orden

Opiliones, pero la construcción de nidos solo se ha reportado en el género neotropical Quindina Roewer, 1914

(Nomoclastidae).

Objetivo: Describir un nuevo tipo de nido utilizado en el cuidado parental, por una nueva especie de Quindina,

y determinar su ubicación en el árbol filogenético del género, con el fin de proponer hipótesis sobre el origen de

los dos tipos de nidos reportados hasta el momento.

https://doi.org/10.15517/rev.biol.trop..v72i1.61426

INVERTEBRATE BIOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

INTRODUCTION

Nomoclastidae Roewer, 1943, is a family of

Neotropical species of harvestmen distributed

from Costa Rica to Northern Brazil, it includes

the subfamilies Zamorinae Kury, 1997 and

Nomoclastinae Roewer, 1943, the first includ-

ing the monotypic genus Zamora Roewer, 1927

and the second comprising the genera Call-

cosma Roewer, 1932, Globitarsus Roewer, 1913,

Kichua Pinto & Bragagnolo, 2015, Liomma

Roewer, 1959, Lisarea Roewer, 1943, Merida-

natus Roewer, 1943, Micropachylus Roewer,

1913, Napostygnus Roewer, 1929, Nomoclastes

Soerensen, 1932, Quindina Roewer, 1914 and

Troya Roewer, 1914 (Kury et al., 2024; Kury &

Villarreal, 2015; Pinto-da-Rocha & Bragagnolo,

2017; Villarreal & Kury, 2023). The family is

defined by the following characters: no sexual

dimorphism in its chelicerae and pedipalps

(except Liomma); low, elliptical ocularium, with

a median depression; ventral plate of the penis

sub-rectangular to trapezoidal in shape; no

lamellar setae on the ventrodistal margin of

the ventral plate; short, stout stylus with a

rounded head, mounted on a long, columnar

glans; glans without processes except in Napos-

tygnus (Pinzón-Morales & Pinto-da-Rocha,

2020). Nomoclastidae currently includes 12

genera (8 monotypic) and 29 species, includ-

ing the one herein described (Kury et al., 2024;

Pinto-da-Rocha & Bragagnolo, 2017; Pinzón-

Morales & Pinto-da-Rocha, 2020; Villarreal &

Kury, 2023).

The genus Quindina: Quindina is charac-

terized by a type of body armor similar to that

of Callcosma and Kichua, with a pair of long

spines on area III and free tergites with a pair

of spines each. It differs from other genera of

the family by most species having a row of tall

tubercles separated from each other on the lat-

eral margins of dorsal scutum, very elongated

tubercles on area III of doral scutum, absence of

lateral margin elevation, and the presence of an

apical prodorsal projection on femur IV (Pinto-

da-Rocha & Bragagnolo, 2017; Pinzón-Morales

& Pinto-da-Rocha, 2020). It is distributed in

Costa Rica, Panama, Colombia and Ecuador.

Eleven species had previously been described

in this genus, four of which from Colombia: Q.

bella Roewer, 1914 (type species) and Q. mar-

ginata (Roewer, 1963), from the Andean region;

Q. discolor Pinzón-M. & Pinto-da-Rocha, 2020

and Q. hermesi Pinzón-M. & Pinto-da-Rocha,

2020, from the Caribbean region (Pinto-da-

Rocha & Bragagnolo, 2017; Pinzón-Morales &

Pinto-da-Rocha, 2020).

Parental care and nest building: Exclu-

sive paternal care evolved independently in

fifteen arthropod lineages, which include

Métodos: Los especímenes y nidos fueron recolectados en La Calera, Cundinamarca, Colombia. Se observó el

cuidado parental, en el campo y en terrarios, de una nueva especie descrita con base en caracteres somáticos y

en la estructura del pene. Se realizó un análisis filogenético, se utilizaron métodos de consenso con pesos iguales,

ponderación implícita y ponderación sucesiva.

Resultados: Describimos un nuevo caso de cuidado parental con un nuevo tipo de nido, denominado “nido disco

pendular”, el cual es construido por machos de Quindina pendula sp. nov.; esta especie se reconoce externamente

por la ausencia de manchas blancas, la presencia de un conjunto de manchas en forma de champiñón en la parte

media del prosoma y la presencia de un fondo amarillo alrededor de los tubérculos. Los resultados del análisis

filogenético mostraron a Q. pendula sp. nov. como hermana de todas las especies de Quindina, excepto Quindina

marginata. Con base en el árbol obtenido, por ponderación sucesiva, se propusieron siete hipótesis filogenéticas,

relacionadas con el origen de los dos tipos de nidos registrados.

Conclusiones: Este estudio presenta un nuevo caso de protección de los huevos para el género Quindina, que

implica la construcción de nidos en artrópodos, lo cual es poco común en la naturaleza.

Palabras clave: Nomoclastinae; protección de huevos; cuidado paterno; nido disco pendular, nido abierto de

barro; bosque andino.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

approximately 1 500 species. This is a very

small fraction of the diversity of this group

(with more than 1.2 million species), making it

probably the rarest form of postzygotic paren-

tal care investment (Requena et al., 2014). Egg

guarding behavior is observed in most orders of

the class Arachnida, but paternal investment in

postzygotic care is restricted to members of the

order Opiliones. For example, in Gonyleptidae,

adult males of Ampheres leucopheus (Mello-

Leitão, 1922) protect batches of eggs, attached

by a mucous substance to the underside of

leaves, and newly hatched nymphs (Hara et al.,

2003). Three species of Progonyleptoidellinae

Soares & Soares, 1985 show a similar behavior,

with eggs being laid on the underside of leaves,

covered by a thick mucous layer, while two spe-

cies of Gonyleptinae Sundevall, 1833 lay eggs

in natural cavities, in trunks or on the ground.

In all these cases, eggs can be found in differ-

ent stages of development with males taking

care of them (Machado et al., 2004). Maternal

care is also present in the order Opiliones and

has been reported for at least five families of

the suborder Laniatores Thorell, 1876. There

are cases of maternal care in which females

care for eggs in cavities and leaves or for

example, two species of Cranaidae, belonging

to the genus Phareicranaus Roewer, 1913, have

females that protect groups of eggs in cavities

found between the roots of trees. The same

type of behavior has been observed in females

of the Gonyleptidae subfamily Goniosomatinae

(Machado & Warfel, 2006).

The aforementioned cases imply paternal

or maternal care of the eggs in surfaces or cavi-

ties, but without the construction of complex

structures to protect the eggs. Rodriguez and

Guerrero (1976) provided the first description

of harvestmen nests for Quindina albomarginis

(Chamberlin, 1925). The authors indicated that

the construction of each nest corresponded to

a single male, and that the material used came

from the outer bark or outer bark remains of

the trunk on which it was built, or from logs

and debris from the vicinity. Mora (1990) com-

plemented these observations and indicated

that, for the construction, the males use bark

removed with the chelicerae and mixed with

mud collected from the crevices of the trunk.

She also noted that material could be collected

from the forest floor, combined with salivary

secretions and later applied on the substrate

to assemble the nest, with a mud wall then

applied to the edges, with an average nest diam-

eter of 33 mm and an average nest wall height

of 8.3 mm. Pinto-da-Rocha and Bragagnolo

(2017) described four species of Quindina from

Panama, three with the same type of nest (Q.

morae Pinto-da-Rocha & Bragagnolo, 2017, Q.

burbayar Pinto-da-Rocha & Bragagnolo, 2017

and Q. kuna Pinto-da-Rocha & Bragagnolo,

2017), which they called “open mud nest” or

“reproductive arena”. Quesada-Hildalgo et al.

(2019) and Rojas et al. (2019) also reported this

type of nest for Q. limbata (Roewer, 1943) in

Costa Rica, calling it “mud-nest” or “cup-like

m u d ne s t ”.

Our aim in this study is to describe a new

species of the genus Quindina, which builds

a new type of nest. In addition, we strive to

determine the position of the species in the

phylogenetic tree of its respective genus and

hypothesize about the origins for the two nest

types recorded in this group.

MATERIALS AND METHODS

Collection and conservation of speci-

mens: We collected the male specimens in the

vereda Mundo Nuevo and females in the vereda

El Líbano, in the municipality of La Calera,

Cundinamarca, Colombia. The specimens were

gathered at the site where the nests were found,

by direct observation of the tree branches, the

underside of trunks and the interior of an aban-

doned hut. After locating the nests, we waited

until nightfall to collect the male individuals

coming back to the nests. Two male individu-

als were fixed in 70 % ethanol; two others were

transported alive, in order to observe their nest

building in terrariums. Females were collected

under logs and rocks near a road in the vereda

El Líbano. The two female specimens collected

were fixed in 70 % ethanol. All specimens were

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

deposited in the collection of the “Museo de

Ciencias de la Universidad El Bosque (MCUB)”.

Measurement and description of the

specimens: We observed the general characters

of the body and appendages under a stereo-

scope, with magnifications of 10-180X. The

penis was observed under an optical micro-

scope, with magnifications of 400X and 1 000X.

In both cases, photographs were taken with

a digital camera attached to one of the eye-

pieces. In the photographs taken from the

stereoscope, a millimeter scale was included. In

the microscope photographs, dimensions were

calculated using the millimeter scale recorded

at 4X magnification. These scales were used

to measure the parts with ImageJ software

(Rueden et al., 2017).

All measurements are given in millimeters.

Illustrations were made with LibreOffice Draw

(The Document Foundation, 2022) and Gimp

software (Kimball et al., 2022). The delinea-

tion of the parts was drawn following the con-

tour of the digital photographs. Descriptions

and terminology follow the ones proposed by

Pinto-da-Rocha and Bragagnolo (2017) and

Pinzón-Morales and Pinto-da-Rocha (2020).

Description of genitalia follows the findings of

Kury and Villarreal (2015) and Kury (2016).

Indications for nomenclature of the structures,

measurements and relative positions of struc-

tures, follow Acosta et al. (2007).

Distribution records: At the type local-

ity (vereda Mundo Nuevo, municipality of La

Calera) a photographic record of the vegetation

type was taken using a drone. The collection

records of the type specimens was comple-

mented by previous observations made by Héc-

tor Lancheros in the rural area of the Bogotá,

Capital District, and other new records made in

the municipalities of Choachí (Department of

Cundinamarca), Saboyá (Department of Boy-

acá) and Une (Department of Cundinamarca),

by biologists Daniel Arias Cuellar, Christian

Franco García and Osvaldo Villarreal, respec-

tively. The distribution map was assembled

using QGIS software (QGIS.org, 2021), based

on the coordinates recorded with Viking soft-

ware (Battaglia et al., 2021).

Description of the nests: The fresh weight

of twelve nests was recorded in the field, using

an electronic scale with 3-digit precision.

Length and width dimensions for each one were

measured using ImageJ software (Rueden et al.

2017), based on photographs taken on the scale

plate, using its diameter as a point of reference.

Thickness was recorded using a digital caliper.

The dry weight of eight nests was recorded after

drying them in a hot air circulation oven at 30

°C. Four nests were used for behavioral obser-

vations in the laboratory, where we paid special

attention to the care provided by adult males

and egg development and hatching. These four

nests were also used to analyze the materials

that compose them. In each nest, the type of

plant material from which it hung was identi-

fied. The remains of liverworts present in nests

were identified down to genus level.

Observations of parental care and devel-

opment: The first observations of parental

care were made directly in the field, from the

beginning of the day until the early evening

hours (from 18:00 to 20:30). For observations

under controlled conditions in the laboratory,

two terrariums were prepared, each containing

a section of branch from which hung a nest in

a plastic box, with the respective adult male.

Observations were made in each terrarium

for two weeks. For developmental observa-

tions (i.e. changes in egg coloration, hatching

and nymph development), two complementary

set-ups were made on absorbent paper in Petri

dishes, each with only one nest. Observations

for each petri dish lasted between 20 days and

one month, until hatching ceased. Adults and

nymphs were fed using cooked rice with peas

and carrots, cooked spaghetti and dead fruit

flies, in both terrariums and petri dishes.

Phylogenetic analysis: The matrix used

for the phylogenetic analysis was originally

published by Kury and Villarreal (2015), includ-

ing 77 morphological characters and 38 taxa

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

(26 from the clade Laminata). It was later

modified by Pinto-da-Rocha and Bragagnolo

(2017) who added 13 species in Nomoclastidae,

adjusted the coding of characters 11 and 12,

and added 16 new characters (78 to 93), two of

which originated from the subdivision of the

original character 13 (characters 78 and 82).

To this, Pinzón-Morales and Pinto-da-Rocha

(2020) added two new species of the genus

Quindina. With the addition of the new species,

the matrix now comprises 93 characters and

54 taxa (Table 1), all belonging to the suborder

Laniatores. The taxa comprise 48 species of the

superfamily Gonyleptoidea (infraorder Grassa-

tores), of which 42 belong to the clade Laminata,

which is divided into 20 species of Nomoclasti-

dae and 22 of Microsetata. Outgroups comprise

five species from three other superfamilies of

the infraorder Grassatores (Samooidea, Eped-

anoidea and Assamioidea) and the root is one

species from the superfamily Triaenonychoidea

of the infraorder Insidiatores (the latter is the

first outgroup) (Kury & Villarreal, 2015). The

description of the characters can be found in

Pinzón-Morales and Pinto-da-Rocha (2020).

Species of the most recently added genera to

Nomoclastidae (Globitarsus, Liomma, Lisarea,

Meridanatus, Micropachylus and Troya) were

not included due to lack of reproductive data

and several morphological data for matrix.

The parsimony analysis was carried out using

the software TNT v.1.5 (Goloboff & Cata-

lano, 2016). The heuristic method (“traditional

search”) was applied with the tree bisection-

reconnection (TBR) option with equal weights.

From the trees with maximum parsimony, two

cladograms were obtained, one using the strict

Nielsen consensus method and another using

majority consensus, with 50 % cutoff. Subse-

quently, the implicit weighting method was

applied, which decreases the weight value of

homoplastic characters in the analysis based on

a concavity constant (k), the value of this con-

stant is negatively related to the decrease in the

weight of these characters in the analysis (Golo-

boff, 1993). To obtain the “optimal” value of the

constant k for our data set, the TNT script setk.

run was used (Goloboff et al., 2008; Villarreal &

García, 2021). Finally, the successive weighting

method was applied with the rescaled consis-

tency index rc (Farris, 1989), using the script

rewt.run. The latter is a technique that decreas-

es the weight of homoplastic characters after

obtaining a cladogram and repeats the pro-

cess until successively equal trees are obtained

(Farris, 1969). Relative Bremer supports were

calculated for the trees obtained with the four

methods, which provide an approximate mea-

sure of the amount of favorable/contradic-

tory evidence for each group (Giribet, 2003;

Goloboff & Farris, 2001), and the tree with the

highest support values was chosen. Sensitivity

plots (Navajo rugs) were used to compare the

coincidence of the groups obtained with the

different methods and hypotheses.

Hypotheses of the origin of the nests: To

propose hypotheses of origin of the nests, the

most parsimonious reconstructions method

was used (Goloboff, 2022, Chapter 3), with the

TNT Characters/Reconstructions option in the

optimize menu. For this, the genus Quindina

section of the chosen tree was used, with an

external group and the character nest presence

with three unordered states: 0 = absent, 1 =

pendular disc nest, 2 = open mud nest, ? = nest

construction unknown. The reconstructions

obtained were stored in the display buffer. The

frequency with which each nest type occurred

in a reference group (each group is one of the

main clades that make up the genus Quindina)

was tabulated for each possible “condition”

(ancestral, derived or in an internal group).

The combination of each nest type with the

reference group was called “characteristic”, and

for each characteristic, the condition with the

highest frequency was listed. This is an illustra-

tive method to indicate the possible nodes of

origin of each nest type, as unknown data for

most species of Quindina does not allow for

making a complete reconstruction of the evolu-

tion of the nest type and its origin.

All figures (illustrations, map, photographs

and cladograms) were diagrammed using

LibreOffice Draw graphics editing software

(The Document Foundation, 2022).

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

Tabl e 1

Data matrix (93 characters, 54 taxa) used for the phylogenetic analysis of Nomoclastidae.

Species

Characters

0000000000 1111111111 2222222222 3333333333 4444444444 5555555555 6666666666 7777777777 8888888888 999

0123456789 0123456789 0123456789 0123456789 0123456789 0123456789 0123456789 0123456789 0123456789 012

Acumontia succinea 0000000000 2-0?000000 0200000200 0000000000 0300-0000- 000000-000 00-0------ -?--000000 0000000000 0??

Stygnomma fuhrmanni 000-100000 2-0?100100 0010000?00 0000000010 1001-0000- 000000-000 00-0------ -?--010000 0000000000 0??

Santobius annulipes 500-100000 2-0?100100 0220000000 0000000010 1001-0000- 000000-000 00-0------ -?--0110?? ?0???????? ???

Pungoica simoni 3101001000 000?001002 01?0100000 0010000012 1001-0000- 0002000000 00-0-00010 00-01100?? ?0???????? ???

Dibunus similis 810-100100 0200100000 1200000100 0010000010 1001-0000- 0002000000 00-0100001 40001110?? ?0???????? ???

Sorensenius pygmaeus 5100000000 000?000000 12?0000000 0010200010 1001-0000- 0002000000 00-0150010 00001110?? ?0???????? ???

Paramitraceras granulatum 5000000000 020?200000 12?0000200 0-10210010 1001-0000- 0002000000 00-0100025 ?00011?000 0000000000 0??

Globibunus rubrofemoratus 1100001000 0001000003 1200000001 0010010110 1102-00000 0201000000 00-01000-- 300011?000 0000000000 000

Ricstygnus quineti 511-101100 020?200000 0020000100 0002111111 1102-00000 0101000000 10-3100014 1301111000 0000000000 0??

Stenostygnellus macrochelis 101-101100 030?200000 0020000100 1002111111 1102-00001 0001000000 00-010001? 1000111000 0000000000 000

Taquara bicoloripes 6100010000 0000001000 0300100000 1010010110 1102-00000 0021000000 00-0100020 110011?000 0000000000 000

Gerdesius mapinguari 6100010000 0301001000 0000000000 1012210111 1102-00000 0001000000 1100100022 ?00111?000 0000000000 000

Cajamarca affinis 2101001100 000?000000 1200000001 1000010110 1203000210 0101010110 10-0040025 141111?000 0000000000 0??

Chacoikeontus clavifemur 3101001100 130?000000 1200000001 1000010110 1203000210 0101010110 10-0040025 141111?000 0000000000 0??

Incasarcus dianae 2101011100 1001000000 1200000201 0010010110 1203000210 0001010110 1131040025 141111?000 0000100000 0??

Taito juruensis 3101002000 2-00011011 0121010401 1012010111 1203000200 0101000210 1122040025 1411111000 0000000000 0??

Cynorta conspersa 3101002100 2-00011011 0121010400 0000000111 1203000200 0101000210 1102040025 1411111000 0000000000 0??

Cosmetus variolosus 3101002200 2-0?011011 0121010400 0000000111 1203000200 0101000210 10-2040025 1411111000 0000000000 0??

Syncranaus cribrum 3101001100 2-00001000 1020000310 1002000111 1203000100 0001010000 1121040025 231111?000 0001100000 0??

Saramacia lucasae 3101001100 2-00001000 1020000310 1002000111 1203000100 0001010000 10-1040025 2311111000 0000000000 0??

Heterocranaus chlorogaster 3011013100 2-00200000 0210001200 1002010111 1213100202 0001111001 1201041125 131111?000 0000000000 0??

Chiriboga albituber 6011001100 2-00200000 0200001200 1002010111 1213100202 0001111001 1201041125 131111?000 0000000000 0??

Zannicranaus monoclonius 4011013100 2-00202000 0220001200 1002010111 1213100202 0001111001 1201041025 131111?000 0000100000 0??

Phareicranaus giganteus 3101003100 1000002000 0300001200 1002010111 1213100202 0001111001 1201041025 ?311111000 0001100000 0??

Phalangodus sp. 2101003100 0000002000 0310000200 1000010111 1213100202 0001011000 1200040025 ?31111?000 0000000000 0??

Licornus tama 4101011110 0311001000 0000000300 1111211111 120300?002 0101210000 ?0?0040025 ?30111?100 0100000000 000

Hutamaia caramaschii 4101011020 031?001000 0000000300 1111211111 1203000002 0001210000 00-0040025 230111?100 0100000000 000

Ampycus telifer 2100011111 2-11001000 0000000300 1112211111 1203000302 1001210000 1210040025 230111?100 0000000000 000

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

Species

Characters

0000000000 1111111111 2222222222 3333333333 4444444444 5555555555 6666666666 7777777777 8888888888 999

0123456789 0123456789 0123456789 0123456789 0123456789 0123456789 0123456789 0123456789 0123456789 012

Nesopachylus monoceros 2101011001 030?001000 0000000300 1112211111 1203000302 1001210000 1210040025 230111?000 0000000000 000

Glysterus metatarsalis 2101011111 030?001000 0000000300 1112211111 1203000302 1001210000 0000040025 230111?000 0000000000 000

Neopachyloides sp. 2101011011 1-1?001000 0000000300 1112211111 1203000302 1001210000 12100?0025 210111?000 0000000000 000

Discocyrtus crenulatus 4201011100 0301001000 1000000303 1110211111 1203000202 0011000000 1101040025 1411111100 0000000000 000

Gonyleptes horridus 4201011100 2-01001000 1000000303 1112-11111 1203000202 0011000000 1101030025 1411111110 0100000010 000

Acanthopachylus aculeatus 4101011000 0101001000 1000000303 1110211111 1203000202 0011000000 1200040025 1411111100 0000000000 000

Zamora granulata 5010001000 0200200000 0200000000 0002110111 120301040- 0001410000 00-0020123 14?111?000 0000000000 000

Nomoclastes quasimodo 50--102000 2-0?300000 0010000000 0000200111 1203011000 0001410000 00-00-0125 020111?000 0000000000 000

Napostygnus bispinosus ?201011011 0000001000 0020000000 0000000112 1203011000 0101410000 0--4030125 ?111?1?000 0000000000 000

Kichua rheimsae 3101001100 2-00001000 1020000000 0000000111 1203011000 0001410000 0--00-0125 020111?000 0000000000 000

Callcosma cofan 3101001100 0300001000 1020000000 0000000111 1203011000 0001410000 0--?0-1125 02-?11?000 0020100000 001

Callcosma gracillima 3101001100 0300001000 1020000000 0000000111 1203011000 0001410000 0--?0-1125 020111?000 0020101000 001

Callcosma abrapatricia 3101001100 1300001000 1000000000 0000000111 1203011000 0001410000 0--40-1125 020111?000 0000111000 001

Callcosma barasana 3101001100 0300001000 1020000000 0000000111 1203011000 0001410000 0--?0?1125 020111?000 0020111000 001

Quindina marginata 3101001100 2-20011000 0000000000 0000000111 1203011000 0001410000 0--?0-0125 ?2??11?200 0100000100 000

Quindina albomarginis 3101001100 1220011000 1000000000 0000000111 1203011000 0001410000 00-40-1125 0201111200 0210100101 011

Quindina albiocularia 3101001100 0320011000 1000000000 0000000111 1203011000 0001410000 0--40-1125 020111?210 0011100111 111

Quindina limbata 3101001100 2-20011000 1000000000 0000000111 1203011000 0001410000 0--40-1125 020111?211 1011100101 111

Quindina burbayar 3101001100 0320011000 1000000000 0000000111 1203011000 0001410000 0--40-1125 020111?200 0011100101 011

Quindina morae 3101001100 0120011000 1000000000 0000000111 1203011000 0001410000 0--40-1125 020111?200 0011100101 011

Quindina bimaculata 3101001100 0-20011000 1000000000 0002000111 1203011000 0001410000 0--?0--125 02??11?201 0100100111 011

Quindina bella 3101001100 0320011000 1000000000 0000000111 1203011000 0001410000 13100-1125 020111?211 0101100111 111

Quindina kuna 3101001100 0120011000 1000000000 0000000111 1203011000 0001410000 0--40-1125 020111?211 0110110111 111

Quindina hermesi 3101001100 0220011000 1000000000 0000000111 1203011000 0001410000 ????061125 020111?211 1000010111 101

Quindina discolor 3101001100 2-20011000 1000000000 0000000111 1203011000 0001410000 1130060125 020111?210 0101101101 011

Quindina pendula sp. nov. 3101001100 0320011000 0000000000 0000000111 1203011000 0001410000 10-0061025 020111?200 0000100100 001

Based on Kury and Villarreal (2015), Pinto-da-Rocha and Bragagnolo (2017) and Pinzón-Morales and Pinto-da-Rocha (2020).

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

RESULTS

Quindina pendula Lancheros, Arias-

Cuellar & Pinto-da-Rocha sp. nov. zoobank.

org:act:69BE1437-BEA0-4006-9528-

329D725401DF

Etymology: The specific epithet, pendula,

is a Latin term meaning “pendant” and refers

to the pendular disc nest, contrasting with the

open mud nest attached to the subtract.

Type material: Holotype (MCUB-R-

AR-000938) adult male preserved in 70 %

ethanol, the nest is preserved dry, Colom-

bia, Cundinamarca, La Calera, vereda Mundo

Nuevo (4°39’55.7’’ N & 73°51’ 16.9” W; 2

693 m elevation), 16.II.2019, Daniel Arias

Cuellar leg. Paratypes: two males (MCUB-R-

AR-000936), from the same locality as the holo-

type, 19.XII.2020, Daniel Arias Cuellar leg.;

two females (MCUB-R-AR-000939), Colombia,

Cundinamarca, La Calera, vereda El Libano

(4°39’44’’ N & 74°0’31’’ W; 3 006 m elevation),

27.III.2021, Nicolás Briceño Avellaneda leg.

Diagnosis: Ocularium with a group of 3-6

tubercles near each eye, towards the center.

Prosoma with dark yellow background on most

tubercles both dorsally and ventrally. Area I

with a group of uniformly tubercles forming

a triangle on each side. Area III with two or

three pale yellow patches on each side. Area IV

with one pale yellow patch on each side. Dorsal

scutum areas and tergites without white spots.

Free tergites predominantly black on the sides

and yellow in the center.

Comparison: Comparison: Resembles Q.

bella, Q. bimaculata, Q. discolor, Q. hermesi, and

Q. marginata, by the lack of enlarged tubercles

on the lateral margins of the dorsal scutum.

It can be distinguished from Q. bella by the

presence of only small yellow tubercles on the

lateral margin of the dorsal scutum (a group of

larger, white tubercles on each side of the lateral

margin in Q. bella); from Q. bimaculata by the

presence of dark yellow background around

most of the tubercles on the dorsal and ventral

body, and in free tergites I-III (yellowish white

tubercles and a white patch with three tubercles

in each side of the lateral margin of the dorsal

scutum in Q. bimaculata); from Q. discolor by

the dark yellow background of the prosome

with black areas (presence of a white patch on

the posterior half of the prosoma in Q. discolor);

from Q. hermesi by presence of only yellow

and black coloration (anterior margin tubercles

white and a white spot on each side of the lat-

eral margin near the back of the dorsal scutum

in Q. hermesi); and from Q. marginata by the

smaller number of tubercles in the ocularium

and the absence of these in the posterior part

of the prosoma, the presence of only yellow,

brown and black coloration and a pair of paral-

lel spines in the free tergite III (presence of 18

tubercles in the center of the ocularium and 23

behind of this, silver tubercles and absence of

a pair of parallel spines in the free tergite III in

Q. marginata).

Description, Male Holotype

(MCUB-R-AR-000938)

Measurements: Carapace length: 1.4; cara-

pace width: 2.2; dorsal scutum length: 3.1; dor-

sal scutum width: 2.8; femur: 1.6, 3.6, 2.4, 3.3;

patella: 0.6, 1, 0.9, 1.1; tibia: 1.1, 2.5, 1.6, 2.1;

metatarsus: 1.9, 3.2, 2.5, 3.6. Variation in male

measurements (Table 2).

Live color (Fig. 1A): Dorsal shield with

brown background, ocularium with black

patches on the lateral areas, middle part of

the prosoma with a mushroom-shaped set of

brown patches. Black patches in 49 % of the

prosoma, the other 51 % of the prosoma brown.

Areas I and II with dark yellow dots. Spines of

area III black with two bright yellow patches on

each side. Most of the dorsal and ventral tuber-

cles have dark yellow patches. Lateral areas with

small irregular black patches. Lateral edges of

the shield are dark yellow. Free tergites black

with light yellow lateral patches. Legs light yel-

low with a pattern of irregular black patches.

Pedipalps and chelicerae are dark yellow with

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

Tabl e 2

Variation in body and appendage size in Quindina pendula sp. nov.

Males (n = 3) Females (n = 2)

Max. Min. Mean Max. Min. Mean

CL 1.6 1.4 1.5 1.2 1.2 1.2

CW 2.6 2.2 2.4 1.8 1.8 1.8

DSL 3.5 3.1 3.3 2.8 2.7 2.8

DSW 3.0 2.8 2.9 2.6 2.4 2.5

Fe I 1.7 1.6 1.7 1.1 1.1 1.1

Ti I 1.1 1.1 1.1 0.8 0.7 0.8

Fe II 3.6 3.3 3.5 2.1 2.1 2.1

Ti II 2.6 2.4 2.5 1.4 1.4 1.4

Fe III 2.5 2.2 2.4 1.5 1.5 1.5

Ti III 1.6 1.5 1.6 1.0 1.0 1.0

Fe IV 3.5 3.2 3.4 2.2 2.1 2.2

Ti IV 2.1 2.0 2.0 1.4 1.3 1.4

Abbreviations: CL = carapace length, CW = carapace width, DSL = dorsal scutum length, DSW = dorsal scutum width, Fe =

femur length, Ti = tibia length, n = number of specimens measured.

Fig. 1. Quindina pendula sp. nov. male holotype (MCUB-R-AR-000938). A-B. Habitus, A, dorsal view; B, lateral view. C.

Chelicerae hand with fixed and movable fingers. D-E. Pedipalp, trochanter to tarsus, C, meso-dorsal view; D, ventral view. F.

Leg I, trochanter to tarsus in ventral view. Scales: 1 mm.

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

black patches. Ventral body dark yellow with

heterogeneous black patches. Free sternites

with black lines and light yellow patches. UV

fluorescence is observed in free tergites I to III

(Fig. 2C and Fig. 2D).

Color in ethanol: Background of the dor-

sal scutum dark yellow. Ocularium with black

patches on the lateral areas. Black patches in

49 % of the prosoma, the other 51 % of the

prosoma dark yellow (Fig. 1A). Area I and II

Fig. 2. Observations of the adult male and nymphs of Quindina pendula sp. nov. in the terrarium. A-B. Adult male tending

the nest in the terrarium. C-D. UV observation for evidence of fluorescent areas. E. 2-day-old nymphs on the nest. F. 21-day-

old nymph. Photos: A, B, C and D. Daniel Arias; E and F. Héctor Lancheros.

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with dark yellow dots. Spines of area III black,

each with two or three pale yellow patches

forward on each side (Fig. 1A). Most of the

dorsal and ventral tubercles have dark yellow

patches. Lateral areas with small irregular black

patches (Fig. 1B). Free tergites with black lateral

patches. Yellow pedipalps with black patches.

Chelicerae with a dark yellow background with

heterogeneous black patches. Legs yellow with

black patches. Ventral part is dark yellow with

heterogeneous black patches. Free sternites

with black lines and dark yellow patches.

Dorsal scutum (Fig. 1A): Type β dorsal

scutum, anterior margin with one tubercle on

each side. Preocular monticle with some gran-

ules. Carapace with one tubercle at each side

of the preocular monticle, and one tubercle

forward or behind of each ozopore. Ocularium

with a small median depression, with two para-

medial groups of 3-6 tubercles. Lateral margins

with a row of 19-20 tubercles from center of

coxa III to almost posterior margin, elevated

posteriorly. Grooves of the dorsal scutum are

well marked. Area I with 7-8 tubercles forming

a triangle on each side. Area II with a row of

eleven tubercles. Area III with a pair of sharp,

slightly divergent spines, directed backwards;

with four tubercles forward of each spine and

one or two tubercles at the base or over these.

Area IV with four tubercles on each side. Pos-

terior margin with nine tubercles. Free tergites:

I and II with a row of twelve tubercles; III with

a pair of parallel spines directed backwards,

with four tubercles between spines and four at

each side. Anal plate with nine small-scattered

tubercles.

Ventral: Coxa I with 13 scattered tubercles.

Coxa II with 28 tubercles. Coxa III with 21

scattered tubercles. Coxa IV with more than

50 tubercles.

Legs (Fig. 1F): I and III with small tuber-

cles on the front, II with one medium-sized and

one large tubercle on the front. Trochanters:

I-IV with small scattered setiferous tubercles

in the dorsal, ventral and lateral parts. Femora

I-IV with rows of various setiferous tubercles;

the apex of the femora II-IV presents two spiny

lateral tubercles, the retrolateral larger than the

prolateral. Tarsal articles: 6(3)/12-13(3)/7/8.

Chelicerae (Fig. 1C): Basal segment with

a few small setiferous tubercles on the bulla.

Movable finger with four teeth; fixed finger

with five teeth.

Pedipalps (Fig. 1D and Fig. 1E): Tro-

chanter with a few small dorsal tubercles, one

ecto-lateral and one ventral setiferous tubercles.

Femur with small scattered dorsal tubercles.

Tibia: mesal iiIi, ectal iIi; Tarsus: mesal IiIi,

ectal IiIi.

Penis (Fig. 3): Ventral plate oval in shape,

lateral margin with a basal constriction, half of

the distal edge curved, three pairs of spatulate

macrosetae C; one pair of spatulate macrosetae

A directed towards the truncus, almost as long

as macrosetae C; one pair of conical macrose-

tae D; three pairs of macrosetae E on ventral

surface of ventral plate. Glans wide and conical,

stylus long, slightly wide at apex, with subme-

dian ventral process. Type 4 microsetae distrib-

uted in the ventral side of the ventral plate.

Female Paratype (MCUB-R-AR-000939)

Measurements: Carapace length: 1.2; cara-

pace width: 1.8; dorsal scutum length: 2.8; dor-

sal scutum width: 2.5; femur: 1.1, 2.1, 1.5, 2.2;

patella: 0.4, 0.6, 0.7, 0.7; tibia: 0.8, 1.4, 1.0, 1.4;

metatarsus: 1.1, 1.8, 1.5, 2.3. Variation in female

measurements (Table 2).

Females differ from males in several mor-

phological aspects and have dark brown spots

instead of black spots (Fig. 4). Anterior mar-

gin of the scutum with two tubercles on each

side. Ocularium with two paramedial groups

of tubercles. Lateral margins without a row of

tubercles. Area III and free tergite III lacking

a pair of spines. Free tergites II and III with

sharper and larger tubercles (Fig. 4B).

The number of tubercles is different in

areas I-IV of the dorsal scutum, the free tergites

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

and the anal plate. Area I with a row of two or

four tubercles on each side. Area II with a row

of three tubercles on each side. Area III with

four to eight tubercles on each side. Area IV

with three to four tubercles on each side. Pos-

terior margin with ten tubercles. Free tergites: I

with a row of ten tubercles; II with a row of 13

tubercles; III with eleven tubercles. Anal plate

with two rows of four small tubercles. Tarsal

counts smaller than males: 5(3)/8(3)/6/6.

Distribution (Fig. 5): Recorded in the

department of Cundinamarca: municipalities

of La Calera (Fig. 6), Choachí (Fig. 7A, Fig. 7B,

Fig. 7C), Tena; and in the Distrito Capital local-

ity of Chapinero (Bogotá). The elevation range

is between 2 040 and 3 150 m and the observed

vegetation corresponds to the Andean Forest

formation. In addition, there are photographic

records of the same type of nests of this spe-

cies in the municipality of Saboyá (department

Fig. 3. Penis of Quindina pendula sp. nov., holotype (MCUB-R-AR-000938). A. dorsal view, B. ventral view, C. ventrolateral

view, D. lateral view. MS = macrosetae, μS 4= type 4 microsetae. Scale: 0.01 mm.

Fig. 4. Comparison of male and female of Quindina pendula sp. nov.. A. Dorsal view of the male holotype (MCUB-R-

AR-000938), B. Dorsal view of a female paratype (MCUB-R-AR-000939). Scale 1mm. Photos: Héctor Lancheros.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

of Boyacá), at an elevation of 2 619 m, these

could belong to the same species, but there is

no record, of the individuals from this locality,

deposited in scientific collections.

Nest characteristics: We report a new type

of nest in Quindina pendula sp. nov. This spe-

cies builds pendular disc-shaped nests in which

the female lays the eggs that are cared for by the

male. Most of these pendular disc nests were

observed on tree branches in well-preserved

Andean Forest areas, generally on branches

colonized by foliose liverworts. Apparently, the

nest is composed of mud. However, upon close

examination, one can observe only plant frag-

ments such as bark segments and bryophytes

(Fig. 8), united by an adherent substance that

is presumably secreted by the harvestmen.

A detailed examination of one of the nests

revealed the presence of small arthropods on

its surface: a Psocoptera nymph of the fam-

ily Ectopsocidae and a mite, apparently of the

order Mesostigmata (Fig. 8E and Fig. 8F).

The nests were found suspended from

various structures such as sections of leafy liv-

erworts (Frullania atrata, F. cf. brasiliensis and

Plagiochila sp.), thin branches of trees, leaves of

bromeliads, among others (Fig. 8B, Fig. 8C, Fig.

9 and Fig. 10). Seven nests were observed in a

single tree, four of these on a branch. Two nests

were observed (Fig. 10) in an abandoned hut

in the forest of the vereda Mundo Nuevo, one

hanging from the lower part of the sill beam

(Fig. 10B and Fig. 10C) and the other inside

near the roof, suspended from a blackberry

plant that was growing partially inside the hut

Fig. 5. Sites where Quindina pendula sp. nov. or pendular disc nests have been recorded in the departments of Cundinamarca

and Boyacá. The color of the dots indicates the type of record; the blue dot represents the record of the nests without the

individuals, so there is no certainty that it is the same species.

14 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

Fig. 6. Municipality of La Calera, vereda Mundo Nuevo. Elevation: 2 693 m. A. General view of the vegetation cover in the

area. B. Relief of the area. C. Abandoned hut where two nests were observed. D. Nest suspended from liverworts on a tree

branch. E. Adult of Quindina pendula sp. nov. protecting the nest. Photos: A, B, C and D. Héctor Lancheros; E. Daniel Arias.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

Fig. 7. A-C. Observations in the municipality of Choachí, Forest near La Chorrera waterfall, elevation: 2 620 m. A. High cover

of liverworts on tree branches in the forest interior. B. Nest hanging from liverworts on a tree branch. C. Male harvestman

found in the nest. D-F. Observations of juvenile individuals in the wild. D-E. Shortly after hatching the nymphs remain in the

nest. F. Juvenile individual found on the ground under a log. Photos: A, B and C. Daniel Arias; D, E and F. Héctor Lancheros.

16 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

Fig. 8. Nest structure of Quindina pendula sp. nov. A. General view of the nest. B-C. Detail showing the elements from which

it hangs, a thin branch and a leaf of Tillandsia sp., we observed that the nest is composed of small fragments of plant bark.

D. Phyllidium of a liverwort that is part of the nest structure. E-F. Mite, probably of the order Mesostigmata, and Psocoptera

nymph of the family Ectopsocidae found in the nest. Photos: Héctor Lancheros.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

Fig. 9. Nests of Quindina pendula sp. nov. found in tree branches, suspended from different structures A. tree with high

epiphytism of foliose liverworts. B. Nest built on the apex of an epiphyte of the genus Tillandsia. C. Nest built around a

liverwort with two support points. D. Nest built around the middle part of a liverwort. E. Nest built on the terminal part of

a slender branch. Photos: Héctor Lancheros.

(Fig. 10D and Fig. 10E). Nests generally hang

from a single point, but occasionally hang from

two (Fig. 9C and Fig. 10C); In most observa-

tions the nest was at the terminal part of the

supporting element, but occasionally it was

observed in the middle part, with a section of

the element protruding at the bottom (Fig. 9C,

Fig. 9D and Fig. 7D).

The observed values of mass and dimen-

sions were as follows: fresh mass 0.12-0.6 g; dry

mass 0.08-0.01 g; length 16-20 mm, width 15-21

mm, average diameter 15-21 mm; thickness

18 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

3.80-4.35 mm; surface area (sum of the two

faces) 353-693 mm². The value of fresh mass

is highly variable due to the different degree

of hydration, which depends on the location

of the nest and the precipitation before data

collection. For dry mass n = 8, for the other

variables n = 12.

Observations on parental care: Adults

were not observed in the nests during the day.

In the field, after 6:00 p.m., we observed that

males remained constantly on or near the nest

(Fig. 6E, Fig. 10D and Fig. 10E). In the lab, we

observed that, at night, the male takes care of

the nest, even if it is damaged, repairing the

nest when it suffered minor damage (Fig. 2A,

Fig. 2B, Fig. 2C and Fig 2D). During the day

the male remained hidden at the bottom of

branches. We also observed that, when the nest

of another male is placed inside the box, the

male continues to take care of it, even if it is not

his own. We have not recorded any females in

or near the nests.

Developmental stages from laying to

hatching: Initially the eggs were completely

covered by nest material, but as they developed

and increased in size, they became partially

exposed. It appears that deterioration of the

nest due to rain also causes them to gradually

become exposed. The fact of being partially

uncovered does not cause direct changes in

the color of the eggs, they only change color

gradually as they advance in their development.

Preliminary observations show four stages of

development from egg laying to hatching: white

Fig. 10. Nests of Quindina pendula sp. nov. found in an abandoned hut inside the forest of the vereda Mundo Nuevo. A. Hut

hidden inside the forest. B. Nest hanging on the lower part of the sill beam. C. Detail of the nest hanging on the sill beam.

D. Adult harvestman tending a nest on a blackberry plant. E. Nest suspended from the blackberry plant (Rubus sp.) inside

the hut. Photos: Héctor Lancheros.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

eggs with no visible differentiated structures;

brown eggs with visible appendages; black eggs;

emergence of the anterior part of the body

(Fig. 11). At least 22 days elapse between lay-

ing and the second stage. Then, brown eggs

take approximately 12 days to develop into

black eggs and 10 to 12 days elapse between

the third and fourth stages. In total, at least

45 days elapse from laying to the beginning of

hatching (Fig. 11).

In the field, the nymphs remain in the

nest shortly after hatching (Fig. 7D and Fig.

7E). They can fall to the ground when hit by

raindrops or can take shelter again in the holes

on the nest that remain after hatching. This was

observed once in the first nest recorded in the

Fig. 11. Nest of Quindina pendula sp. nov. with eggs at different stages of development, the circles on the absorbent paper

are 1 mm in diameter, the eggs have a diameter of 0.55 mm. A-B. 10 days after collection, with black and white eggs, two

hatchings are also observed (orange circles). C-D. 22 days after collection, on one side of the nest there are eggs in the second

and third stage of development, brown and black, respectively; also, an egg shell is observed after hatching (green circle). E-F.

22 days after collection, on the other side of the nest there are eggs in the first, third and fourth stage of development: white,

black and initiating hatching (red circles), respectively. Photos: Héctor Lancheros.

20 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

vereda Mundo Nuevo during the day, when the

adult male was absent. A juvenile individual was

found on the ground, under a log; this may be

the environment in which they remain before

reaching maturity and initiate nest construc-

tion (Fig. 7F). As in the natural environment, in

the terrarium, nymphs remained in the nest for

the first few days (Fig. 2E and Fig. 2F).

Phylogenetic analysis: Using the equal

weights method, we obtained 30 equally parsi-

monious trees with 380 steps. From these, we

obtained a strict consensus tree of 416 steps

and a majority consensus tree of 386 steps.

Using the implicit weighting method, with k

= 9.2188, obtained with the setk.run script, we

obtained a tree of 381 steps. Using successive

weighting, with the rc index, we obtained a tree

of 381 steps. With all methods, we recovered

Nomoclastidae as a monophyletic unit, sister to

Microsetata; in the latter, the relative position of

the species is similar to that obtained by Kury

and Villarreal (2015), with the exception of the

species in the Ampycus telifer, Neopachyloides

sp., Glysterus metatarsalis and Nesopachylus

monoceros clade, with the strict consensus,

implicit weighting and successive weighting

methods (Appendix 1A, Appendix 1B, Appen-

dix 1C, Appendix 1D). The relative position of

the other Gonyleptoidea families, as well as that

of the outgroups, was the same as that obtained

Kury and Villarreal (2015) using the successive

weighting method but slightly different with

the implicit weighting method, the latter dif-

fers in the position of Sorensenius and Dibunus

(Appendix 1C, Appendix 1D).

Concerning the relative position of the

genera within Nomoclastidae, the different

methods showed Zamora as sister to the other

genera. Nevertheless, the methods do not agree

on their relative positions. The strict consensus

tree with equal weights showed a polytomy

between all species of Callcosma and the genera

Kichua, Napostygnus, Nomoclastes and Quindi-

na (Appendix 1A). The majority consensus tree

with equal weights showed a polytomy between

Napostygnus, Nomoclastes and the clade (Kich-

ua (Callcosma Quindina)) (Appendix 1B). The

successive and the implicit weighting tree shows

a well-defined topology, with Nomoclastes sister

to the group ((Kichua (Napostygnus Callcos-

ma)) Quindina) (Appendix 1C, Appendix 1D);

in addition, the highest relative Bremer sup-

port values were obtained with the successive

weighting method (Appendix 1D).

Regardless, the main object of this study is

the genus Quindina, the topologies we obtained

with the implicit and successive weighting

methods show Quindina marginata as sister to

the remaining species of the genus (including

Q. pendula sp. nov.). Similarly, we recovered

Quindina pendula sp. nov. as sister to the

remaining 10 species of the genus. Sensitivity

plots (Navajo rugs) indicate that the two topol-

ogies coincide in seven of the twelve species of

the genus Quindina, the relative position vary-

ing within the clade formed by Q. hermesi, Q.

bella, Q. kuna, Q. limbata and Q. albiocularia.

We obtained the highest values of Bremer rela-

tive support with the topology recovered by the

successive weighting method (values between

53 and 74 for each clade within the genus), so

we chose this cladogram for the comparison

with the other methods (with the sensitivity

plots) and the approach of the possible hypoth-

eses of the origin of the pendular disc-shaped

and open mud nests (Fig. 12).

Hypotheses of the origin of the nests: We

have obtained seven hypotheses for the origin

of the two types of nests in the genus Quindina

using the method of most parsimonious recon-

structions with the tree obtained by the suc-

cessive weighting (Fig. 13). The characteristics

observed in the hypotheses are summarized in

Table 3. For each characteristic (nest type for

each reference group) we give the frequency

of each condition (ancestral, derived or in an

internal group), the sum being equal to seven.

The last column shows the condition that we

observed most frequently (in most hypotheses).

The obtained results differ on the origin

of the nests and the type of nest that first

appeared. The lack of data for other nomoclas-

tid genera hindered us to know if these genera

share nest construction (Fig 13F, Fig. 13G)

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

Fig. 12. Cladogram of Nomoclastidae obtained by successive weighting with rc, including the new species. Sensitivity plots

(Navajo rugs) indicate, with red squares, agreement with groups obtained by other methods: strict Nelsen consensus (NE).,

majority consensus > 50 % (MA) and implicit weighting with k = 9.2188 (IW). Numbers in each branch indicate relative

Bremer supports. Next to each species there are drawings indicating the type of nest observed in each; nest descriptions are

presented in the boxes on the right. The other species of Gonyleptoidea have been omitted as they are not the subject of

discussion in this paper (Appendix 1D).

Tabl e 3

Characteristics present in the seven hypotheses with the frequency of occurrence of each condition.

Characteristic Variable

Condition Most common

condition

Ancestral Derived In an internal

group

1. Presence of pendular disc nest in the genus

Quindina.

E C A B G

5In an internal group

2. Presence of pendular disc nest in the sister

group of Q. marginata.

D F

E C A G

4In an internal group

3. Presence of pendular disc nest in Q. pendula. H

D B F

E C A G

4 0 Derived

4. Presence of open mud nests in the genus

Quindina.

C A D B F

5In an internal group

5. Presence of open mud nest in the sister group

of Q. marginata.

E G

A D B F

4In an internal group

6. Presence of open mud nest in the sister group

of Q. pendula.

E C G

A D B F

4 0 Derived

H = hypotheses in which this condition is met (Fig. 13); F = frequency (number of hypotheses in which it is met).

22 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

Fig. 13. Hypothesized origin of nests in the genus Quindina. A. Independent origin of the pendular disc nest in Q. pendula

and the open mud nest in the sister group of this. B. Origin of the pendular disc nest in the common ancestor of the sister

group of Q. marginata and modification to the open mud nest in the sister group of Q. pendula. C. Origin of the open mud

nest in the common ancestor of the sister group of Q. marginata and modification toward the pendular disc nest in Q.

pendula. D. Origin of the pendular disc nest in the common ancestor of Quindina spp. and modification to the open mud

nest in the sister group of Q. pendula. E. Origin of the open mud nest in the common ancestor of all Quindina species and

modification toward the pendular disc nest in Q. pendula. F. Origin of the pendular disc nest in an ancestor external to the

genus Quindina with modification toward the open mud nest in the sister group of Q. pendula. G. Origin of the open mud

nest in an ancestor external to the genus Quindina with modification toward the pendular disc nest in Q. pendula.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

or if this is exclusive for Quindina spp. (Fig.

13A, Fig. 13B, Fig. 13C, Fig. 13D, Fig. 13E).

The absence of reproductive data for the first

derived species of Quindina, Q. marginata, is

also an obstacle in understanding how nest

construction evolved within the group. At this

moment it is impossible to know if the pendular

disc nest is exclusive for Quindina pendula sp.

nov., or if it evolved first and then was replaced

by an open mud nest. The only consensus

among all hypotheses is that most species of

Quindina share an open mud nest as a strategy

to lay eggs and raise nymphs. Thus, this clade is

composed of species for which we observed an

open mud nest in the field (Q. albomarginis, Q.

morae, Q. burbayar, Q. kuna, and Q. limbata)

and other species that we hypothesize also build

this kind of nest.

DISCUSSION

Previously described nest-building har-

vestmen are distributed across the territories

of Costa Rica (Q. limbata) and Panama (Q.

albomarginis, Q. burbayar, Q. kuna and Q.

morae) (Pinto-da-Rocha & Bragagnolo, 2017),

making Q. pendula sp. nov. the first nest-pro-

ducing species recorded in Colombia. In addi-

tion, this new species represents the first record

of a pendular disc nest, built with fragments

of bark and other plant remains. The pendular

disc nest differs significantly in shape, substrate

type and substrate attachment, from the previ-

ously reported nest type, which is an open mud

nest. This last kind of nest is totally attached

to the substrate, surrounded by walls and built

with bark fragments and other materials. Both

nest types do share some similarities, specifical-

ly in the type of materials which compose them

(Mora, 1990; Rodríguez & Guerrero, 1976).

Nevertheless, the composition of the previ-

ously reported nests needs to be re-examined

through detailed microscopic observations in

order to verify its accuracy.

The presence of small arthropods on the

surface of the pendular disc nest, such as pso-

cids and mites, could be related to their use of

the nest as a food source. Insects of the order

Psocoptera are mainly phytophagous, feeding

on algae, fungi, pollen and fallen plant parts

(Gillott, 2005, Chapter 8). On the other hand,

Mesostigmata mites are mainly free-living

predators, where some are phoretic on insects

and some consume pollen, nectar and fungi

(Dhooria, 2016, Chapter 5). The Ectopsocidae

(Psocoptera) may roam the nest consuming

plant debris or the mycelium of fungi growing

on its surface. Meanwhile, the mites may also be

in the nest to eat mycelium or, instead, act as a

predatory species, consuming small arthropods

such as psocoptera or harvestmen nymphs.

The addition of Q. pendula sp. nov. to the

phylogenetic analysis result in great differences

in the relationships within Quindina when

compared with Pinzón-Morales and Pinto-

da-Rocha (2020); the implicit and successive

weighting methods coincide only in Q. mar-

ginata as sister group of the remaining species

in the genus and in the clade (Q. albomarginis

(Q. morae Q. burbayar)). However, the topology

of the different nomoclastid genera is identical

between our findings with these two methods

and the previous study.

Aside from Quindina, no reproductive

studies have tackled other nomoclastid genera.

Field observations did not find nests in the

area of occurrence for Kichua rheimsae Pinto-

da-Rocha & Bragagnolo 2017, Callcosma abra-

patricia Pinto-da-Rocha & Bragagnolo 2017

or C. gracilima Roewer 1932 (Cabra, Rheims,

pers. comm.; Pinto-da-Rocha pers. obs.). How-

ever, the lack of observation of nests in these

other species is not sufficient evidence of their

absence, given that the respective field trips

lasted only a few days. It is more likely that all

or at least most of the species of the genus Quin-

dina build nests, since this behavior has been

observed in half of the known species. Further-

more, the ones that have no records (Q. mar-

ginata, Q. discolor, Q. bimaculata, Q. hermesi, Q.

bella and Q. albiocularia) have few observations

(Pinto-da-Rocha & Bragagnolo, 2017; Pinzón-

Morales & Pinto-da-Rocha, 2020). The fact that

we did not find the nestless species on the same

branch, but on different branches (interspersed

with species that do build nests) reinforces the

24 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

idea that the absence of nest reports for these

species is due to a lack of field observation.

We also call attention to the fact that

44 years elapsed between the description of

Q. albomarginis (Chamberlin, 1925) and the

publication of the first article describing open

mud nests (Rodríguez & Guerrero, 1976). In

the case of Q. limbata, 71 years separate the

species description (Roewer, 1943) from the

first published description of its nest (Rojas

& Solano, 2014). The other three species with

known nests were reported by Pinto-da-Rocha

and Bragagnolo (2017). At this point, it is dif-

ficult to know if the pendular disc nest is an

autapomorphy of Q. pendula sp. nov. or if it

arose earlier, since we have no available data for

either Q. marginata or the other nomoclastid

genera. More knowledge of parental care in Q.

marginata would help us clarify the origin of

the nests in the genus Quindina, as this is the

sister group for the remaining species of the

genus. In the event that it builds a pendular

disc nest, hypotheses about its origin in the

most recent common ancestor of the genus

or even in an ancestor external to the genus

would be more plausible. On the other hand, if

Q. marginata builds open mud nests, it would

be more likely that the pendular disc nest is an

autapomorphy of Q. pendula sp. nov. Finally, in

the case that Q. marginata does not build nests,

it would be more probable that the origin of

the nests is in one of the internal groups of the

genus Quindina.

This study enriches our knowledge on

parental care behavior and nest construction

in the genus Quindina. Until recently, only one

type of nest had been reported for this genus

in the scientific literature (Pinto-da-Rocha &

Bragagnolo, 2017; Quesada-Hildalgo et al.,

2019; Rojas et al., 2019) and now we know that

there are variations in the shape and structure

of the nests. Likewise, we can observe that our

knowledge about the species within this genus,

as well as for the whole Nomoclastidae family,

has increased steadily in the last ten years (Kury

et al., 2024; Kury & Villarreal, 2015; Pinto-da-

Rocha & Bragagnolo, 2017; Pinzón-Morales &

Pinto-da-Rocha, 2020; Quesada-Hidalgo et al.,

2019; Requena et al., 2014; Rojas et al., 2019;

Rojas & Solano, 2014; Villarreal & Kury, 2023).

Beyond the genus Quindina and order Opilion-

es as a whole, the results obtained in this study

are of great importance in our understanding of

egg care and nest construction in arthropods.

The construction of hanging nests with frag-

ments of plant material has not been previously

reported in other groups of arachnids or in any

of the classes within Arthropoda, so our find-

ings are relevant for any further studies about

the evolution of parental care in this phylum.

Ethical statement: the authors declare that

they all agree with this publication and made

significant contributions; that there is no con-

flict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

See supplementary material

a53v72n1-MS1

ACKNOWLEDGMENTS

We express our gratitude to all the people

who supported the development of this work:

Nicolás Briceño, who assisted us with the col-

lection of specimens; Manuel David Cortés

who identified the bryophyte species; Osvaldo

Villarreal, who provided us with the setk script

and recorded the species and nests in the

municipality of Tena (department of Cundina-

marca); Christian Franco García, who recorded

the pendular disc nest in the municipality of

Saboyá (department of Boyacá); Daniela Ahu-

mada, who helped us with the extraction and

observation of the male genitalia; Conchita

Pinzón, who offered initial feedback on the

phylogenetic analysis and species description;

Paula Nuñez, who aided in the elaboration of

the illustrations; Víctor Rodríguez Saavedra,

who registered the specimens in the collection

of the Museo de Ciencias de la Universidad El

Bosque MCUB; Cristina Rheims, who kindly

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e61426, enero-diciembre 2024 (Publicado Nov. 20, 2024)

reviewed the language of an early version of

the manuscript; David Hernández, who kindly

checked and reviewed the language for the

final manuscript; and the anonymous review-

ers, who provided their valuable feedback. We

also give thanks to the Willi Hennig Society

for providing us with the TNT v.1.5 soft-

ware. The participation of Héctor Lancheros

in this study was done within the framework

of the “Ecological Interactions of Plants in

Colombian Ecosystems: Phase I” project, from

the Biology Research Group GRIB of Uni-

versidad El Bosque. This study was funded

by CAPES, CNPq (306722/2018-6), FAPESP

(BIOTA, 2013/50297-0), NSF (DOB 1343578),

and NASA to Ricardo Pinto-da-Rocha.

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