Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

Diversity of centipedes and millipedes (Arthropoda: Myriapoda)

in tropical dry forest fragments in Bolívar, Colombia

Martín Carrillo-Pallares1; https://orcid.org/0000-0003-3406-1699

Wilder Zapata-M.1; https://orcid.org/0000-0003-2011-6943

Daniela Martínez-Torres2; https://orcid.org/0000-0001-8865-927X

Gabriel R. Navas-S.3*; https://orcid.org/0000-0001-9554-6345

Adriana Bermúdez-Tobón1; https://orcid.org/0000-0002-5247-4561

1. Grupo de Investigación en Biología Descriptiva y Aplicada, Programa de Biología, Universidad de Cartagena, Campus

San Pablo, Cartagena de Indias, Bolívar, Colombia; m-carrillo1996@hotmail.com; wilder214_@hotmail.com; abermu-

dezt@unicartagena.edu.co

2. Grupo de Investigación en Aracnología y Miriapodología, Instituto de Ciencias Naturales, Sede Bogotá, Universidad

Nacional de Colombia, Bogotá D.C., Colombia; sdmartinezt@unal.edu.co

3. Grupo de Investigación Hidrobiología, Programa de Biología, Universidad de Cartagena, Campus San Pablo,

Cartagena de Indias, Bolívar, Colombia; gnavass@unicartagena.edu.co (*Correspondencia).

Received 20-II-2025. Corrected 10-V-2025. Accepted 12-VIII-2025.

ABSTRACT

Introduction: Tropical dry forests are among the most endangered Neotropical ecosystems, characterized

by high biodiversity and endemism but increasingly threatened by anthropogenic activities. In Colombia,

particularly in the Caribbean region, these forests have been extensively fragmented. Despite their ecological

significance, studies on myriapod communities remain scarce. Myriapods play an essential role in soil dynamics

and nutrient cycling. However, their diversity and ecological associations in Bolívar’s tropical dry forests remain

poorly understood.

Objective: To assess the diversity and distribution of centipedes (Chilopoda) and millipedes (Diplopoda) in

four tropical dry forest fragments within Bolívar and examine their relationships with environmental variables.

Methods: Fieldwork was conducted in October and November 2020 across four forest fragments in Turbaco,

San Jacinto, Magangué, and El Carmen. Specimens were collected using free search, soil monoliths, pitfall traps,

and Winkler traps in 50 × 50 m plots. Taxonomic identifications and ecological analyses were performed to

assess the diversity and structure of the studied communities. Ecological indices were analyzed, and ANOSIM

was used to contrast differences between locations, NMDS was used to visualize patterns of dissimilarity, and

Canonical Correspondence Analysis (CCA) was used to evaluate the relationship between species and environ-

mental variables.

Results: A total of 1 106 individuals were recorded, comprising 8 orders, 15 families, and 32 morphospecies.

Diplopoda exhibited greater richness and abundance than Chilopoda. Polydesmida and Spirostreptida were the

dominant millipede orders, while Geophilomorpha was the most abundant centipede order. Species richness was

highest in San Jacinto and Turbaco, while Magangué and El Carmen exhibited lower diversity. The CCA revealed

that soil pH, temperature, and humidity influenced species distribution, reinforcing the role of myriapods as

bioindicators.

Conclusion: This study provides the first systematic assessment of myriapod diversity in Bolívar’s tropical dry

forests, emphasizing their ecological significance and the urgent need for conservation efforts.

Key words: Chilopoda; Diplopoda; ecology; environmental variables; Neotropic; sampling methods.

https://doi.org/10.15517/10.15517/9w79j526

TERRESTRIAL ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

INTRODUCTION

A Tropical Dry Forest are ecosystems dis-

tributed between 0 and 1 000 m in altitude,

characterized by temperatures ranging from

25 to 38 °C (Murphy & Lugo, 1986) and annu-

al precipitation between 250 and 2 000 mm.

These forests experience pronounced seasonal

droughts lasting between four and six months

(Grainger, 1996; Mayaux et al., 2005; Miles

et al., 2006; Pennington, 2006). Due to their

unique structural and ecological characteristics,

these forests exhibit high levels of alpha diver-

sity and endemism, with species displaying a

wide range of adaptive strategies to withstand

extreme drought conditions and seasonal cli-

matic fluctuations (Alvarado-Solano & Otero,

2015; Dryflor et al., 2016; Zapata et al., 2023).

In Colombia, the original extent of Tropical

Dry Forest was estimated to be approximately

80000 km² (Etter, 1993). Later assessments

by Instituto de Hidrología, Meteorología y

Estudios Ambientales et al., (2007) estimated

a current coverage of 76 581 km², primarily

distributed across six major regions: The Carib-

bean, the inter-Andean valleys of the Cauca and

Magdalena rivers, the North Andean region

(Santander and Norte de Santander), the Patía

Valley, and the Llanos (Arauca and Vichada).

This represents roughly 7 % of the national ter-

ritory (Díaz, 2006). However, these forests have

been extensively fragmented by anthropogenic

pressures, with 97 % of their original coverage

impacted by activities such as livestock farm-

ing, mining, deforestation, and monoculture

agriculture, accelerating the rate of habitat deg-

radation (Álvarez Hincapié et al., 2021; García-

Herrera et al., 2019; Miles et al., 2006).

Among Colombia’s remaining Tropical

Dry Forests, the Caribbean region contains the

largest remnants, covering 1 355 km² (Rodrí-

guez et al., 2012). However, most of these

RESUMEN

Diversidad de ciempiés y milpiés (Arthropoda: Myriapoda) en

fragmentos de bosque seco tropical en Bolívar, Colombia

Introducción: Los bosques secos tropicales se encuentran entre los ecosistemas neotropicales más amenazados,

caracterizados por una gran biodiversidad y endemismo, pero cada vez más amenazados por las actividades

antropogénicas. En Colombia, particularmente en la región del Caribe, estos bosques se han fragmentado amplia-

mente. A pesar de su importancia ecológica, los estudios sobre las comunidades de miriápodos siguen siendo

escasos. Los miriápodos desempeñan un papel esencial en la dinámica del suelo y el ciclo de los nutrientes. Sin

embargo, su diversidad y sus asociaciones ecológicas en los bosques secos tropicales de Bolívar siguen siendo

poco conocidas.

Objetivo: Evaluar la diversidad y distribución de ciempiés (Chilopoda) y milpiés (Diplopoda) en fragmentos de

cuatro bosques secos tropicales de Bolívar y examinar su relación con variables ambientales.

Métodos: El trabajo de campo se llevó a cabo en octubre y noviembre de 2020 en cuatro fragmentos de bosques

en Turbaco, San Jacinto, Magangué y El Carmen. Las muestras se recolectaron mediante búsqueda libre, mono-

litos de suelo, trampas de caída y trampas Winkler, en parcelas de 50 × 50 m. Se analizaron índices ecológicos y

se utilizó ANOSIM para contrastar diferencias entre localidades, NMDS para visualizar patrones de disimilitud

y análisis canónico de correspondencias (CCA) para evaluar la relación entre especies y variables ambientales.

Resultados: Se registraron en total 1 106 individuos, que comprendían 8 órdenes, 15 familias y 32 morfoespecies.

Los diplópodos exhibieron mayor riqueza y abundancia que los quilópodos. Polydesmida y Spirostreptida fueron

los órdenes de milpiés dominantes, mientras que Geophilomorpha fue el orden de ciempiés más abundante. La

riqueza de especies fue mayor en San Jacinto y Turbaco, mientras que Magangué y El Carmen mostraron una

menor diversidad. El CCA reveló que el pH, la temperatura y la humedad del suelo influyeron en la distribución

de las especies, lo que refuerza el papel de los miriápodos como bioindicadores.

Conclusión: Este estudio proporciona la primera evaluación sistemática de la diversidad de miriápodos en los

bosques secos tropicales de Bolívar, enfatizando su importancia ecológica y la urgente necesidad de esfuerzos de

conservación.

Palabras clave: Chilopoda; Diplopoda; ecología, variables ambientales, neotrópico; métodos de muestreo

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

forests exist as highly fragmented and heavily

altered patches (Díaz, 2006). The Department

of Bolívar is a key area for forest conservation,

comprising 4.1 % of the national territory and

ranking third in Tropical Dry Forest coverage,

following Atlántico (5.7 %) and Cesar (4.9 %)

(Pizano et al., 2014). Given the ecological sig-

nificance of these relictual forests, it is essential

to investigate arthropod diversity to better

understand ecosystem dynamics and inform

future conservation and restoration strategies

(Bello et al., 2014; Ramírez, 2018).

Among arthropods, Myriapods (subphy-

lum Myriapoda) are particularly relevant as

soil ecosystem engineers. This group comprises

four classes: Chilopoda (centipedes), Diplop-

oda (millipedes), Symphyla, and Pauropoda

(Edgecombe, 2011; Shelley, 2002). Myriapods

contribute significantly to soil health by increas-

ing porosity and water retention, promoting

nutrient cycling, and accelerating decomposi-

tion processes (Bueno-Villegas, 2012). Due

to their close association with microhabitat

conditions, myriapods are often used as bio-

indicators for evaluating soil quality, environ-

mental change, and ecosystem integrity (Adis

et al., 1996; Druce et al., 2004). Additionally,

centipedes serve as biological control agents,

preying on soil mesofauna, macroinvertebrates,

and occasionally small vertebrates (Lewis, 1981;

Molinari et al., 2005). They also represent an

important food source for birds, mammals,

reptiles, insects, and arachnids (Cantú-Salazar

et al., 2005; Cupul-Magaña, 2014).

Despite their ecological importance, myr-

iapod diversity in Colombian Tropical Dry

Forests remains poorly studied. Most research

has focused on the Andean region, where four

orders, seven families, twelve genera, and forty

species of Chilopoda (Chagas-Jr et al., 2014;

Prado-Sepúlveda et al., 2016), and seven orders,

20 families, 76 genera, and approximately 215

species of Diplopoda have been documented

(Castillo-Ávila et al., 2015; Martínez-Torres &

Flórez, 2015). In the Caribbean region, early

contributions were made by Carl (1914) and

Chamberlin (1921), who described several

myriapod species in Magdalena and Atlántico.

However, in Bolívar, myriapod records remain

scarce, with only a few inventories present-

ed at scientific meetings. These inventories

report four orders, six families, and nine mor-

phospecies of Chilopoda, along with seven

orders, eleven families, and over 20 morpho-

species of Diplopoda (Carrillo-Pallares et al.,

2019; Sociedad Colombiana de Entomología

[SOCOLEN], 2016).

Given these knowledge gaps, this study

aimed to assess the diversity of centipedes

and millipedes in various Tropical Dry Forest

fragments within the Department of Bolívar.

Additionally, we investigate how species from

Chilopoda and Diplopoda correlate with envi-

ronmental variables, including ambient humid-

ity, ambient temperature, soil temperature,

and soil pH. Our findings contribute to the

taxonomic and ecological understanding of

myriapods in Bolívar and Colombia, provid-

ing baseline data for biodiversity conservation

and management.

MATERIALS AND METHODS

Study Area: The Department of Bolívar

is located in Northern Colombia, within the

Caribbean plain region (Fig. 1). It covers 25 978

km², with altitudes ranging from sea level to

2 700 m above sea level (MASL). Bolívar is

positioned between 07º00’03’’-10º48’37’’ N &

73º45’15’’-75º42’18’’ W, sharing borders with

the Caribbean Sea and the Department of

Atlántico to the North, the Magdalena River

to the East, the departments of Santander and

Antioquia to the South, and the departments of

Antioquia, Córdoba, Sucre, and the Caribbean

Sea to the West. The department accounts for

approximately 20 % of the Caribbean region’s

total area and 2.3 % of Colombia’s national ter-

ritory (Galvis, 2009). The region experiences an

average temperature ranging from 19 to 30 °C,

with annual precipitation between 911 and

1 157 mm. Bolívar comprises 45 municipalities

and 348 townships, with a mosaic of land uses,

including Tropical Dry Forest fragments at dif-

ferent successional stages, shrublands, gallery

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

forests, and deforested areas due to livestock

and agricultural expansion (Pizano et al., 2014).

Field Sampling: Was conducted between

October and November 2020 at four sites: Tur-

baco, San Jacinto, Magangué, and El Carmen

(Table 1). Each location contained remnants of

these forests at varying degrees of conservation

(Fig. 2). The sampling design followed the

COBRA (Conservation-Oriented Rapid Biodi-

versity Assessment) protocol (Cardoso et al.,

2009; Malumbres-Olarte et al., 2017), incor-

porating modifications by Morales (2019) and

Galvis (2019).

At each site, a 50 × 50 m plot was estab-

lished, where myriapod specimens were

Fig. 1. Sampling locations in the department of Bolívar, Colombia.

Tabl e 1

General characteristics of the conserved patches of tropical dry forest sampled for myriapods in the Bolívar department,

Colombia.

Locality,

Municipality

Coordinates Elevation

(MASL)

Description

Cañaveral,

Turbaco

10°22’07.1” N-75°21’30.1” W 104 It is located near a water source and remains minimally

disturbed by anthropogenic activities.

Brasilar,

San Jacinto

9°54’33.1” N-75°11’06.1” W 435 It is surrounded by mountainous areas and located near a

water source.

Henequén,

Magangué

9°12’29.9” N-74°53’45.9” W 56 In the vicinity of the flood zone, surrounding areas are

disturbed by anthropogenic activities such as livestock

farming, access roads, monocultures, and semi-urban areas.

Tolemaida,

El Carmen

9°38’07.4” N-75°07’29.3” W 253 Surrounding areas, affected by anthropogenic activities such

as armed conflict, monocultures, and livestock, are in a state

of succession within mountainous areas. There are no nearby

water sources.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

collected using the following techniques: Free

Search: Four researchers conducted active

manual searches for one hour during both day-

time and nighttime shifts. Searches focused on

decaying logs, tree trunks, leaf litter, and other

microhabitats known to harbor myriapods.

Soil Monolith Extraction: A metal sampling

frame (25 cm × 25 cm × 10 cm) was used to

extract ten soil monoliths per sampling day.

These were randomly selected within the plot.

The extracted soil was spread on a white cloth

and examined under artificial light to locate

specimens. Winkler Traps: Twelve 1 m² leaf

litter samples were randomly collected within

each plot. The samples were processed using

Winkler extractors, placed in Winkler bags, and

left at room temperature for three days before

analysis. Pitfall Traps: A total of 48 pitfall traps

Fig. 2. Locations where myriapods were captured in the department of Bolívar, Colombia. a. Turbaco. b. San Jacinto. c.

Magangué. d. El Carmen.

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

(disposable 10 cm diameter plastic cups) were

placed in groups of four, evenly distributed

along the perimeter of each quadrant. Traps

were covered with disposable plates positioned

5 cm above ground level to prevent rainwater

ingress. The traps were filled with propylene

glycol and remained in place for 15 days before

specimen retrieval.

To assess habitat conditions, we recorded

four environmental variables at each sampling

site: Relative humidity (%) and ambient tem-

perature (°C) were measured using a UNI-T

UT333 digital meter. Soil temperature (°C) was

recorded every three hours using a digital soil

thermometer. Soil pH was measured with a

portable pH meter.

Laboratory Processing and Taxonomic

Identification: Collected specimens were fixed

and preserved in 70 % ethanol, labeled, and

transported to the University of Cartagena

laboratories. Taxonomic identification was per-

formed to the lowest possible level, based on

morphological traits. Identification followed

standard taxonomic keys, including those by

Hoffman et al. (2002), Cupul-Magaña (2012),

Chagas-Jr et al. (2014), and Prado-Sepúlveda

et al. (2016). Additionally, expert taxonomists

provided verification.

For species confirmation, collected speci-

mens were compared with reference collections

housed at the Institute of Natural Sciences

(ICN), National University of Colombia. The

studied specimens are currently stored at the

University of Cartagena research laboratories

and the ICN collections.

For ecological analyses, individuals were

categorized into morphospecies, ensuring that

each morphospecies included at least one adult

individual. Sex differentiation was not consid-

ered in abundance estimates.

Data Analysis: To evaluate sampling

effectiveness, we generated species accumula-

tion curves using non-parametric estimators

(Chao1 and ACE) based on abundance data.

These models incorporated: Singletons (spe-

cies represented by a single individual) and

Doubletons (species represented by two indi-

viduals). Species accumulation curves were

constructed using R Studio and EstimateS v. 9.1

software (Colwell et al., 2012; Colwell, 2013).

To compare alpha diversity among sites,

we calculated Hill numbers at three diversity

orders (Moreno et al., 2011): Species richness

(q = 0): Number of observed species. Shannon

diversity (q = 1): The exponent of the Shannon-

Wiener index, weighting species by their rela-

tive abundance. Simpson diversity (q = 2): The

inverse of the Simpson index, emphasizing

dominant species. Confidence intervals were

computed for each diversity order using Spade

software and R Studio (Chao et al., 2016).

To assess differences among sites, we per-

formed an Analysis of Similarities (ANOSIM)

with 9 999 permutations and a significance

threshold of p < 0.05. The null hypothesis

(H₀) stated that there were no differences

among sites.

Additionally, we conducted a classification

and ordination analysis using the Chao-Jaccard

index to visualize grouping patterns among

study sites. This was implemented in PAST v.

3.01 and R Studio (Chao et al., 2016; Hammer

et al., 2001).

To identify morphospecies responsible for

site dissimilarities, we performed a non-metric

multidimensional scaling (NMDS) analysis

using species abundance data.

Finally, to explore the potential influence

of environmental variables (ambient humidity,

ambient temperature, soil temperature, and

soil pH) on morphospecies abundance, we

conducted a Canonical Correspondence Analy-

sis (CCA). Morphotypes with fewer than 10

individuals were excluded from this analysis to

minimize bias.

RESULTS

A total of 1 106 myriapod individuals were

collected during the study, comprising 326

individuals of Chilopoda and 780 individuals

of Diplopoda. The centipedes were distributed

across three orders, six families, and 11 mor-

phospecies, while the millipedes represented

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

five orders, nine families, and 21 morphospe-

cies, indicating greater richness and abundance

within the Diplopoda (Table 2).

Among the Chilopoda, the order Geophi-

lomorpha exhibited the highest abundance,

with 162 individuals, representing 14.6 % of the

total collected specimens. This was followed

closely by Scolopendromorpha, which had 158

individuals (14.3 %). In contrast, Lithobio-

morpha was the least represented, with only

Tabl e 2

Taxonomic composition and abundances of centipedes and millipedes in the evaluated localities’ Tropical Dry Forest

fragments, using different collection techniques.

Class Order Family Morpho-

species

Abundance Techniques Total

Tur SaJ Mag Car

Chilopoda Geophilomorpha Geophilidae Geo1 8 17 8 8 Free Search

Soil Monolith

Pitfall Traps

Winkler Traps

  Oryidae Ory1 0500Soil Monolith

Pitfall Traps

Winkler Traps

   Ory2 17 54 10 30 Free Search

Winkler Traps

111

  Schendylidae Sch1 0500Free Search

Soil Monolith

Winkler Traps

Lithobiomorpha Henicopidae Hen1 1050Free Search 6

Scolopendromorpha Scolopendridae Sco1 0 1 1 0 Free Search

Soil Monolith

   Sco2 0 0 0 9 Free Search 9

   Sco3 6 1 7 16 Free Search

Soil Monolith

Winkler Traps

   Sco4 5 2 0 5 Free Search 12

  Scolopocryptopidae Scp1 15 60 4 25 Free Search

Soil Monolith

Winkler Traps

104

   Scp2 1 0 0 0 Free Search 1

Diplopoda Polydesmida Chelodesmidae Che1 22 116 0 8 Free Search

Winkler Traps

146

   Che2 3000Free Search

Soil Monolith

   Che3 23 4 26 2 Free Search

Soil Monolith

Pitfall Traps

   Che4 3260Free Search

Winkler Traps

  Cyrtodesmidae Cyr1 0500Soil Monolith

Winkler Traps

   Cyr2 0200Winkler Traps 2

  Fuhrmannodesmidae Fhu1 105 18 0 0 Soil Monolith 123

   Fhu2 39 25 0 0 Free Search

Soil Monolith

Pitfall Traps

Winkler Traps

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

six specimens, accounting for less than 1 %

of the total abundance. Within the Chilopoda

families, Oryidae was the most abundant, com-

prising 10.5 % of the total individuals, while

Schendylidae was the least abundant, contrib-

uting less than 1 %.

For Diplopoda, the order Polydesmida

was the most dominant, with 473 individu-

als, representing 42.7 % of the total collect-

ed specimens. The second most abundant

order was Spirostreptida, with 184 individu-

als (16.5 %), while Siphonophorida exhibited

the lowest abundance, with only 17 individu-

als (2 %). Regarding family-level composition,

Fuhrmannodesmidae was the most abundant,

comprising 21.9 % of the total individuals, fol-

lowed by Chelodesmidae (19.4 %). The families

Pyrgodesmidae and Cyrtodesmidae were the

least represented, each accounting for less than

1 % of the total Diplopoda individuals.

The species accumulation curves (Fig. 3)

indicated a sampling efficiency of 97 %, as

estimated by ACE and Chao1 non-parametric

estimators. The proportion of Singletons and

Doubletons (species with only one or two

individuals recorded) accounted for approxi-

mately 10 % of the total sample, confirming

the adequacy and representativeness of the

sampling effort. The negative trend in the

Singletons curve suggested a decline in newly

Class Order Family Morpho-

species

Abundance Techniques Total

Tur SaJ Mag Car

   Fhu3 0030Free Search

Pitfall Traps

   Fhu4 0 53 0 0 Free Search

Soil Monolith

Winkler Traps

  Pyrgodesmidae Pyr1 8 0 0 0 Free Search 8

Polyxenida Lophoproctidae Pol1 5735Free Search

Soil Monolith

Winkler Traps

   Pol2 0900Free Search

Soil Monolith

Siphonophorida Siphonophoridae Sip1 6600Free Search

Soil Monolith

   Sip2 0100Free Search 1

   Sip3 0040Free Search

Soil Monolith

Winkler Traps

 Spirobolida Rhinocricidae Rhi1 3 0 0 74 Free Search

Soil Monolith

Pitfall Traps

Winkler Traps

Spirostreptida Pseudonannolenidae Pse1 1 1 95 12 Free Search

Soil Monolith

Winkler Traps

109

   Pse2 0 0 61 0 Free Search

Soil Monolith

  Spirostreptidae Spt2 7000Free Search 7

   Spt3 3310Free Search

Soil Monolith

  281 397 234 194 - 1 106

* Turbaco (Tur), San Jacinto (SaJ), Magangué (Mag), and El Carmen (Car).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

detected species, reinforcing the completeness

of the myriapod assemblages sampled within

the Tropical Dry Forest fragments of Bolívar.

The diversity profiles (Fig. 4) revealed that

San Jacinto and Turbaco exhibited the highest

effective species richness across all diversity

orders. In terms of species richness (q0), El

Carmen had the lowest number of species,

with 11 recorded morphospecies, followed by

Magangué with 14, Turbaco with 20, and San

Jacinto with 22. When considering order 1

diversity (q1), which accounts for common spe-

cies, San Jacinto had the highest effective species

count, reaching 9.42, making it 1.01 times more

diverse than Turbaco, 1.34 times more diverse

than El Carmen, and 1.59 times more diverse

than Magangué. In terms of species dominance

(q2), which emphasizes dominant species,

Magangué had the lowest diversity value, with

only 3.95 effective species, representing 60.9 %

of San Jacinto’s diversity, 72.0 % of Turbaco’s,

and 79.7 % of El Carmen’s diversity.

A notable variation was observed between

Magangué and El Carmen, where Magangué

exhibited greater species richness, yet El Car-

men had a higher proportion of common

and dominant species. The observed diver-

sity trends closely aligned with expected values,

confirming the robustness of the sampling

effort. However, broader confidence intervals

were noted, particularly for species richness

(q0), indicating potential underrepresentation

of rare species in the dataset.

The ANOSIM analysis detected significant

differences among the study sites (R = 0.8208, p

= 0.0001), suggesting that each locality harbors

distinct myriapod assemblages. The classifica-

tion (Fig. 5A) and NMDS ordination analyses

(Fig. 5B) further identified three distinct clus-

ters. The first group comprised samples from

San Jacinto and Turbaco, which exhibited the

highest similarity in species composition, likely

due to minimal anthropogenic disturbance in

these areas. The second group corresponded to

El Carmen, which exhibited moderate dissimi-

larity from the previous group. The third and

most distinct cluster was formed by Magangué,

which showed the greatest divergence from the

other study sites, suggesting that its myriapod

assemblage was the most dissimilar.

Analysis of environmental factors revealed

that Turbaco and Magangué recorded the high-

est air temperatures, reaching 28.1 and 27.4 °C,

respectively, while San Jacinto and El Carmen

exhibited lower temperature values of 25.6 and

Fig. 3. Accumulation curves of myriapod species richness from Tropical Dry Forest fragments from the department of

Bolívar, Colombia.

Fig. 4. Diversity profiles (Diversity orders q0 for species

richness, q1 for common species, q2 for dominant species)

observed and expected for myriapods from Tropical Dry

Forest. fragments in the department of Bolívar. Confidence

intervals are presented as the projections of each curve on

the graph.

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

26.0 °C. Similarly, soil temperature followed

the same trend, with Turbaco and Magangué

reaching 27.9 and 27.6 °C, and San Jacinto and

El Carmen showing lower values. In terms of

relative humidity, Turbaco exhibited the highest

percentage, at 93.6 %, followed by El Carmen

at 92.8 %, San Jacinto at 90.3 %, and Magangué

at 88.0 % (Fig. 6A). Soil pH varied across sites,

with Turbaco and Magangué displaying acidic

soils with pH values of 6.9 and 6.6, respectively,

whereas El Carmen had a neutral pH of 7.0,

and San Jacinto exhibited the highest pH value

of 7.1 (Fig. 6B).

The Canonical Correspondence Analysis

(CCA) (Fig. 7) identified a strong association

between species distribution and environmen-

tal variables. The first CCA axis explained 64.2

% of the total variance, indicating a strong

environmental influence on myriapod com-

position across sites. The species Pse1, Spt2,

Che3, and Che4 were positively associated with

high air and soil temperatures, particularly in

Magangué, which had the highest recorded

temperatures. This pattern suggests that these

species exhibit a preference for warm environ-

ments. In contrast, environmental humidity

Fig. 5. Classification A. and ordination B. of samples of myriapod assemblages carried out in fragments of Tropical Dry

Forests in the department of Bolívar. Note the formation of three groups: Group 1 (San Jacinto and Turbaco), El Carmen,

and Magangué.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

was positively correlated with El Carmen, rein-

forcing the presence of species associated with

more humid conditions.

Additionally, morphospecies Ory2, Geo1,

and Scp1 exhibited a strong association with soil

pH variations, with their abundance increasing

in response to changes in pH across the four

evaluated localities. These findings highlight

the significant role of environmental variables

in shaping the composition and distribution of

myriapod communities within the Tropical Dry

Forest fragments of Bolívar.

DISCUSSION

Class Chilopoda: In this study, we iden-

tified three of the four Chilopoda orders

previously documented in Colombia within

the department of Bolívar. Geophilomor-

pha emerged as the most predominant order,

encompassing all three families reported for the

country (Foddai et al., 2000; Prado-Sepúlveda

et al., 2016). At the family level, 60 % of those

documented for Colombia were represented

(Chagas-Jr et al., 2014), with Scolopendridae

(Scolopendromorpha) being the most diverse.

This contrasts with previous studies, such as

those conducted by Castillo (2017) and Prado-

Sepúlveda et al. (2016), where Scolopocryptopi-

dae exhibited the highest richness.

Previous research identified the genera

Newportia Gervais, 1 847, and Scolopocryp-

tops Newport, 1 844, along with the species

Scolopocryptops ferrugineus Linnaeus, 1 767,

Fig. 6. Variation of environmental parameters concerning the abundance of each location. A. Ambient humidity, ambient

temperature, and soil temperature, vs abundances. B. pH vs abundance.

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

Rhisyda Wood, 1 862, Rhisyda celeris (Humbert

& Saussure, 1869), Scolopendra arthrorhabdoi-

des Ribaut, 1 912, and Scolopendra gigantea

Linnaeus, 1 758 in Bolívar. Additionally, the

genus Cormocephalus Newport, 1 844 (order

Scolopendromorpha) and Notiphilides Lat-

zel, 1 880 (order Geophilomorpha) have been

recorded for the Colombian Caribbean region

(Chagas-Jr et al., 2014).

Class Diplopoda: Five of the seven Dip-

lopoda orders registered for Colombia were

found in the department of Bolívar, with Poly-

desmida and Spirostreptida being the most

representative. This pattern aligns with findings

from studies in Colombia and the Neotropics,

which highlight the dominance of these orders

in various ecosystems, such as the Otún River

basin (Ruiz-Cobo et al., 2010), the Andean

humid forest of Icononzo, and the Eastern Hills

of Bogotá (Castillo-Ávila et al., 2015; Prado

& Herrera, 2013), as well as the high tropical

rainforest of Los Tuxtlas in Mexico (Bueno &

Rojas, 1999).

At the family level, 69.2 % of the Diplopoda

families documented in Colombia were rep-

resented (Castillo-Ávila et al., 2015; Martínez-

Torres & Flórez, 2015; Ruiz-Cobo et al., 2010).

Additionally, the number of recorded families

has increased compared to prior inventories

conducted in Bolívar and presented at scien-

tific conferences (SOCOLEN, 2016; Carrillo-

Pallares et al., 2019). The new records also

expand upon previously unregistered families

in ICN databases and the Universidad de Carta-

gena collections (consulted in December 2020).

However, despite their presence in neighboring

areas, Aphelidesmidae and Paradoxosomatidae

were absent from this study.

Furthermore, For Paradoxosomatidae,

Hoffman (1999), Pérez-Asso & Pérez-Gelabert

(2001), Shelley & Lehtinen (1998), and Suriel

(2012) suggested that species such as Oxidus

gracilis (Koch, 1847) and Orthomorpha coarc-

tata (De Saussure, 1860) were likely introduced

via the greenhouse plant trade from Asia. The

absence of these species in Bolívar may indicate

environmental or ecological barriers limiting

their establishment in the studied fragments.

Fig. 7. Canonical correspondence analysis (ACC) between the environmental variables (ambient humidity, ambient

temperature, soil pH, and soil temperature) and the abundances of the morphospecies of the evaluated localities.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

Myriapod Diversity in Tropical Dry For-

est Fragments: The species accumulation curve

suggests that the sampling effort successfully

captured a robust representation of myriapods,

supporting the reliability of the dataset for

characterizing Tropical Dry Forest assemblages

in Bolívar. The decreasing trend in Singleton

and Doubleton curves, which eventually inter-

sect, suggests that the probability of discover-

ing additional rare species is low. However,

further sampling in new localities containing

forest fragments would be valuable in assessing

whether diversity patterns remain consistent.

The ANOSIM analysis revealed significant

differences among localities, a result further

supported by classification and ordination anal-

yses (Fig. 5). The similarity observed between

San Jacinto and Turbaco can be attributed to

their low levels of anthropogenic disturbance

and the presence of well-preserved Tropical Dry

Forest remnants. The shared biotic and abiotic

conditions in these locations, including prox-

imity to water sources (Aguilera, 2014; Menco,

2013), may facilitate species coexistence and

interactions, leading to higher local diversity

(González-Montaña et al., 2017; Wardle, 2006).

Conversely, El Carmen and Magangué exhib-

ited greater dissimilarity, likely due to intensive

habitat modification. These areas have been

heavily affected by armed conflict, extensive

livestock farming, monocultures, and semi-

urban expansion (Aguilera, 2014; Daniels-Puel-

lo, 2016). Such disturbances can degrade forest

structure, disrupt ecological interactions, and

lead to biodiversity loss (Rendón-Carmona et

al., 2009). Despite these transformations, some

morphospecies were shared across multiple

sites, suggesting that certain myriapods possess

ecological plasticity, allowing them to adapt to

fragmented forest environments.

The CCA analysis showed that the mor-

phospecies Ory1 and Geo1 (order Geophi-

lomorpha) were associated with soil pH

variations in San Jacinto. Their abundance

increased in soils with higher pH values, likely

reflecting their preference for microhabitats

within surface soil layers, where they construct

galleries. Since soil chemistry can directly

influence species distribution, this result aligns

with previous studies on macroinvertebrate

communities, where chilopod densities were

linked to soil pH, nitrogen, and carbon content

(Díaz et al., 2014). Given these relationships,

further investigations into soil properties would

enhance our understanding of myriapods as

bioindicators of soil health (Rendón et al., 2011;

Ruiz-Cobo et al., 2010).

Interestingly, the morphospecies Fhu1

(order Polydesmida) and Rhy1 (order Spi-

robolida) did not show significant variations

in abundance across environmental gradients.

However, numerous individuals of these spe-

cies were observed engaging in courtship and

copulation behaviors during the study. Similar

patterns have been reported in the Andean

humid forest of Icononzo, Tolima, where Spi-

rostreptidae morphospecies displayed compa-

rable reproductive behaviors (Castillo, 2017).

Conservation Implications: Tropical Dry

Forests are recognized for their high arthropod

richness and serve as valuable ecological mod-

els for assessing biotic and abiotic interactions

(Janzen, 1980; Janzen, 1988). In Colombia,

previous biodiversity assessments have focused

primarily on insect communities, such as ants

(Formicidae) (Chacón et al., 2012; Simanca-

Fontalvo et al., 2013), dung beetles (Scarabaei-

nae) (Rangel-Acosta et al., 2016), and certain

spider groups (Escorcia et al., 2012). This study

represents the first systematic evaluation of

myriapods in these forest fragments in Bolívar,

contributing new taxonomic and ecological

insights for this arthropod group.

The rapid loss of Tropical Dry Forest veg-

etation has direct consequences for myriapod

communities, as many species depend on stable

microhabitats for survival. The observed diver-

sity variations across forest fragments under-

score the importance of conserving remaining

forest patches, which harbor unique species

assemblages. If these remnants were lost, a

significant portion of the region’s myriapod

diversity would disappear, further diminishing

ecosystem integrity.

14 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

Despite ongoing anthropogenic pressures,

the Tropical Dry Forest remnants of Bolívar

continue to support high arthropod diversity,

including myriapods. The results of this study

indicate that species richness and abundance

remain high, likely due to microhabitat het-

erogeneity across different successional stages.

This underscores the ecological significance of

these forests and highlights their priority sta-

tus for conservation efforts. Given the limited

research on Neotropical myriapods, further

assessments are necessary to expand taxonomic

knowledge and refine conservation strategies

for these ecosystems.

Author contribution statement: Martín

Carrillo-Pallares and Wilder Zapata-M contrib-

uted to the conception, design, execution, and

writing of this study. Daniela Martínez-Torres,

Gabriel R. Navas-S, and Adriana Bermúdez-

Tobón led the project design, performed speci-

men collection, identification, data analysis,

and contributed to manuscript writing and final

proofreading. All authors read and approved

the final version of the manuscript and agree to

its submission.

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 acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

ACKNOWLEDGMENTS

The authors thank the University of Carta-

gena for funding this research (Resolution No.

00473 de 2022) and the Institute of Natural

Sciences of the National University of Colom-

bia for its institutional support. Special thanks

to Sebastián Galvis and Fabio Cupul for their

help in specimen identification, and to Profes-

sor Eduardo Flórez for his contributions to the

ICN myriapodology collection. We also extend

our gratitude to field assistants Diego Bolaño,

Andrés Florez, José Martínez, Waldo León,

Dairo Rodríguez, David Vergara, Ada Luz San-

doval, and Gloribeth Ríos for their valuable

collaboration. Finally, we thank the local com-

munities of El Carmen Turbaco, San Jacinto,

and Magangue for their support, willingness,

and contributions.

REFERENCES

Adis, J., Minelli, A., de Morais, J. W., Pereira, L. A., Barbie-

ri, F., & Rodrigues, J. M. G. (1996). On abundance

and phenology of Geophilomorpha (Chilopoda) from

Central Amazonian Upland forests. Ecotropica, 2,

165–175.

Álvarez Hincapié, C. F., Maldonado-Ocampo, J., Sanjuan,

T., Álvarez-Dávila, E., Ajiaco, R. E., Buitrago, L.,

González, M. A., Herrera-R, G. A., Ortíz Gallego, R.,

Plata, C., Polanco, A., Rosselli, L., Uribe, U., Alterio,

H., Barrero, E. Y., & Laverde, O. (2021). Estado de

la biodiversidad en Colombia (pp. 100–369). En R.

Gómez-S., M. E. Chaves, W. Ramírez, M. Santama-

ría, G. Andrade, C. Solano, & S. Aranguren (Eds.),

Evaluación nacional de biodiversidad y servicios eco-

sistémicos de Colombia. Instituto de Investigación

de Recursos Biológicos Alexander von Humboldt,

Programa de las Naciones Unidas para el Desarrollo,

Centro Mundial de Monitoreo para la Conservación

del Programa de las Naciones Unidas para el Medio

Ambiente, & Ministerio Federal de Medio Ambiente,

Conservación de la Naturaleza y Seguridad Nuclear

de la República Federal de Alemania.

Alvarado-Solano, D. P., & Otero, J. T. (2015). Distribución

espacial del bosque seco tropical en el Valle del Cauca,

Colombia. Acta Biológica Colombiana, 20, 141–153.

https://doi.org/10.15446/abc.v20n2.46703

Aguilera, M. (2014). La economía de los Montes de María.

Economía & Región, 8(1), 81–141.

Bello, J. C., Báez, M., Gómez, M. F., Orrego, O., & Nägele,

L. (2014). Biodiversidad 2014: Estado y tendencias de

la biodiversidad continental de Colombia. Instituto

Alexander von Humboldt.

Bueno, J., & Rojas, P. (1999). Fauna de milpiés (Arthropoda:

Diplopoda) edáficos de una selva alta de los Tuxtlas,

Ver. México. Acta Zoológica Mexicana, 76, 59–83.

https://doi.org/10.21829/azm.1999.76761700

Bueno-Villegas, J. (2012). Diplópodos: los desconocidos

formadores de suelo. Biodiversitas, 102, 1–5.

Cantú-Salazar, L., Hidalgo-Mihart, M. G., López-González,

C. A., & González-Romero, A. (2005). Diet and food

resource use by pygmy skunk (Spilogale pygmaea) in

the tropical dry forest of Chamela, Mexico. Journal

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

of Zoology, 267(3), 283–289. https://doi.org/10.1017/

S0952836905007417

Cardoso, P., Crespo, L. C., Carvalho, R., Rufino, A. C., &

Henriques, S. S. (2009). Ad-hoc vs. standardized and

optimized arthropod diversity sampling. Diversity,

1(1), 36–51. https://doi.org/10.3390/d1010036

Carl, J. (1914). Die Diplopoden von Columbien nebst

Beiträgen zur Morphologie der Stemmatoiuliden.

Mémoires de la Société neuchâteloise des Sciences natu-

relles, 5, 821–993.

Carrillo-Pallares, M., Martínez-Torres, S. D., & Galvis-

Jiménez, S. (2019). Primer aporte a la diversidad de

miriápodos (Arthropoda: Myriapoda) del departa-

mento de Bolívar, Colombia. En Libro de resúmenes

del V Congreso Colombiano de Zoología (p. 46).

Castillo-Ávila, C. C., Prado, C., & García, A. (2015). First

record of the intake of lichenized fungi by diplopo-

ds (Myriapoda) in Eastern subregion of Sumapaz,

Colombia. Boletín de la Sociedad Entomológica Arago-

nesa, 57, 358–360.

Castillo, C. C. (2017). Caracterización de la miriapodofauna

(Arthropoda: Myriapoda) de bosque húmedo andino

en el municipio de Icononzo-Tolima, Colombia [Tesis

de pregrado]. Universidad Distrital Francisco José de

Caldas, Bogotá.

Chagas-Jr, A., Chaparro, E., Galvis, S., Triana, H. D., Flórez,

E., & Sícoli, J. C. (2014). The centipedes (Arthropo-

da, Myriapoda, Chilopoda) from Colombia: Part I.

Scutigeromorpha and Scolopendromorpha. Zootaxa,

3779(2), 133–156. 10.11646/zootaxa.3779.2.2

Chamberlin, R. V. (1921). Results of the Bryant Walker

expeditions of the University of Michigan to Colom-

bia 1913 and British Guiana 1914. Occasional Papers

of the Museum of Zoology, University of Michigan,

97, 1–39.

Chacón, P., Osorio-García, A. M., Achury, R., & Bermúdez-

Rivas, C. (2012). Hormigas (Hymenoptera: Formici-

dae) del Bosque seco Tropical (Bs-T) de la cuenca alta

del río Cauca, Colombia. Biota Colombiana, 13(2),

165-181.

Chao, A., Ma, K. H., Hsieh, T. C., & Chiu, C.-H. (2016).

User’s guide for online program SpadeR (Species-rich-

ness Prediction and Diversity Estimation in R). Natio-

nal Tsing Hua University. https://doi.org/10.13140/

RG.2.2.20744.62722

Colwell, R. K., Chao, A., Gotelli, N. J., Lin, S, Mao, C.

X., Chazdon, R. L., & Longino, J. T. (2012). Models

and estimators linking individual-based and sample-

based rarefaction, extrapolation and comparison of

assemblages. Journal of Plant Ecology, 5(1), 3–21.

https://doi.org/10.1093/jpe/rtr044

Colwell, R. K. (2013). EstimateS: Statistical estimation

of species richness and shared species from samples

(versión 9.1) [Software] https://www.robertkcolwell.

org/pages/1407-estimates

Cupul-Magaña, F. G. (2012). Los ciempiés escutigeromor-

fos (Scutigeromorpha), escolopendromorfos (Scolo-

pendromorpha) y geofilomorfos (Geophilomorpha)

de la selva tropical caducifolia de la reserva de Cha-

mela, Jalisco, México. Insecta Mundi, 0208, 1–17.

Cupul-Magaña, F. G. (2014). Los ciempiés escolopendro-

morfos (Chilopoda: Scolopendromorpha) de México:

clave para géneros. Revista Colombiana de Entomolo-

gía, 40(2), 286–291.

Daniels-Puello, A. (2016). La transformación de la estruc-

tura productiva de los Montes de María: de despen-

sa agrícola a distrito minero-energético. Memorias,

Revista digital de historia y arqueología desde el Caribe

colombiano, 12(28), 52–83. https://doi.org/10.14482/

memor.29.8278

Díaz, J. M. (2006). Bosque seco tropical Colombia. Libros de

la Colección Ecológica.

Díaz, M., Rionda, M. H., Duhour, A. E., & Momo, F. R.

(2014). Artrópodos del suelo: Relaciones entre la

composición faunística y la intensificación agrope-

cuaria. Ecología Austral, 24, 327–334. https://doi.

org/10.25260/EA.14.24.3.0.10

Druce, D., Hamer, M., & Slotow, R. (2004). Sampling strate-

gies for millipedes (Diplopoda), centipedes (Chilopo-

da) and scorpions (Scorpionida) in savanna habitats.

African Zoology, 39(2), 293−304. https://doi.org/10.10

80/15627020.2004.11657225

Dryflor, B., Banda-R, K., Delgado-Salinas, A., Dexter, K.

G., Linares-Palomino, R., Oliveira-Filho, A., Prado,

D., Pullan, M., Quintana, C., Riina, R., Rodríguez,

G. M., Weintritt, J., Acevedo-Rodríguez, P., Adarve,

J., Álvarez, E., Aranguren, A., Arteaga, J. C., Aymard,

G., Castaño, A., … Pennington, R. T. (2016). Plant

diversity patterns in neotropical dry forests and their

conservation implications. Science, 353(6306), 1383–

1387. https://doi.org/10.1126/science.aaf5080

Edgecombe, G. D. (2011). Phylogenetic relationships of the

Myriapoda. In A. Minelli (Ed.), The Myriapoda (Vol.

1, pp. 1–20). Brill.

Escorcia, R. Y., Martínez, N. J., & Silva, J. P. (2012). Estudio

de la diversidad de arañas de un Bosque Seco Tropical

(Bs-T) en Sabanalarga, Atlántico, Colombia. Boletín

Científico Centro de Museos Museo de Historia Natural

de la Universidad de Caldas, 16(1), 247–260.

Etter, A. (1993). Diversidad ecosistémica en Colombia

hoy. En S. Cárdenas & H. D. Correa (Eds.), Nuestra

diversidad biológica (pp. 44–61). CEREC/Fundación

Alejandro Ángel Escobar.

Foddai, D., Pereira, L. A., & Minelli, A. (2000). A catalogue

of the geophilomorph centipedes (Chilopoda) from

16 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

Central and South America including Mexico. Ama-

zoniana, 14(1–2), 59–185.

Instituto de Hidrología, Meteorología y Estudios Ambien-

tales, Instituto Geográfico Agustín Codazzi, Instituto

de Investigación de Recursos Biológicos Alexander

von Humboldt, Instituto de Investigaciones Marinas

y Costeras José Benito Vives De Andréis, Instituto

Amazónico de Investigaciones Científicas Sinchi, &

Instituto de Investigaciones Ambientales del Pacífico

Jhon von Neumann. (2007). Ecosistemas continentales,

costeros y marinos de Colombia. Imprenta Nacional

de Colombia.

Galvis, L. A. (2009). Geografía económica del Caribe Conti-

nental (p. 87). Banco de la República.

Galvis, J. S. (2019). Estimando la diversidad de ciempiés

(Chilopoda, Myriapoda) en un bosque de niebla de los

Andes de Colombia: Aporte al establecimiento de un

protocolo de muestreo [Tesis de pregrado, Universidad

Nacional de Colombia].

García-Herrera, L. V., Ramírez-Fráncel, L. A., & Reinoso-

Flórez, G. (2019). Mamíferos del departamento del

Tolima: distribución y estado de conservación. Revis-

ta UDCA Actualidad & Divulgación Científica, 22(2).

González-Montaña, L. A., Wilches-Álvarez, W., & Rocha-

Caicedo, C. (2017). Diversidad alfa y beta de artró-

podos epigeos (Arthropoda) en tres sectores sobre

la Cordillera Oriental de Colombia. Revista Colom-

biana de Entomología, 43(2), 292–300. https://doi.

org/10.25100/socolen.v43i2.5962

Grainger, A. (1996). An evaluation of the FAO tropical forest

resource assessment, 1990. The Geographical Journal,

162(1), 73–79. https://doi.org/10.2307/3060217

Hammer, Ø., Harper, D. A. T., & Ryan, P. D. (2001). PAST:

Paleontological Statistics Software Package for Edu-

cation and Data Analysis. Palaeontologia Electronica,

4(1), 1–9. https://palaeo-electronica.org/2001_1/past/

past.pdf

Hoffman, R. L. (1999). Checklist of millipeds of North and

Middle America. Virginia Museum of Natural History.

Hoffman, R., Golovatch, S. I., Adis, J., & Morrais, J.

W. (2002). Diplopoda. In J. Adis (Ed.), Amazonian

Arachnida and Myriapoda (pp. 505–533). Pensoft

Publishers.

Janzen, D. H. (1980). Specificity of seed-attacking beetles

in a Costa Rican deciduous forest. Journal of Ecology,

68(3), 929–952. https://doi.org/10.2307/2259466

Janzen, D. H. (1988). Management of habitat fragments

in a tropical dry forest: Growth. Annals of the Mis-

souri Botanical Garden, 75(1), 105–116. https://doi.

org/10.2307/2399468

Lewis, J. G. E. (1981). The biology of centipedes (p. 476).

Cambridge University Press.

Malumbres-Olarte, J., Scharff, N., Pape, T., Coddington, J.

A., & Cardoso, P. (2017). Gauging megadiversity with

optimized and standardized sampling protocols: A

case for tropical forest spiders. Ecology and Evolution,

7, 494–506. https://doi.org/10.1002/ece3.2626

Martínez-Torres, D., & Flórez, E. (2015). Clase Diplopoda.

En E. Flórez, C. Romero-Ortíz, & D. S. López (Eds.),

Los artrópodos de la Reserva Natural Río Ñambí (Vol.

15, pp. 263–288). Serie de Guías de Campo del Insti-

tuto de Ciencias Naturales.

Mayaux, P., Holmgren, P., Achard, F., Eva, H., Stibig, H., &

Branthomme, A. (2005). Tropical forest cover chan-

ge in the 1990s and options for future monitoring.

Philosophical Transactions of the Royal Society B, 360,

373–384. https://doi.org/10.1098/rstb.2004.1590

Menco, D. (2013). Tenencia y usos de la tierra rural en

Montes de María-Colombia. Observatorio de la Eco-

nomía Latinoamericana, (185). https://ideas.repec.

org/a/erv/observ/y2013i18520.html

Miles, L., Newton, A. C., DeFries, R. S., Ravilious, C., May,

I., Blyth, S., Kapos, V., & Gordon, J. E. (2006). A

global overview of the conservation status of tropical

dry forests. Journal of Biogepography, 33(3), 491–505.

https://doi.org/10.1111/j.1365-2699.2005.01424.x

Molinari, J., Gutiérrez, E. E., de Ascencão, A. A., Nassar, J.

M., Arends, A., & Márquez, R. J. (2005). Predation by

giant centipedes, Scolopendra gigantea, on three spe-

cies of bats in a Venezuelan cave. Caribbean Journal of

Science, 41(2), 340–346.

Morales, A. Y. (2019). Estructura y caracterización de la

comunidad de milpiés (Arthropoda: Diplopoda) en tres

fragmentos boscosos urbanos del norte de Armenia,

Quindío-Colombia [Tesis de pregrado, Universidad

del Quindío].

Moreno, C. E., Barragán, F., Pineda, E., & Pavón, N. P.

(2011). Reanálisis de la diversidad alfa: alternati-

vas para interpretar y comparar información sobre

comunidades ecológicas. Revista Mexicana de Biodi-

versidad, 82(4), 1249–1261. https://doi.org/10.22201/

ib.20078706e.2011.4.745

Murphy, P. G., & Lugo, A. E. (1986). Ecology of tropical dry

forest. Annual Review of Ecology, Evolution, and Syste-

matics, 17(1), 67–88. https://doi.org/10.1146/annurev.

es.17.110186.000435

Pennington, R. T., Lewis, G. P., & Ratter, J. A. (2006). An

overview of the plant diversity, biogeography and con-

servation of Neotropical savannas and seasonally dry

forests. In R. T. Pennington, G. P. Lewis, & J. A. Ratter

(Eds.), Neotropical savannas and seasonally dry forests

(pp. 17–45). CRC Press.

Pérez-Asso, A. R., & Pérez-Gelabert, D. E. (2001). Checklist

of the millipeds (Diplopoda) of Hispaniola. Boletín de

la Sociedad Entomológica Aragonesa, 28, 67–80.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025546, enero-diciembre 2025 (Publicado Set. 18, 2025)

Pizano, C., Cabrera, M., & García, H. (2014). Bosque seco

tropical en Colombia: Generalidades y contexto. En

C. Pizano & H. García (Eds.), El bosque seco tropical

en Colombia (pp. 37–47). Instituto de Investigación de

Recursos Biológicos Alexander von Humboldt.

Prado, C. C., & Herrera, C. A. (2013). Contribución al estu-

dio de los miriápodos en la Quebrada La Vieja, Bosques

Orientales, Bogotá [Tesis de pregrado, Universidad

Distrital Francisco José de Caldas].

Prado-Sepúlveda, C., Triana, H. D., & Galvis, S. (2016). Los

ciempiés (Myriapoda: Chilopoda) de bosque andino

en el municipio de Icononzo (Colombia, Tolima) y

clave para las familias presentes en Colombia. Boletín

de la Sociedad Entomológica Aragonesa, 58, 188–196.

Ramírez, V. A., Aranda, L., & Rico, V. (2018). La resiliencia

del bosque seco tropical: un seguro de vida para su

conservación. Biodiversitas, 137, 13–16.

Rangel-Acosta, J. L., Blanco-Rodríguez, O. R., & Mar-

tínez-Hernández, N. J. (2016). Escarabajos copro-

necrófagos (Scarabaeidae: Scarabaeinae) en diferentes

usos del suelo en la Reserva Campesina La Montaña

(RCM) en el departamento del Atlántico, Colombia.

Boletín Científico Centro de Museos Museo de Historia

Natural de la Universidad de Caldas, 20(1), 78–97.

https://doi.org/10.17151/bccm.2016.20.1.7

Rendón-Carmona, H., Martínez-Yrízar, A., Balvanera, P., &

Pérez-Salicrup, D. (2009). Selective cutting of woody

species in a Mexican tropical dry forest: Incompa-

tibility between use and conservation. Forest Ecolo-

gy and Management, 257(2), 567–579. https://doi.

org/10.1016/j.foreco.2008.09.031

Rendón, S., Artunduaga, F., Ramírez, R., Quiroz, J. A., &

Leiva, E. I. (2011). Los macroinvertebrados como

indicadores de la calidad del suelo en cultivos de

mora, pasto y aguacate. Revista Facultad Nacional de

Agronomía Medellín, 64(1), 5793–5802.

Rodríguez, G. M., Banda-R, K., Reyes, S. P., & Estupiñan, A.

C. (2012). Lista comentada de las plantas vasculares

de bosques secos prioritarios para la conservación

en los departamentos de Atlántico y Bolívar (Caribe

colombiano). Biota Colombiana, 13(2), 7–39.

Ruiz-Cobo, D. H., Bueno-Villegas, J., & Feijoo-Martínez,

A. (2010). Uso de la tierra y diversidades alfa, beta

y gamma de diplópodos en la cuenca del río Otún,

Colombia. Universitas Scientiarum, 15(1), 59–67.

http://dx.doi.org/10.11144/javeriana.SC15-1.luaa

Shelley, R. M. (2002). A revised, annotated, family-level

classification of the Diplopoda. Arthropoda Selecta,

11(3), 187–207.

Shelley, R. M., & Lehtinen, P. T. (1998). Introduced milli-

peds of the family Paradoxosomatidae on Pacific

Islands (Diplopoda: Polydesmida). Arthropoda Selecta,

7(2), 81−94.

Simanca-Fontalvo, R., Fajardo-Herrera, R. J., & Martínez,

N. J. (2013). Fauna de hormigas (Hymenoptera: For-

micidae) en dos remanentes de Bosque Seco Tropical

(Bs-T) en Corrales de San Luis, Atlántico, Colombia.

Boletín del Museo de Entomología de la Universidad

del Valle, 14(1), 1–14.

Sociedad Colombiana de Entomología. (2016). Libro de

resúmenes 43° Congreso de la Sociedad Colombiana de

Entomología (287 p.). SOCOLEN.

Suriel, C. (2012). Presencia de Oxidus gracilis, Chon-

dromorpha xanthotricha y Asiomorpha coarctata

(Diplopoda: Paradoxosomatidae) en La Hispaniola.

Solenodon, 10, 23–31.

Wardle, D. A. (2006). The influence of biotic interac-

tions on soil biodiversity. Ecology Letters, 9(7).

870–886. https://onlinelibrary.wiley.com/doi/

epdf/10.1111/j.1461-0248.2006.00931.x

Zapata, W., Vergara-Moreno, D., Carrillo-Pallares, M.,

Segovia-Paccini, A., Navas-S., G. R., & Malumbres-

Olarte, J. (2023). Seasonal diversity of spider assem-

blages (Araneae: Arachnida) in the “Guillermo

Piñeres” Botanical Garden, Turbaco–Colombia. Neo-

tropical Biodiversity, 9(1), 17–28. https://doi.org/10.10

80/23766808.2022.2157948