Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e51166, enero-diciembre 2023 (Publicado May. 16, 2023)

The characteristics of stingless bee nests (Apidae: Meliponini)

in the Cycloop Mountain Nature Reserve, Indonesia

David Reinhard Jesajas1; https://orcid.org/0000-0001-8294-9612

Tri Atmowidi2*; https://orcid.org/0000-0002-2179-3914

Berry Juliandi2; https://orcid.org/0000-0003-0348-5675

Sih Kahono3; http://orchid.org/0000-0001-7222-3180

1. Student of Animal Bioscience Study Program, Graduate School, IPB University, Bogor 16680, Indonesia;

juandaud10@gmail.com

2. Department of Biology, Faculty of Mathematics and Natural Sciences, IPB University, Bogor 16680, Indonesia;

atmowidi@apps.ipb.ac.id (*Correspondence), bjuliandi@apps.ipb.ac.id

3. Research Center for Applied Zoology-BRIN, Cibinong 16912, Indonesia; sihkahono@gmail.com

Received 29-V-2022. Corrected 06-II-2023. Accepted 02-V-2023.

ABSTRACT

Introduction: Stingless bees are widespread in tropical and subtropical regions. In Indonesia, the distribution of

stingless bees are grouped in three regions, namely Indo-Malayan, Wallacea, and Indo-Australian. Ten species

of stingless bees have been recorded in Papua, seven of which are endemic. The Cycloop Mountains Nature

Reserve (CMNR) is one of the conservation area in Papua, Indonesia, for flora and fauna. Unfortunately, the

study of the diversity of stingless bees in Papua has been limited.

Objective: To measure the diversity, nesting sites, nest entrance characteristics and nest architecture of sting-

less bees.

Methods: Observation of the stingless bee nests in the nature reserves and in the residential areas used a road

sampling method and information from local people, respectively. A total of 22 colonies were studied.

Results: Two species of stingless bee were found, namely Tetragonula sapiens (Cockerell, 1911) and

Heterotrigona (Platytrigona) planifrons (Smith, 1865). The current study showed new distribution records for

T. sapiens and H. planifrons in the CMNR. The nesting site of T. sapiens was commonly found in house founda-

tion, while that of H. planifrons was in coconut palm cavities. The nest entrance of T. sapiens varied, i.e., ellipti-

cal, oval, rounded, irregular, horizontally or vertically elongated. Meanwhile, the nest entrance of H. planifrons

was vertically elongated. The brood cells of T. sapiens varied, i.e., vertical, horizontal, or semi-clusters, while

in H. planifrons was layered vertically.

Conclusions: Two species of stingless bees found, T. sapiens and H. planifrons, showed a new distribution

records and T. sapiens was a dominant species in Papua. The nest entrance of the species varied in shape, color,

and texture.

Key words: Cycloop Mountains Nature Reserve; diversity; nest characteristics; stingless bee.

RESUMEN

Ubicación y características de los nidos de abejas sin aguijón (Apidae: Meliponini)

en la Reserva Natural Cycloop Mountain, Indonesia.

Introducción: Las abejas sin aguijón están muy extendidas en las regiones tropicales y subtropicales. En

Indonesia, la distribución de las abejas sin aguijón se agrupa en tres regiones: Indo-Malayan, Wallacea e

https://doi.org/10.15517/rev.biol.trop..v71i1.51166

INVERTEBRATE BIOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e51166, enero-diciembre 2023 (Publicado May. 16, 2023)

INTRODUCCIÓN

The stingless bees (Apidae: Meliponini)

are a cosmopolitan group of bees in the tropics

and subtropics. Stingless bees are distributed in

tropical regions of the world across the Neo-

tropical, Afrotropical, and Australasian realms

(Michener, 2000) and are the highest diversity

among bees consist of more than 600 species

(Rasmussen, 2008; Rasmussen & Cameron,

2007). Indonesia, as part of the Indo-Malayan

and Australasian ecoregions, has 46 record-

ed stingless bee species across the islands

of Sumatra, Java, Borneo, Timor, Sulawesi,

Ambon, Maluku, and Papua (Kahono et al.,

2018). The species of this group are small to

medium sized with a vestigial stinger and live

in colonies (Michener, 1974). These beehives

can be found under the soil surface, in tree and

wood cavities, house wall cavities, or hollows

of bamboo trees (Dollin et al., 1997; Engel et

al., 2019; Michener, 1974; Sakagami et al.,

1983). Stingless bees have a high prospect

as a pollinator of agricultural crops related

to its small body size, have a high foraging

activity, and high adaptation to environmental

stress (Kahono, 2015). Stingless bees play an

important role in pollinating of various plant

species (Michener, 2007), include in mustard

(Atmowidi et al., 2007), Tetragonula laeviceps

in strawberry (Fragaria x annanassa) and Het-

erotrigona itama in melon (Cucumis melo) in

the greenhouse (Atmowidi et al., 2022). Nest-

ing preference and nest characteristics, such as

the morphology of nest entrance, nest architec-

ture, and nest height from the ground are sup-

porting characters that can be used to identify

stingless bee species. Nest entrances vary in

shape, texture, length, ornamentation, and color

according to each species (Anaktototy et al.,

2021; Kelly et al., 2014). For example, the nest

entrance of Heterotrigona itama (Cockerell,

1918) is funnel-shaped and in Geniotrigona

thoracica (Smith, 1857) is rounded (Kelly et

al., 2014). The internal nest architecture con-

sists of brood cells, honey and pollen pots (Efin

et al., 2019; Franck et al., 2004; Michener,

1974; Sayusti et al., 2021).

The Cycloops Mountains Nature Reserve

(CMNR) located in the Jayapura region of

Papua Province, Indonesia. It is a tropical for-

est area with high biodiversity and ecological

functions that are important for human life.

Human activities in this area have occurred for

a long time that impact the biodiversity, includ-

ing stingless bees. Forest disturbance in CMNR

Indo-Australian. Se han registrado diez especies de abejas sin aguijón en Papua, siete de las cuales son endémi-

cas. La Reserva Natural de las Montañas Cycloop (CMNR) es una de las áreas para la conservación de flora y

la fauna en Papua, Indonesia. Desafortunadamente, el estudio de la diversidad de abejas sin aguijón en Papua

ha sido limitado.

Objetivo: Medir la diversidad, los sitios de anidación y describir la arquitectura y características de entrada al

nido de las abejas sin aguijón.

Métodos: Se observaron los nidos de abejas sin aguijón en reservas naturales y áreas residenciales, mediante el

método de muestreo de caminos e información de la población local, respectivamente. Se estudiaron 22 colonias.

Resultados: Se encontraron dos especies de abejas sin aguijón, Tetragonula sapiens (Cockerell, 1911) y

Heterotrigona (Platytrigona) planifrons (Smith, 1865). Este estudio mostró nuevos registros de distribución de T.

sapiens y H. planifrons en el CMNR. El sitio de anidación de T. sapiens se encontró comúnmente en los cimien-

tos de las casas, mientras que el de H. planifrons estaba en las cavidades de las palmas de coco. La forma de la

entrada al nido de T. sapiens varió: elíptica, ovalada, redondeada, irregular, alargada horizontal o verticalmente.

Mientras tanto, la entrada al de H. planifrons se alarga verticalmente. Las celdas de cría de T. sapiens variaron,

entre verticales, horizontales o semi-racimos, mientras que en H. planifrons eran verticales.

Conclusiones: Se encontraron dos especies de abejas sin aguijón, T. sapiens y H. planifrons, que mostraron nue-

vos registros de distribución y T. sapiens fue una especie dominante en Papúa. La entrada al nido de las especies

varió en forma, color y textura.

Palabras clave: reserva natural montañosa Cycloop; diversidad; características de nido; abejas sin aguijón.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e51166, enero-diciembre 2023 (Publicado May. 16, 2023)

is mainly caused by community socio-econom-

ic activity (Ngutra, 2017). Limited informa-

tion and knowledge are a limiting factor for

the local community. People in Papua haven’t

cultivated yet the stingless bees for producing

honey and propolis. Community knowledge is

key factor affecting stingless bee conservation.

Until now, poor knowledge of taxonomy and

phylogeny of these bees in the studied región

are based for conducting this research. There-

fore, studying the diversity and nests charac-

teristics of stingless bees in the CMNR area

is needed. This study aims to determine the

species richness, characteristics of the nesting

sites, and nest architecture of stingless bees in

the CMNR, Papua, Indoesia.

MATERIALS AND METHODS

Observation of nest characteristic of

stingless bees: The observations of stingless

bees in the CMNR were conducted at three

locations, namely Maribu village (including

Mount Batu, Dumukribun, Dawari, and Maribu

Kampung), Dosay village (including Kausei

and Dansari), and Kemiri village (including

Kemiri-2 of Forestry Residential) (Table 1).

Nest of stingless bee was documented using a

Nikon D5300 camera. The nest characteristics

at Mount Batu and Dawari were observed by

a road sampling method (Garton et al., 2004).

While in Maribu Kampung, Dosay village, and

Kemiri village, sampling was based on infor-

mation from local people. The nest features

TABLE 1

Study sites and the number of colonies of stingless in Maribu, Dosay, and Kemiri villages of Papua

Maribu Village Coordinate/Altitude (mdpl) Colony

number Species Nesting sites

Gunung Batu

(Dumukribun)

02°27’31.7” S & 140°23’35.8” E/364 1T. sapiens Mountains-coconut tree

Dawari 02°28’47.5” S & 140°21’48.9” E/142 2T. sapiens Local farm-dry wood

02°28’54.1” S & 140°21’43.7” E/126 3Local farm-coconut tree

02°29’05.0” S & 140°21’37.3” E/160 4T. sapiens Local farm-dry wood

02°28’42.1” S & 140°21’47.0” E/133 5T. sapiens Local farm-house foundation

Maribu Kampung 02°29’00.9” S & 140°22’15.8”E/118 6T. sapiens Housing-coconut tree

02°29’06.0” S & 140°22’13.7” E/112 7T. sapiens Housing-coconut tree

02°29’03.6” S & 140°22’16.4” E/122 8T. sapiens Housing-fern tree

02°29’06.5” S & 140°22’12.0” E/109 9T. sapiens Housing-house foundation

02°29’08.5” S & 140°22’09.2” E/112 10 T. sapiens Housing-fern tree

02°29’20.3” S & 140°22’05.1”E/80 11 T. sapiens Housing-house wall

02°29’11.1” S & 140°22’10.6”E/107 12 T. sapiens Housing-house wall

Dosay Village

Kausei 02°30’42.1” S & 140°24’05.8”E/127 1H. planifrons Local farm-coconut tree

02°30’41.9” S & 140°24’07.3”E/128 2T. sapiens Housing-house foundation

02°30’42.2” S & 140°24’07.4” E/128 3T. sapiens Housing-house foundation

02°30’42.3” S & 140°24’07.5”E/129 4T. sapiens Housing-house foundation

02°30’43.6” S & 140°24’07.8”E/125 5T. sapiens Housing-house foundation

Dansari 02°30’24.8” S & 140°24’29.5”E/140 6T. sapiens Housing-soil

Kemiri Village

Kemiri-2

(Forestry Residential)

02°33’17.8” S & 140°29’06.0”E/117 1T. sapiens Housing-house wall

02°33’10.7” S & 140°29’02.8”E/125 2T. sapiens Housing-house foundation

02°33’11.1” S & 140°29’03.0”E/125 3T. sapiens Housing-house foundation

02°33’11.1” S & 140°29’03.4”E/133 4T. sapiens Housing-house foundation

Total colony (%)

22 21 (95 %) 1 (5 %)

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e51166, enero-diciembre 2023 (Publicado May. 16, 2023)

observed were nesting site, nest entrance

measurements (length, diameter, height from

ground), shape, and texture (Dollin et al., 1997;

Kelly et al., 2014). The coordinates of nest-

ing sites were determined using GPS (Garmin

etrex 10).

Collection, preservation, and identifica-

tion of stingless bee specimens: Five indi-

viduals of worker bees from each colony were

collected. The samples were then preserved

using the dry preservation method (Borror et

al., 1989). Samples were stored in bottles and

labeled. Identification of stingless bee speci-

mens were based on Sakagami et al. (1990),

Dollin et al. (1997), Rasmussen et al. (2017),

Engel et al. (2019), and Engel (2019). The

identification was carried out at the Labo-

ratory of Animal Biosystematics and Ecol-

ogy, Department of Biology, IPB University,

Bogor, Indonesia. Then, the specimens were

verified and deposited at the Research Cen-

ter for Ecology and Ethnobiology, Research

and Innovation Agency (BRIN), Cibinong,

West Java, Indonesia.

Data analysis: The t-test analysis was

used to compare the nest entrance variation

within species of T. sapiens, consisting of the

width and height of the nest entrance, funnel

length, and the height from the ground using

Paleontological Statistics version 4.09 (Ham-

mer et al. 2001).

RESULTS

Richness of stingless bee: Two species

of stingless bees were found in the three vil-

lages, namely Tetragonula sapiens and Het-

erotrigona (Platytrigona) planifrons (Fig. 1).

A total of twenty-two colonies were found,

i.e., one in Maribu Village including Gunung

Batu (Dumukribun), four in Dawari, seven in

Maribu Kampung, five in Dosay village and

Kausei, one in Dansari, and five in Kemiri

Village includes the Kemiri-2 of forestry resi-

dential (Table 1). Tetragonula sapiens was the

dominant species (21 colonies; 95 %) found

in all observation sites, while, Heterotrigona

planifrons (one colony; 5 %) was only found in

Kausei village.

Nesting sites: The dominant nesting site

of T. sapiens was in house foundations (nine

colonies, 43 %), followed by coconut tree cavi-

ties (four colonies, 19 %), house walls (three

colonies, 14 %), dry wood (two colonies,

10 %), fern tree (two colonies, 10 %), and

in the soil (one colony, 5 %). The nest of H.

planifrons (one colony) was found in a coconut

tree cavity.

Nest entrance characteristics: The nest

entrance tunnel of T. sapiens varied, being

horizontally elongated (4 colonies), oval (6

colonies), round (2 colonies), irregular (2 col-

onies), ellipses (4 colonies), and vertically

Fig. 1. Worker of two species of stingless bees found in Cycloop Mountains Nature Reserve. A. Tetragonula sapiens and

B. Heterotrigona planifrons.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e51166, enero-diciembre 2023 (Publicado May. 16, 2023)

elongated (2 colonies), while in H. planifrons

it was vertically elongated (1 coloni). The color

of T. sapiens entrances was brown, dark brown,

blackish, light brown with grayish on the tube-

end, brownish-gray, dark brown, blackish gray

with blackish on the tube-end, with hard to soft

textures. In H. planifrons, the color of the nest

entrance tunnel was light brown with a hard

texture (Fig. 2). The nest entrance opening of

H. planifrons was 5.5 cm in height and 7.5 cm

in wide, while in T. sapiens ranged 0.6-3 cm

in height and 0-3.5 cm in wide (Table 2). The

differences in the diameter of the nest entrance

opening in T. sapiens and H. planifrons may

be due to differences in body size, colony size,

and nest age. In Dosay-Kausei villages, we

found the vertically elongated nest entrance of

H. planifrons with characteristics was 2.0 cm in

width and 5.5 cm in height, light brown color,

7.5 cm of funnel length, and no ornamenta-

tion (Fig. 2) with a thick and hard-texture in

tube-end, respectively and the nest height from

ground was 162 cm (0-325 cm) (Table 2).

Nest architecture: The brood cells of T.

sapiens in three villages (four colonies) were

a vertically cluster, horizontally cluster, and

semi-cluster. The brood cells were connected

to other cells by lamellate pillar structures (Fig.

3, Table 3). The brood cells were oval in shape

Fig. 2. Types of nesting entrance opening of T. sapiens in Maribu, Dosay, and Kemiri villages. A.-D. elongated horizontal

with dark brown, blackish and slightly blackish gray on tube-end. E.-J. oval with brown, blackish brown, light brown on

tube end, blackish gray, and brownish black. K.-L. round with dark brown. M.-N. irregular with dark brown and brownish

gray. O.-R. ellipses with brown, dark brown, and blackish gray. S. Elongated vertically with dark brown. T. elongated

vertical with light brown hard texture in H. planifrons. Scale = 1 cm).

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e51166, enero-diciembre 2023 (Publicado May. 16, 2023)

TABLE 3

Nest architecture of T. sapiens and H. planifrons (Modification of Wille & Michener, 1973; Dollin, et al., 1997).

Atribut T. sapiens H. (P. ) planifrons

Maribu Village Kemiri Village Dosay Village

Nest characteristics Colony 2 Colony 5 Colony 10 Colony 22 Colony 13

Entrance tunnel shape horizontal wide horizontal wide vertical elongated horizontal wide vertical elongated

Entrance width (cm) 1.6 2 0.8 2.1 2

Entrance height (cm) 0.6 1 2.6 1 5.5

Funnel length (cm) 1 0 4.7 3.5 7.5

Funnel texture hard hard hard hard hard

Pollen pot space length (cm) 7.5 11.9 12.3 11.2 73

Pollen pot diameter (cm) 0.6-0.8 0.6-1 0.5-0.9 0.8-1.6 5.2-6.8

Pollen pot shape irregular* irregular* irregular* irregular* irregular*

Pollen pot color black dark brown brown dark brown black brown

Honey pot space length (cm) 15.5 4.8 11.5 11 30

Honey pot diameter (cm) 0.6-0.9 0.7-0.9 0.7-1.1 0.6-1.2 5.1-6.9

Honey pot shape irregular* irregular* irregular* irregular* irregular*

Honey pot color black dark brown blackish brown dark brown black brown

Length of pollen pot space +

honey pot space (cm)

18 12.3 25.9 22.2 103

Nest space length (cm) 44 18.6 50 52 143

Brood cells

Arrangement vertical cluster semi-cluster vertical cluster horizontal cluster layer vertical comb

Room length (cm) 26.6 13.5 23.3 32.2 40

Cell height (cm) 0.31-0.36 0.26-0.31 0.3-0.4 0.39-0.44 1.58-2.22

Cell width (cm) 0.19-0.26 0.17-0.22 0.04-0.05 0.21-0.30 0.55-0.62

Shape oval oval oval oval oval

Color light cream-

blackish brown

light cream-

blackish brown

light cream-

blackish brown

light cream-

blackish brown

cream-

dark brown

Pillar columnar columnar columnar lamellate columnar lamellate

Nesting habitat

Substratum (nesting site) dry wood dry wood fern stem house foundation coconut tree

Height from ground surface (cm) 290 70 91 15 162

* round, oval, and ellipse coincide with each other.

TABLE 2

The average of nest entrance characteristics of stingless bees in Maribu, Dosay, and Kemiri villages of Papua.

Species,

Nest locations

Nest entrance

Width (cm) Height (cm) Tube length (cm) Height from

ground surface (cm)

M R SD M R SD M R SD M R SD

T. sapiens

Maribu (n=11) 1.43a0.6-2.4 0.54 1.54a0.6-2.9 0.75 1.24a0-4.7 1.54 118.6ab 0-325 120

Dosay (n=5) 2.14a1.4-2.8 0.53 1.76a1.1-3 0.74 2.02a0.2-3.4 1.24 9.3a1-25 10.2

Kemiri (n=4) 1.92a0.8-2.9 0.86 1.27a0.7-2.5 0.82 1.9a0.1-3.5 1.75 27a8-63 24.6

H. planifrons

Dosay (n=1) 2 2 0 5.5 5.5 0 7.5 7.5 0 162 162 0

Note: M=Mean; R=Range; SD=Standard deviation; Different letters in the same column indicate a significant difference

between the means based on the t-test.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e51166, enero-diciembre 2023 (Publicado May. 16, 2023)

and light brown in color (brood cells with adult

larvae) and dark brown (brood cells with young

larvae). The pollen pots of T. sapiens were

irregular (0.5-1.1 cm in width and 7.5-12.3 cm

in length) with brown, dark brown, and black

colors. The honey pots had an irregular shape

(0.6-1.2 cm in width) and were dark brown,

blackish, and black. The nest heights of T.

sapiens from ground level ranged from 15 to

290 cm (Table 3).

In Dosay-Kausei village, the brood cells

of H. planifrons had a vertically elongated with

a layered vertical comb with a lamellate pillar

structure that connect it to the tree trunk. The

brood cells of the species was oval in shape,

light brown (old cells) and dark brown (young

cells), with height ranging from 1.58-2.22 cm

and a width of 0.55-0.62 cm (Fig. 3). The pol-

len pots were irregular, while the honey pots

were an irregular shape with blackish (Table 3).

DISCUSSION

Morphologically, stingless bees are dif-

ficult to identify (Sakagami & Inoue, 1985).

The current study identified two species as

Tetragonula sapiens (Australasian stingless

bee) and Heterotrigona (Platytrigona) plani-

frons (Papuasian stingless bee). This study

added a new distribution record for T. sapiens

and H. planifrons in Papua, specifically in the

CMNR area of Jayapura Regency. Previous

studies reported T. sapiens was distributed in

the Solomon islands and Papua New Guinea

(Dollin et al., 1997), while H. planifrons has

been reported from the Central Mamberamo

District, Bovendigul District, Merauke Dis-

trict, Manokwari of Papua, and in the Bulolo

District, Morobe, and Oro Provinces of Papua

New Guinea (Engel, 2019). Previous studies

in Papua reported four species of Tetragonula,

namely T. fuscobalteata (Cameron, 1908), T.

sapiens (Cockerell, 1911) (Dollin et al., 1997),

T. clypearis (Friese, 1908), and T. biroi (Friese,

1898) (The Papua Insect Foundation, 2008),

and four species of Heterotrigona, namely

H. planifrons (Smith, 1865), H. lamingto-

nia (Cockerell, 1929), H. flaviventris (Friese,

1908) and H. keyensis (Friese, 1901) (Engel et

al., 2019; Engel, 2019).

The nesting sites of stingless bee are usu-

ally in a tree cavity, soil, and hollow walls

(Slaa et al., 2000). Some species, such as of the

genera Tetragonula, Tetragonisca, Partamona,

Paratrigona, and Plebeia build nests on tree

trunks or solid substrates (Roubik, 2006). In

this study, we found that the nesting site of

T. sapiens was dominatly in house founda-

tion (9 colonies), followed by coconut trees (4

colonies), house walls (3 colonies), dry wood

(2 colonies), fern trees (2 colonies), and soil

(1 colony). In contrast, the nesting site of H.

planifrons was in a coconut tree cavities. The

limited nesting sites of H. planifrons (1 colony

found) may be a result of burning the nest due

to their aggressive biting.

The nests of aggressive stingless bees

are usually found individually in specific tree

Fig. 3. The internal nest architecture of four colonies of T. sapiens in Maribu, Dosay, and Kemiri villages, Papau: A.-B.

vertical cluster, C. semi-cluster, D. horizontal cluster, and E. layered vertical comb in H. planifrons (insert for detail).

ne=nest entrance, bc=brood cells, pc=pollen cells, hc=honey cells, lp=lamellate pillars. Scale=1cm.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e51166, enero-diciembre 2023 (Publicado May. 16, 2023)

trunks, such as palm trees (Roubik, 2006).

Meanwhile, the nests of non-aggressive sting-

less bee colonies are generally close to other

stingless bee species (Roubik, 1983). The cur-

rent study found more colonies building nests

close-together in sap-producing trees, such

as Psidium or cashew (Anacardium occiden-

tale: Anacardiaceae), merbau (Intsia bijuga:

Fabaceae), mango (Mangifera indica: Anacar-

diaceae), jackfruit (Artocarpus heterophyllus:

Moraceae), and wood milk (Alstonia scholaris:

Apocynaceae). Resin in the nest entrance pro-

tects the colony from predators (Roubik, 2006).

The texture of the nest entrance is related to the

various resins collected from gummy plants.

Variation in the nest entrance is influenced by

nest age, genetics, predators, parasites, sym-

bionts, rain, wind, and light (Sakagami et al.,

1983; Wille & Michener, 1973). This study

found a pronged and a branched nest entrance

of T. sapiens in Maribu village. A branching

nest entrance has also been found in Lepido-

trigona ventralis (Smith, 1857) (in Thailand),

Lepidotrigona, Plebeia, Scaptotrigona, and

Tetragona (Roubik, 2006).

The results showed that the nest entrance

of T. sapiens was a short funnel, whereas in

H. planifrons it was an elongated. Unfortunat-

ley, we only found one colony of the species

and no variation of nest entrance character-

istics was shown. We suppost that there are

variation of the nest entrance characteristics

in the species. So, more exploration in a wide

areas in Papua should be done in the future.

Nest entrances with a short funnel are usu-

ally found in stingless bees of the Neotropics

(T. cilipes) and Asia (T. collina). Similarly, an

elongated nest entrance was also found in two

Asian stingless bees with aggressive behavior,

namely H. canifrons and H. itama (Sakagami

et al., 1983). The color and texture of the nest

entrances of T. sapiens varied, such as brown,

dark brown, blackish, cream (grayish on the

tube-end), brownish, brownish-black, black-

ish gray (blackish on the tube-end), with a

hard and soft texture. In H. planifrons (Smith,

1865), the color of nest entrance is light brown

with a hard texture. The color variation of

the nest entrance may be due to the different

resin sources (Sakagami et al., 1983; Wille &

Michener, 1973). Nest entrance variation in

T. sapiens may be related to polyphenism, in

which two or more phenotypes result from the

same genotype (Mayr, 1963). Polyphenism is

influenced by environmental conditions that

cause phenotypic preferences in neurochemical

and hormonal pathways related to nest-building

behavior (Simpson et al., 2011).

The brood cells of T. sapiens are con-

nected by a columnar lamellate pillar (Fig. 3,

Table 3), oval shape and light- and dark-brown,

while the pollen and honey pots were irregular,

probably an adaptation to substrate types. The

modification of nest architecture in stingless

bees is closely related to the adaptability of a

species to its environment. Species with high

intraspecific and phenotypic variations are usu-

ally more adaptive to environmental changes

than endemic species with low intraspecific

variations and limited distribution in certain

geographic area (Cohet et al., 1980; Lavergne

& Molofsky, 2007). In this study, T. sapiens

found in all sampling sites with various nest

entrance characters and able to live at varying

altitudes (107-364 m.a.s.l). In contrast, H. plan-

ifrons showed a limited distribution, at just one

sampling site at an altitude of 127 m.a.s.l. The

observations and interviews with local com-

munities at West Sentani to Sentani districts

showed H. planifrons had a limited distribution

and was only found in the Dosay village.

Ethical statement: the authors declare

that they all agree with this publication and

made significant contributions; that there is no

conflict 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 acknowledge-

ments section. A signed document has been

filed in the journal archives.

ACKNOWLEDGMENTS

We acknowledge to PT Freeport Indonesia

for funding this research and head of Cycloops

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e51166, enero-diciembre 2023 (Publicado May. 16, 2023)

Mountains Nature Reserve for allowing the

authors to conduct this research. We also thank

Laboratory of Entomology, Research Can-

ter for Ecology and Ethnobiology, National

Research and Innovation Agency (BRIN) for

verified stingless bee specimens.

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