Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61cmxq06, enero-diciembre 2025 (Publicado Ago. 20, 2025)

Distribution, richness, and abundance of Lycaenidae

(Lepidoptera: Papilionoidea) in the Loxicha Region, Oaxaca,

Mexico: A regional analysis on the Pacific slope

Armando M. Luis-Martínez1; https://orcid.org/0000-0002-1044-3986

Robert K. Robbins2; https://orcid.org/0000-0003-4137-786X

Marysol Trujano-Ortega1; https://orcid.org/0000-0001-8911-8504

Jorge E. Llorente-Bousquets1; https://orcid.org/0000-0003-0876-0533

Arturo Arellano-Covarrubias1; https://orcid.org/0000-0001-9515-5782

Omar Ávalos-Hernández1*; https://orcid.org/0000-0002-5476-9400

1. Museo de Zoología (Entomología), Departamento de Biología Evolutiva, Facultad de Ciencias, Universidad Nacional

Autónoma de México, Ciudad de México, México; alm@ciencias.unam.mx, marysol_trujano@yahoo.com.mx, lloren-

tebousquets@gmail.com, arellano.covarrubias@gmail.com, omaravalosh@ciencias.unam.mx (*Correspondence)

2. Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, District of

Columbia, USA; robbinsr@si.edu

Received 21-III-2025. Corrected 23-VII-2025. Accepted 12-VIII-2025.

ABSTRACT

Introduction: The Sierra Madre del Sur in Mexico, known for its geological complexity and elevational gradient,

hosts diverse ecosystems. The Loxicha region in Oaxaca, with five vegetation types and a 2 500 m elevation range,

has the richest documented butterfly fauna on the Pacific slope. This study focuses on Lycaenidae butterflies,

exploring their distribution, richness, and abundance along vegetation and elevational transects.

Objective: To document geographic, elevational, and ecological patterns of Lycaenidae butterflies on the Pacific

slope Loxicha region as part of a larger project on the distribution of diurnal butterflies in the state of Oaxaca,

Mexico. Also, to compare Loxicha fauna with other regions to recognize biogeographical patterns at a larger scale.

Methods: Spatial and temporal patterns of Lycaenidae diversity are based on systematic sampling conducted

between 2005 and 2014 along elevational (80 to 2 500 m) and vegetational transects. Exclusive and characteristic

species for each elevation and vegetation type are recognized.

Results: A total of 123 species of Lycaenidae were recorded: nine species of Polyommatinae and 114 of Theclinae.

The highest species richness occurs in May in tropical semi-deciduous forest. It decreases with increasing eleva-

tion. The results are compared with four other butterfly families. Lycaenid community similarity in the Sierra

Madre del Sur and Pacific slope is correlated with elevation and vegetation type, though variability within

each grouping is significant. Higher elevation faunal isolation also plays a role, as seen in other mountainous

Papilionoidea taxa.

Conclusions: Lycaenidae is one of the most diverse and least known butterfly families in Mexico. This is the

fourth study on the Loxicha Region’s lepidopteran fauna, confirming its high diversity among five Papilionoidea

families. Systematic studies aid in understanding speciation and endemism, particularly for stenotopic species.

The distribution of Lycaenidae in the Loxicha region is the most comprehensive to date for any region in Mexico.

Key words: elevational gradient; Eumaeini; phenology; Sierra Madre del Sur.

https://doi.org/10.15517/61cmxq06

INVERTEBRATE BIOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61cmxq06, enero-diciembre 2025 (Publicado Ago. 20, 2025)

INTRODUCTION

Mexico’s Sierra Madre del Sur is recog-

nized for its complex geological history and for

an elevational gradient that generates ecosys-

tem heterogeneity (Espinosa et al., 2016). This

complexity is reflected in the Loxicha region

(Oaxaca), which consists of five distinctive

vegetation types and an elevation gradient of 2

500 m. It is an underlying reason why we have

been investigating its butterfly richness along

elevation and vegetation transects (Arellano-

Covarrubias et al., 2018; Luis-Martínez et al.,

2016; Luis-Martínez et al., 2020; Luis-Martínez

et al., 2022). It has the richest documented but-

terfly fauna on Mexico’s Pacific slope for the

four families treated so far.

Lycaenidae present challenges for sampling.

Adults tend to be more rarely encountered than

other butterflies, for example, at about half the

rate as that of Riodinidae (Lamas et al., 2023).

Bait traps with rotting carrion seem to be inef-

fective in Mesoamerican localities (Busby et al.,

2019). These characteristics potentially bias the

documentation of Lycaenidae spatial and tem-

poral patterns compared to other butterflies.

New World Lycaenidae exhibit other eco-

logical characteristics that distinguish the fam-

ily. The caterpillars show significant variation

in feeding habits, including extreme polyphagy

(Chew & Robbins, 1984), florivory (Silva et al.,

2014), detritivory (Duarte & Robbins, 2009,

Robbins et al., 2010), and fungivory (Nishida

& Robbins, 2020). Many species are facultative-

ly myrmecophilous (DeVries, 1990; DeVries,

1991; Kaminski et al., 2012). More than 90 %

of the species have adult males with secondary

sexual organs (Valencia-Montoya et al., 2021).

RESUMEN

Distribución, riqueza y abundancia de Lycaenidae (Lepidoptera: Papilionoidea) en la Región Loxicha,

Oaxaca, México: Un análisis regional en la vertiente del Pacífico

Introducción: La Sierra Madre del Sur en México, conocida por su complejidad geológica y gradiente de eleva-

ción, alberga ecosistemas diversos. La región de Loxicha en Oaxaca, con cinco tipos de vegetación y un rango de

elevación de 2 500 m, tiene la fauna de mariposas documentada más rica en la vertiente del Pacífico. Este estudio

se enfoca en las mariposas de la familia Lycaenidae, explorando su distribución, riqueza y abundancia a lo largo

de transectos de vegetación y elevación.

Objetivo: Documentar los patrones geográficos, altitudinales y ecológicos de Lycaenidae en la región de Loxicha

en la vertiente del Pacífico, como parte de un proyecto más amplio sobre la distribución de mariposas diurnas en

el estado de Oaxaca, México. También, comparar la fauna de Loxicha con otras regiones para reconocer patrones

biogeográficos a mayor escala.

Métodos: Los patrones espaciales y temporales de diversidad de Lycaenidae se basan en muestreos sistemáticos

realizados entre 2005 y 2014 a lo largo de transectos de elevación (80 a 2 500 m) y vegetación. Se reconocen espe-

cies exclusivas y características de cada altitud y tipo de vegetación.

Resultados: Se registraron un total de 123 especies de Lycaenidae: nueve especies de Polyommatinae y 114 de

Theclinae. La mayor riqueza de especies se observa en mayo en bosques tropicales semi-deciduos. Esta disminuye

con el aumento de la elevación. Los resultados se comparan con otras cuatro familias de mariposas. La similitud

de la comunidad de Lycaenidae en la Sierra Madre del Sur y la vertiente del Pacífico está correlacionada con la

elevación y el tipo de vegetación, aunque la variabilidad dentro de cada agrupación es significativa. El aislamiento

faunístico en elevaciones altas también juega un papel, como se observa en otros taxones de Papilionoidea de

sistemas montañosos.

Conclusiones: Lycaenidae es una de las familias de mariposas más diversas y menos conocidas de México. Este

es el cuarto estudio sobre la fauna lepidópteros de la región de Loxicha, confirmando su alta diversidad entre

cinco familias de Papilionoidea. Los estudios sistemáticos ayudan a comprender la especiación y el endemismo,

particularmente para las especies estenotópicas. La distribución de Lycaenidae en la región de Loxicha es la más

completa hasta la fecha para cualquier región de México.

Palabras clave: gradiente altitudinal; Eumaeini; fenología; Sierra Madre del Sur.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61cmxq06, enero-diciembre 2025 (Publicado Ago. 20, 2025)

The characteristics of Lycaenidae suggest

that their spatial and temporal distributions

might differ from those of other Papilionoidea.

This study specifically examines the distribu-

tion, richness, and abundance of Lycaenidae

along vegetation and elevational transects in

the Loxicha region and is the first such investi-

gation in any of Mexico’s most diverse regions.

The results are compared with those of other

regions of the Pacific and Atlantic slopes in

Mexico. An updated checklist with photographs

of Lycaenidae from Loxicha is presented, serv-

ing as a reference for future identification

within the family.

Documentation of the Mexican lycaenid

butterfly fauna has progressed for more than

a century. Godman and Salvin (1887-1901)

monographed the diurnal butterfly fauna of

Mesoamerica, recording 124 species of Mexi-

can Lycaenidae. Hoffmann’s (1933), Hoffmann’s

(1936), Hoffmann’s (1940) and Hoffmann’s

(1941) checklist of Mexican Papilionoidea

included distribution by state, physiograph-

ic unit, elevation, and climatic subtype. He

included 221 Lycaenidae. Subsequently, Llor-

ente-Bousquets et al. (2006) recorded 255 spe-

cies and subspecies of Mexican Lycaenidae,

which was later refined to 252 species and

subspecies, representing 13 % of the Mexican

Papilionoidea (Llorente-Bousquets et al., 2014).

Knowledge of the lycaenid fauna of Oaxa-

ca has similarly steadily improved. Hoffmann

(1940) and Hoffman (1941) recorded 164 lycae-

nid species in Oaxaca. More recently, Luis-Mar-

tínez et al. (2016) listed 180 lycaenid species,

partitioned taxonomically into 169 Theclinae,

10 Polyommatinae, and 1 Lycaeninae.

MATERIAL AND METHODS

Previous data: The MARIPOSA database

(Luis-Martínez et al., 2005) from the Zool-

ogy Museum of the Faculty of Science, UNAM

has more than 65 000 records documenting

the geographic distribution of Lycaenidae and

other butterflies in Mexico. Some data are avail-

able on the CONABIO website (Gobierno de

México, n. d.), and we are uploading more. The

data range from the late 19th century (Godman

& Salvin, 1887-1901) to the present and include

more than 2 300 sites. Using the database and

publications on Lycaenidae from other Mexican

regions (de la Maza & de la Maza, 1993, de la

Maza & de la Maza, 2015; Luis-Martínez et

al., 1991; Raguso & Llorente-Bousquets, 1991;

Ross, 1966, Ross, 1976; Vargas-Fernández et

al., 1994; Vargas-Fernández et al., 1999), the

localities and regions (groups of localities) with

the greatest recorded Lycaenidae richness in

Mexico are listed (Table 1).

Faunistic inventory and localities: Since

2005, we conducted systematic butterfly sam-

pling in the Loxicha region (Fig. 1), com-

prising 318 sampling days over seven years

(2005, 2007, 2008, 2011–2014) at 21 localities

(Table 2) (Arellano-Covarrubias et al., 2018;

Luis-Martínez et al., 2016; Luis-Martínez et al.,

2020; Luis-Martínez et al., 2021; Luis-Martínez

et al., 2022). The surveys involved 16 par-

ticipants, averaging five per locality per day,

most of whom had extensive experience in

sampling butterflies.

Localities were grouped by elevation and

vegetation to analyze distribution patterns,

which facilitates comparison with other locali-

ties. Vegetation type classification follows Rze-

dowski (1978) and Llorente-Bousquets (1984):

tropical deciduous forest (TDF), tropical sub-

deciduous forest (TSDF), montane cloud forest

(CF) with three subtypes, and oak-pine forest

(OPF). Additionally, in some sites at elevations

between 2 000 and 2 400 m, a vegetation type

primarily comprising pine-oak forest with ele-

ments of high-altitude montane cloud forest

was recognized, mostly located at the bottom

of ravines. This high-altitude oak-pine and

montane cloud forest (OPCF) is distinct from

OPF and CF, although adjacent to them, and

was considered a separate category (Arellano-

Covarrubias et al., 2018; Luis-Martínez et al.,

2020; Luis-Martínez et al., 2021; Luis-Martínez

et al., 2022). Sampling focused on effective

sites, such as sunny areas with small white

inflorescences, where species of the subfamily

Theclinae were primarily observed.

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Taxonomy: Identification was carried out

by comparison with the Lepidoptera Collection

at the Museum of Zoology, Faculty of Sciences,

UNAM (MZFC) and the National Museum

of Natural History, Smithsonian Institution

(USNM). The updated species list by subtribe

follows Robbins et al. (2022).

Deposition and permits: Specimens were

deposited in the Papilionoidea Collection (Lep-

idoptera) at the Museum of Zoology under

registration number DFE.IN.071.0798 issued

by the Secretariat of Environment, Natural

Resources, and Fisheries (SEMARNAP). The

specimens were collected under the scientific

Table 1

Single sites and regions (Reg. is a group of sites with more than one elevation or vegetation zone) with highest species richness

of Lycaenidae in Mexico.

Altitude (m) Site / Region spp. P / T / Total

Atlantic Slope

0-1 700 Reg. Los Tuxtlas-Catemaco3,4,5* - Ver. 8 / 129 / 137

100-2 800 Reg. Sierra de Juárez2* - Oax. 8 / 117 / 125

500-1 500 Reg. Macizo Central6 - Chis. 7 / 116 / 123+1^

100-1 500 Reg. Selva Lacandona6 - Chis. 4 / 115 / 119

0-2 500 Reg. Cuenca de Tulijá6 - Chis. 4 / 109 / 113

250-1 000 Reg. Cuenca del Grijalva6 - Chis. 4 / 108 / 112

140-200 Site. Río Lacantún1 - Chis. 5 / 103 / 108

200-350 Site. Cuetzalapan* - Ver. 2 / 101 / 103

350 Site. Catemaco* - Ver. 3 / 95 / 98

350-950 Reg. Barranca de Patla* - Pue. 5 / 86 / 91

650 Site. Tequezquitla* - Pue. 5 / 84 / 89

1 350 Site. San Antonio Buenavista (Sta Rosa)* - Chis. 4 / 81 / 85

350 Site. Presidio* - Ver. 4 / 77 / 81

1 200-1 600 Reg. Xalapa* - Ver. 7 / 71 / 78

900 Site. Carretera 175, Metates2* - Oax. 4 / 66 / 70

1 150-1 350 Site. Coatepec* - Ver. 4 / 64 / 68

300 Site. Tapalapan* - Ver. 3 / 64 / 67

860 Site. Cerro El Vigía, Santiago Tuxtla* - Ver. 2 / 64 / 66

1 200-1 500 Reg. Sierra Madre6 - Chis. 5 / 59 / 64

500-1 500 Reg. Depresión Central5 - Chis. 6 / 54 / 60

Pacific slope

80-2 850 Reg. Loxicha - Oax. 9 / 114 / 123

400 Site. Rancho Hagia Sofía* - Oax. 7 / 84 / 91

300-2 500 Reg. Sierra de Atoyac7* - Gro. 7 / 81 / 88

750 Site. Rancho “El Zorrillo”, Cañada Húmeda* - Mich. 6 / 74 / 80

260 Site. Chiquihuitillo* - Mich. 6 / 65 / 70

250-1 800 Reg. Sierra de Manantlán8* - Jal. / Col. 8 / 59 / 67

2 250 Site. Santa Rosa, Comitán* - Chis. 6 / 60 / 66

900 Site. Planta Hidroeléctrica Cupatitzio* - Mich. 7 / 58 / 65

1 200-1 500 Reg. Soconusco6 - Chis. 4 / 58 / 62+1^

650 Site. Río Santiago, 4 km W* - Gro. 8 / 53 / 61

850 Site. Arteaga* - Mich. 6 / 55 / 61

P: Polyommatinae; T: Theclinae; Chis.: Chiapas; Col.: Colima; Gro.: Guerrero; Jal.: Jalisco; Mich.: Michoacán; Oax.: Oaxaca;

Ver.: Veracruz; ^ Species recorded from the Lycaeninae subfamily. Source of data: *, MARIPOSA database (Luis-Martínez

et al., 2005); 1, de la Maza & de la Maza (1993); 2, Luis-Martínez et al. (1991); 3, Ross (1966); 4, Ross (1976); 5, Raguso &

Llorente-Bousquets (1991); 6, de la Maza & de la Maza (2015); 7, Vargas-Fernández et al. (1994); 8, Vargas-Fernández et al.

(1999).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61cmxq06, enero-diciembre 2025 (Publicado Ago. 20, 2025)

Fig 1. Location of the geographic units on the Pacific slope analyzed: Loxicha Region, Oaxaca; Sierra de Atoyac de Álvarez,

Guerrero; Omiltemi Region, Guerrero; locality Ixcateopan de Cuauhtémoc, Guerrero; Sierra de Manantlán, Jalisco-Colima.

The Sierra Madre del Sur province (grey) and state lines are outlined.

Table 2

Number of specimens and species by locality.

Altitude (m) Vegetation / Site P T Total

  Tropical deciduous forest (TDF)

0-500 80-100 Parque Nacional Huatulco, río Cacaluta 53 / 4 37 / 16 90 / 20

100 Parque Nacional Huatulco 151 / 5 53 / 20 204 / 25

Tropical subdeciduous forest (TSDF)

380-500 Azulillo 298 / 8 212 / 37 510 / 45

410 Rancho Hagia Sofía 303 / 7 1 738 / 84 2041 / 91

  Cloud forest (CF)

500-1 000 530-700 Río Molinos* 24 / 4 13 / 9 37 / 13

600-1 200 Copalita, río Copalita* 3 / 2 19 / 15 22 / 17

750-900 Magdalena, El Lirio 57 / 5 95 / 27 152 / 32

900 Copalita, Los Plátanos* 0 / 0 1 / 1 1 / 1

940 Copalita, Siete Veneros* 29 / 4 5 / 2 34 / 6

1 000-1 500 1 000 San Mateo Piñas* 0 / 0 3 / 3 3 / 3

1 100-1 200 Pluma Hidalgo, 4 km NW “La Curva” 128 / 9 146 / 34 274 / 43

1 100-1 250 Finca Aurora-Finca San Isidro 65 / 5 105 / 18 170 / 23

1 200 Copalita, Llano de Ocote* 0 / 0 18 / 4 18 / 4

1 200-1 530 Portillo del Rayo-Finca El Encanto 47 / 6 54 / 16 101 / 22

1 470-1 550 La Soledad-Buenavista 54 / 6 63 / 21 117 / 27

1 500-1 650 La Pasionaria 11 / 3 114 / 16 125 / 19

  Oak-pine - cloud forest (OPCF)

1 500-2 000 1 950-2 150 Puente Arroyo “El Guajolote” 53 / 5 26 / 12 79 / 17

>2 000 2 280-2 400 San José del Pacífico, 1 km S 281 / 6 264 / 32 545 / 38

Oak-pine forest (OPF)

2 700-2 800 Manzanal-Doncella* 14 / 3 4 / 3 18 / 6

2 700-2 800 Camino a San Agustín Loxicha* 0 / 0 2 / 2 2 / 2

2 820-2 850 Nevería-La Ciénega* 1 / 1 2 / 2 3 / 3

  Total 1 572 / 9 2 974 / 114 4 547 / 123

P: Polyommatinae. T: Theclinae. specimens / number of species. * Localities considered without exhaustive sampling.

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collecting permit issued to Jorge Enrique Llor-

ente Bousquets by the Ministry of Environment

and Natural Resources (SEMARNAT), with

permit number FAUT-0148. Specimen data

were incorporated into the MARIPOSA data-

base (Luis-Martínez et al. 2005).

Species richness by vegetation and eleva-

tion: Species richness was estimated for each

vegetation type and for five elevation zones:

0-500, 500-1 000, 1 000-1 500, 1 500-2 000, >

2 000 m. The non-parametric estimator Chao1

(Colwell & Coddington, 1994) was used with

the SpadeR package (Chao et al., 2016). A con-

fidence interval (95 %) with the standard error

was calculated for each estimate using the Boot-

strap technique. Significant differences in spe-

cies richness among months, vegetation types,

and elevation zones were evaluated by the over-

lap of confidence intervals for the estimates.

Additionally, for the elevation zones, eco-

logical diversity was estimated as the exponen-

tial of the Shannon-Wiener index (H’), using

the Chao & Shen (2003) method implemented

in SpadeR (Chao et al., 2016). This diversity

measure is also known as the Effective Number

of Species (NES) (Jost, 2006). The index takes

into account both species richness and their

relative abundance.

Exclusive and characteristic species: Spe-

cies recorded only in a single vegetation type

or in a single elevation zone, were considered

stenoecious sensu Udvardy (1969), occupying

a narrow range of the habitats available, and

referred as “exclusive”. Those with 80 to 99 %

of their records in a particular vegetation type

or elevation zone were considered amphiecious

and referred as “characteristic”, occupying two

or more types of habitats with a broader ecolog-

ical tolerance than stenoecious species, but still

not as flexible as euryecious species. The rest of

the species were considered euryecious, occu-

pying a wide range of the habitats available.

Faunal similarity with other Mexican

Pacific Slope localities: To determine how veg-

etation and elevation determine the occurrence

of lycaenid species on Mexico’s Pacific slope, we

calculated a distance matrix dendrogram from

species abundance data using the Bray-Curtis

index through the “vegan” package (Oksanen et

al., 2022) in R 4.0.0 (R Core Team, 2023). The

analysis included data from 31 localities (Fig.

1): 12 from the Loxicha region in Oaxaca (this

study); 10 from Guerrero: El Faisanal, Las Paro-

tas, Nueva Delhi, Puente de los Lugardo, and Río

Santiago 4 km W in Sierra de Atoyac de Álvarez

(Vargas-Fernández et al., 1994); Cañada W of La

Gruta del Borrego, Potrerillo, Presa de la Perra,

and Rancho Omiltemi in the Omiltemi region

(Luis-Martínez & Llorente-Bousquets, 1993),

and Ixcateopan de Cuauhtémoc. Also included

were six localities from the Sierra de Manantlán

in Jalisco-Colima: Agua Dulce, Ahuacapán, La

Calera, Los Mazos 9 mi SW Autlán, Platanarillo,

and Zenzontla (Vargas-Fernández et al., 1994).

The localities were selected based on vegetation

type, geographic location (region), elevation

(100-2 600 m), and sampling coverage.

RESULTS

Mexican Lycaenidae richness by local-

ity: The 17 individual localities and 14 regions

(groups of localities) with the highest Lycae-

nidae species richness in Mexico are listed

(Table 1). Two localities exceed 100 species:

Río Lacantún in Chiapas with 108 (45 %)

species (de la Maza & de la Maza, 1993) and

Cuetzalapan in Veracruz with 103 (43 %) spe-

cies (Luis-Martínez et al., 2005). The region

(group of localities) with the highest number

of species is Los Tuxtlas and the surroundings

of Catemaco, Veracruz, with 137 species (57 %

of the total recorded in the country); this is one

of the regions with the most extensive sampling

efforts primarily from 1960 to 1990 (Raguso &

Llorente-Bousquets, 1991). More than 94 % of

the 2 159 documented Mexican localities have

10 or fewer lycaenid species recorded. These

data demonstrate the historic insufficiency of

sampling effort in most of Mexico.

The Loxicha region: The MARIPOSA

database (Luis-Martínez et al., 2005) had

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61cmxq06, enero-diciembre 2025 (Publicado Ago. 20, 2025)

Family Lycaenidae [Leach], [1815])

Subfamily Theclinae Swainson, 1831

Tribe Eumaeini Doubleday, 1847

Subtribe Eumaeina Doubleday, 1847

Eumaeus Hübner, [1819]

1. Eumaeus childrenae (Gray, 1832)

2. Eumaeus toxea (Godart, [1824])

Micandra Staudinger, 1888

3. Micandra cyda (Godman & Salvin, 1887)

Subtribe Rhammina Prieto & Busby, 2021

Lathecla Robbins, 2004

4. Lathecla latagus (Godman & Salvin, 1887)

Subtribe Timaetina Prieto & Busby, 2021

Nesiostrymon Clench, [1964]

5. Nesiostrymon dodava (Hewitson, 1877)

Subtribe Atlidina Martins & Duarte, 2021

Brangas Hübner, [1819]

6. Brangas neora (Hewitson, 1867)

7. Brangas getus (Fabricius, 1787)

Atlides Hübner, [1819]

8. Atlides halesus (Cramer, 1777)

9. Atlides gaumeri (Godman, 1901)

10. Atlides polybe (Linnaeus, 1763)

11. Atlides inachus (Cramer, 1775)

12. Atlides carpasia (Hewitson, 1868)

Arcas Swainson, 1832

13. Arcas cypria (Geyer, 1837)

Pseudolycaena Wallengren, 1858

14. Pseudolycaena damo (H. Druce, 1875)

Denivia K. Johnson, 1992

15. Denivia theocritus (Fabricius, 1793)

16. Denivia hisbon (Godman & Salvin, 1887)

Subtribe Evenina Faynel & Grishin, 2021

Evenus Hübner, [1819]

17. Evenus regalis (Cramer, 1775)

Subtribe Jantheclina Robbins & Faynel, 2021

Contrafacia K. Johnson, 1989

18. Contrafacia bassania (Hewitson, 1868)

19. Contrafacia ahola (Hewitson, 1867)

Allosmaitia Clench, [1964]

20. Allosmaitia strophius (Godart, [1824])

200 specimen records from the Loxicha region

between 1968 and 1992 from 100 to 2 280 m

elevation. From these historic specimens, 41

genera were recorded, with five Polyommatinae

species and 53 Theclinae species. As a result

of the faunistic work carried out for this study

from 2005 to 2014, an additional 4 547 speci-

mens were sampled, belonging to 67 genera

with nine species of Polyommatinae and 114 of

Theclinae (from 80 to 2 850 m elevation) (Fig.

2, Fig. 3, Fig. 4). A nomenclatural list of the

species follows.

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Laothus K. Johnson, Kruse & Kroenlein, 1997

21. Laothus erybathis (Hewitson, 1867)

Subtribe Paiwarriina Lamas & Robbins, 2021

Kolana Robbins, 2004

22. Kolana ligurina (Hewitson, 1874)

Subtribe Cupatheclina Lamas & Grishin, 2021

Cupathecla Robbins, 2002

23. Cupathecla cupentus (Stoll, 1781)

Subtribe Parrhasiina Busby & Robbins, 2021

Parrhasius Hübner, [1819]

24. Parrhasius polibetes (Stoll, 1781)

25. Parrhasius orgia (Hewitson, 1867)

26. Parrhasius moctezuma (Clench, 1971)

Ignata K. Johnson, 1992

27. Ignata gadira (Hewitson, 1867)

Michaelus Nicolay, 1979

28. Michaelus jebus (Godart, [1824])

29. Michaelus hecate (Godman & Salvin, 1887)

30. Michaelus ira (Hewitson, 1867)

Apuecla Robbins, 2004

31. Apuecla maeonis (Godman & Salvin, 1887)

Dicya K. Johnson, 1991

32. Dicya carnica (Hewitson, 1873)

Symbiopsis Nicolay, 1971

33. Symbiopsis sp.

Subtribe Ipideclina Martins & Grishin, 2021

Ipidecla Dyar, 1916

34. Ipidecla miadora Dyar, 1916

Subtribe Calycopidina Duarte & Robbins, 2010

Gigantorubra Johnson, 1993

35. Gigantorubra collucia (Hewitson, 1877)

Ziegleria K. Johnson, 1993

36. Ziegleria hesperitis (Butler & H. Druce, 1872)

37. Ziegleria hoffmani K. Johnson, 1993

Arzecla Duarte & Robbins, 2010

38. Arzecla ceromia (Hewitson, 1877)

Kisutam K. Johnson & Kroenlein, 1993

39. Kisutam syllis (Godman & Salvin, 1887)

Rubroserrata K. Johnson & Kroenlein, 1993

40. Rubroserrata mathewi (Hewitson, 1874)

Electrostrymon Clench, 1961

41. Electrostrymon hugon (Godart, [1824])

42. Electrostrymon joya (Dognin, 1895)

Pendantus K. Johnson & Kroenlein, 1993

43. Pendantus denarius (Butler & H. Druce, 1872)

44. Pendantus guzanta (Schaus, 1902)

Mercedes K. Johnson, 1992

45. Mercedes clarina (Hewitson, 1874)

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46. Mercedes atnius (Herrich-Schäffer, [1853])

47. Mercedes demonassa (Hewitson, 1868)

48. Mercedes calus (Godart, [1824])

Calycopis Scudder, 1876

49. Calycopis isobeon (Butler & H. Druce, 1872)

Subtribe Strymonina Tutt, 1907

Thereus Hübner, [1819]

50. Thereus orasus (Godman & Salvin, 1887)

51. Thereus oppia (Godman & Salvin, 1887)

52. Thereus ortalus (Godman & Salvin, 1887)

Rekoa Kaye, 1904

53. Rekoa meton (Cramer, 1779)

Heterosmaitia Clench, [1964]

54. Heterosmaitia palegon (Cramer, 1780)

55. Heterosmaitia zebina (Hewitson, 1869)

56. Heterosmaitia marius (Lucas, 1857)

57. Heterosmaitia stagira (Hewitson, 1867)

Arawacus Kaye, 1904

58. Arawacus togarna (Hewitson, 1867)

59. Arawacus sito (Boisduval, 1836)

60. Arawacus jada (Hewitson, 1867)

Strymon Hübner, 1818

61. Strymon melinus (Hübner, 1818)

62. Strymon albata (C. Felder & R. Felder, 1865)

63. Strymon bebrycia (Hewitson, 1868)

64. Strymon yojoa (Reakirt, [1867])

65. Strymon cestri (Reakirt, [1867])

66. Strymon bazochii (Godart, [1824])

67. Strymon istapa (Reakirt, [1867])

68. Strymon serapio (Godman & Salvin, 1887)

69. Strymon megarus (Godart, [1824])

70. Strymon ziba (Hewitson, 1868)

Panthiades Hübner, [1819]

71. Panthiades bathildis (C. Felder & R. Felder, 1865)

72. Panthiades sierrae (Dyar, 1916)

73. Panthiades ochus (Godman & Salvin, 1887)

Subtribe Strephonotina K. Johnson, Austin, Le Crom & Salazar, 1997

Tmolus Hübner, [1819]

74. Tmolus echion (Linnaeus, 1767)

75. Tmolus crolinus Butler & Druce, 1872

Nicolaea K. Johnson, 1993

76. Nicolaea dolium (H.H. Druce, 1907)

Crimsinota K. Johnson, 1993

76. Crimsinota velina (Hewitson, 1868)

Ministrymon Clench, 1961

77. Ministrymon clytie (W.H. Edwards, 1877)

78. Ministrymon arola (Hewitson, 1868)

79. Ministrymon zilda (Hewitson, 1873)

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80. Ministrymon inoa (Godman & Salvin, 1887)

81. Ministrymon phrutus (Geyer, 1832)

82. Ministrymon azia (Hewitson, 1873)

83. Ministrymon una (Hewitson, 1873)

Gargina Robbins, 2004

84. Gargina gargophia (Hewitson, 1877)

85. Gargina caninius (H.H. Druce, 1907)

86. Gargina thoria (Hewitson, 1869)

Siderus Kaye, 1904

87. Siderus philinna (Hewitson, 1868)

Theclopsis Godman & Salvin, 1887

88. Theclopsis mycon (Godman & Salvin, 1887)

Ostrinotes K. Johnson, Austin, Le Crom & Salazar, 1997

89. Ostrinotes keila (Hewitson, 1869)

Strephonota K. Johnson, Austin, Le Crom & Salazar, 1997

90. Strephonota tephraeus (Geyer, 1837)

91. Strephonota syedra (Hewitson, 1867)

Celmia K. Johnson, 1991

92. Celmia celmus (Cramer, 1775)

Subtribe Callophryidina Tutt, 1907

Radissima Johnson, 1992

94. Radissima umbratus (Geyer, 1837)

Thaeides K. Johnson, Kruse & Kroenlein, 1997

95. Thaeides theia (Hewitson, 1870)

Ocaria Clench, 1970

96. Ocaria petelina (Hewitson, 1877)

97. Ocaria ocrisia (Hewitson, 1868)

Chlorostrymon Clench, 1961

98. Chlorostrymon simaethis (Drury, 1773)

99. Chlorostrymon telea (Hewitson, 1868)

Magnastigma Nicolay, 1977

100. Magnastigma elsa (Hewitson, 1877)

Cyanophrys Clench, 1961

101. Cyanophrys fusius (Godman & Salvin, 1887)

102. Cyanophrys herodotus (Fabricius, 1793)

103. Cyanophrys miserabilis (Clench, 1946)

104. Cyanophrys longula (Hewitson, 1868)

105. Cyanophrys agricolor (Butler & H. Druce, 1872)

Callophrys Billberg, 1820

106. Callophrys guatemalena Clench, 1981

Erora Scudder, 1872

107. Erora quaderna (Hewitson, 1868)

108. Erora nitetis (Godman & Salvin, 1887)

109. Erora carla (Schaus, 1902)

110. Erora gabina (Godman & Salvin, 1887)

111. Erora opisena (H.H. Druce, 1912)

112. Erora muridosca (Dyar, 1918)

Semonina Robbins, 2004

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Fig. 2. Lycaenidae from the Loxicha Region. The numbers correspond to the species list (Line: scale 1 cm). Only species not

shown in Fig. 8 and Fig. 9 are included.

113. Semonina semones (Godman & Salvin, 1887)

Chalybs Hübner, [1819]

114. Chalybs hassan (Stoll, 1790)

Subfamily Polyommatinae Swainson, 1827

Leptotes Scudder, 1876

115. Leptotes cassius cassidula (Boisduval, 1870)

116. Leptotes marina (Reakirt, 1868)

Zizula Chapman, 1910

117. Zizula cyna (W.H. Edwards, 1881)

Brephidium Scudder, 1876

118. Brephidium exilis exilis (Boisduval, 1852)

Cupido Schrank, 1801

119. Cupido comyntas (Godart, [1824])

Celastrina Tutt, 1906

120. Celastrina echo gozora (Boisduval, 1870)

Hemiargus Hübner, 1818

121. Hemiargus ceraunus astenidas (Lucas, 1857)

Echinargus Nabokov, 1945

122. Echinargus isola (Reakirt, [1867])

Icaricia Nabokov, 1945

123. Icaricia acmon (Westwood, [1851])

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Encounter rates: Most species sampled

were rarely encountered, which is a widely

observed pattern among animals and plants

(Kunin & Gaston, 1993). Thirty-five species (28

%) were sampled once or twice, representing an

encounter rate of less than 0.03 %. Alternatively,

the ten most commonly encountered species

made up 51 % of the individuals, namely

Cupido comyntas (413 specimens), Leptotes

cassius cassidula (378), Celastrina echo gozora

Fig. 4. Lycaenidae from the Loxicha Region (continued). The numbers correspond to the species list (Line: scale 1 cm).

Fig. 3. Lycaenidae from the Loxicha Region (continued). The numbers correspond to the species list (Line: scale 1 cm).

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(323), Calycopis isobeon (262), Pseudolycae-

na damo (242), Hemiargus ceraunus astenidas

(222), Theclopsis mycon (169), Ministrymon

arola (157), Arawacus sito (148), and Zizula

cyna (122). Half the species are Polyommatinae

and half are Theclinae.

Patterns of species richness: The Lycae-

nidae richness of the Loxicha region represents

53 % of the Mexican fauna (Llorente-Bousquets

et al., 2014) and 71 % at the state level (Luis-

Martínez et al., 2016). The richness of the

Loxicha region is equivalent to that of the

Central Chiapas Massif (de la Maza & de la

Maza, 2015) and is slightly less than the Los

Tuxtlas-Catemaco region, Veracruz (with the

highest richness: 137 species) and the Sierra

de Juárez, Oaxaca (125 species) (Luis-Martínez

et al., 1991). Llorente-Bousquets et al. (2014)

reported 106 Lycaenidae species for the Sierra

Madre del Sur, with 17 species fewer than the

Loxicha region. Loxicha hosts four of the eight

Mexican endemic Theclinae species: Ipidecla

miadora (7 specimens), Laothus erybathis (43),

Semonina semones (4), and Symbiopsis sp. (2)

(Trujano-Ortega et al., 2024).

The Loxicha locality with the highest rich-

ness was Rancho Hagia Sofía, with 84 Thecli-

nae and seven Polyommatinae (Table 2). The

sample included 80 % of the species and 45

% of the specimens in the Loxicha region. It

ranks fourth among the 18 most species-rich

localities in Mexico, and is the richest on the

Pacific slope, surpassing Rancho “El Zorrillo”

and Cañada Húmeda in Michoacán by 11 spe-

cies (Table 1). The locality also had the greatest

sampling effort with 47 fieldwork days.

A contrast to Rancho Hagia Sofía is Azulil-

lo, with an equivalent number of vegetation and

elevational zones. During 43 days of sampling

for Theclinae, we recorded about half as many

species and 12 % of the specimens compared to

Rancho Hagia Sofía (Table 2). In contrast, for

Polyommatinae, Azulillo had one more species

than Rancho Hagia Sofía and nearly the same

number of specimens.

The remaining 20 localities complete the

richness of the Loxicha region with 33 species

and 55 % of the specimens. Notably, Azulillo

and Pluma Hidalgo (4 km NW “La Curva”)

show similar richness with 45 and 43 species,

respectively, even though Azulillo is located at

lower elevation (Table 2). The locality of San

José del Pacífico stands out for its richness with

38 species; it had the second-highest number

of specimens after Rancho Hagia Sofía and is

the richest among the localities above 2 000 m

in Mexico (e.g., Luis-Martínez & Llorente-

Bousquets, 1990, Luis-Martínez & Llorente-

Bousquets, 1993; Luis-Martínez et al. 1991;

Vargas-Fernández et al., 1994; Vargas-Fernán-

dez et al., 1999).

Phenology: Both abundance and estimat-

ed richness show the same annual variation

(Fig. 5, SMT 1). According to the estimates,

richness is highest at the beginning of the rainy

season (May-June), remaining consistent with

no significant differences until October (Fig. 5,

SMT 1). At the start of the dry season (Novem-

ber), there is a significant decrease in both rich-

ness and abundance, with minimum values in

February. The estimates indicate sufficient sam-

pling for most months, although the confidence

intervals are wide, particularly for December,

due to lower abundance and a higher propor-

tion of rarely encountered species.

Vegetation type: The greatest number of

Loxicha Lycaenidae inhabit tropical subdecidu-

ous forest (124 estimated species) and montane

cloud forest (106 estimated species) (Fig. 6,

SMT 1). High-elevation oak-pine and montane

cloud forest are less rich (62 estimated species).

The wide confidence intervals for richness

estimates in tropical deciduous forest, tropical

subdeciduous forest, and high-altitude oak-

pine and montane cloud forest are due to the

high frequency of rarely encountered species.

In contrast, no species were rarely encountered

in oak-pine forest, and the confidence interval

was low.

Elevation: Lower elevations (0-500 m) with

103 estimated species and the 1 000-1 500 m

zone with 123 species had the highest estimated

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richness (Fig. 7). The 0-500 m range is the best

sampled, with 91 % of the species sampled,

according to the estimation. In contrast, for

the 1 000-1 500 m range, only 56 % of the

species were sampled. Additionally, the wide

confidence interval at this elevation (a high fre-

quency of rarely encountered species) may be

due to the montane cloud forest. On the other

hand, ecological diversity (Effective Number

of Species) is high at the lowest elevations and

decreases with elevation (Fig. 7). This ecologi-

cal diversity is affected by the proportion of rare

and dominant species present in each elevation

zone. In general, encounter rates are low, with

Fig. 5. Observed species richness, estimated richness, and abundance of Lycaenidae by month. Both parameters show a peak

at the beginning of the rainy season in May and a decrease at the start of the dry season in November. The estimated richness

indicates good sampling levels for each month, except December.

Fig. 6. Estimation of species richness by vegetation type. CF shows high richness considering it covers a smaller area. The

letters indicate significant differences (p < 0.05) in the estimation of species richness based on the overlap of confidence

intervals. TDF: tropical deciduous forest; TSDF: tropical subdeciduous forest; CF: cloud forest; OPCF: oak-pine – cloud

forest; OPF: oak-pine forest.

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33 % of the 123 sampled species represented

by five or fewer specimens. We note that the

elevation range with the fewest “unseen” spe-

cies is 1 500-2 000 m, which may be a sampling

artifact because only one locality occurs at this

elevation (Table 2).

Exclusive and characteristic species by

vegetation type: Forty-four species (Table 3,

Fig. 8) were sampled in only one vegetation

type, but 25 were represented by only one or

two specimens (Table 3). The exclusive species

are all Theclinae species, with a majority (29

species, 65 %) limited to tropical subdeciduous

forest. The most abundant exclusive species in

tropical subdeciduous forest are Rekoa meton

(37 specimens), Gargina thoria (24), and G.

caninius (17). In montane cloud forest, 12

exclusive species were recorded, with the most

abundant being Arawacus togarna (5). In Oak-

Pine cloud forest, only three exclusive species

were present: Erora quaderna, E. nitetis, and

Thereus ortalus.

For characteristic species (80 to 99 %

of specimens in a specific vegetation type),

tropical subdeciduous forest has the highest

number, with 26 species, followed by montane

cloud forest with six species and oak-pine cloud

forest with five (Table 3, Fig. 9). Three Poly-

ommatinae species were each characteristic of

one vegetation type: Icaricia acmon (tropical

subdeciduous forest), Zizula cyna (montane

cloud forest), and Celastrina echo gozora (high-

altitude oak-pine and montane cloud forest).

Despite its small extent, the montane cloud

forest had 12 exclusive and six characteris-

tic species, with the most frequently encoun-

tered being Zizula cyna, Pendantus denarius,

Eumaeus toxea, and E. childrenae. The other

two vegetation types (tropical deciduous forest

and oak-pine forest) did not have exclusive or

characteristic species.

Community composition: The distance

matrix dendrogram based on species abun-

dance data results in three groupings of fau-

nal similarity characterized by elevation and

vegetation type (Fig. 10). The first group con-

sists of seven localities above 1 800 m eleva-

tion, including the two highest localities in the

Fig. 7. Species richness and diversity by elevation. Observed species richness is higher in the lowest elevational range,

although the estimation shows no differences with intermediate altitudes, which had lower sampling effort. Both richness and

ecological diversity decrease with elevation. Letters indicate significant differences (p < 0.05) in species richness estimation,

based on the overlap of confidence intervals.

16 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61cmxq06, enero-diciembre 2025 (Publicado Ago. 20, 2025)

Fig. 8. Distribution of ‘exclusive’ species recorded only in one type of vegetation. The numbers correspond to the species list.

Table 3

‘Exclusive’ and ‘characteristic’ species by vegetation type.

Tropical

Subdeciduous

Forest

Atlides carpasia, Allosmaitia strophius, Arzecla ceromia (13), Atlides gaumeri (9), Atlides polybe (11),

Brangas getus (7), Brangas neora (1), Cyanophrys fusius, Chalybs hassan, Cyanophrys herodotus, Cyanophrys

miserabilis, Chlorostrymon telea, Cupathecla cupentus (1), Denivia theocritus, Erora carla, Electrostrymon

hugon, Electrostrymon joya (2), Erora gabina (2), Erora muridosca (2), Erora opisena (6), Evenus regalis (12),

Gigantorubra collucia, Gargina caninius (17), Gargina thoria (24), Heterosmaitia marius, Heterosmaitia

stagira, Rekoa meton (37), Heterosmaitia zebina (14), Icaricia acmon, Ipidecla miadora, Ignata gadira (7),

Kisutam syllis (1), Lathecla latagus (1), Ministrymon arola, Mercedes clarina, Mercedes demonassa, Michaelus

jebus, Ministrymon phrutus, Magnastigma elsa (2), Mercedes calus (2), Michaelus hecate (14), Ministrymon

inoa (1), Ministrymon zilda (2), Nicolaea dolium (11), Pseudolycaena damo, Panthiades sierrae, Parrhasius

orgia (6), Radissima umbratus, Siderus philinna, Strephonota tephraeus, Semonina semones (4), Strymon

melinus (2), Symbiopsis sp. (2), Tmolus echion, Ziegleria hesperitis (1)

Cloud forest

Arawacus togarna (5), Atlides inachus (1), Celmia celmus (1), Contrafacia bassania (2), Denivia hisbon

(3), Eumaeus childrenae, Eumaeus toxea, Mercedes atnius (1), Ministrymon una (1), Nesiostrymon dodava

(1), Ocaria petelina (1), Panthiades ochus, Pendantus denarius, Strephonota syedra (1), Thaeides theia (1),

Theorema eumenia (1), Thereus orasus, Zizula cyna

Oak-Pine

cloud forest

Arawacus jada, Atlides halesus, C. guatemalena, Celastrina echo gozora, Erora nitetis (2), Erora quaderna (8),

Parrhasius moctezuma, Thereus ortalus (1)

‘Characteristic’ species (underlined) have 80 to 99 % of their records in one type of vegetation. The number of records for the

‘exclusive’ species is indicated in parentheses.

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Loxicha region and four localities in Omiltemi.

In addition to elevation, this group is also char-

acterized by the predominance of oak forests

and high-altitude oak-pine, which are environ-

ments exclusive to this group.

The second group is characterized by the

presence of tropical subdeciduous forest and

tropical deciduous forest. Eight localities occur

under 1 000 m elevation, but there are also nine

localities at this elevation in the third group.

In other words, vegetation type explains more

than elevation explains similarity between

localities in the second group.

The third group includes the five localities

from the Sierra de Atoyac, nine from the Loxi-

cha region, and Los Mazos from the Sierra de

Manantlán region. This group contains locali-

ties between 1 000 and 1 500 m mixed with

lower-altitude localities. The common char-

acteristic of 12 of the 16 localities is the pres-

ence of montane cloud forest. Thus, similar to

group 2, the similarity among these localities,

which belong to three different regions, are

mainly determined by vegetation type.

DISCUSSION

Species richness: The Loxicha region

fauna represents about 11 % of the Neotropical

Eumaeini fauna (Robbins, 2004). It is the high-

est documented Lycaenidae species richness on

Mexico’s Pacific slope, as was the case for other

butterfly families (Arellano-Covarrubias et al.,

2018; Luis-Martínez et al., 2016, Luis-Martínez

et al., 2020; Luis-Martínez et al., 2022). The

only Mexican regions with a greater Lycaenidae

species richness are Los Tuxtlas-Catemaco in

Veracruz and the Sierra de Juárez in Oaxaca on

the Atlantic slope, both regions with well-doc-

umented sampling during the 1960s to 1980s

(Luis-Martínez et al., 1991; Raguso & Llorente-

Bousquets, 1991; Ross, 1966; Ross, 1976).

Fig. 9. Distribution of ‘characteristic’ species, which account for 80 to 99 % of their records in one type of vegetation. The

numbers correspond to the species list.

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Among the Loxicha region localities, Hagia

Sofía stands out for its richness, accounting for

73 % of the species. It has a diversity similar to

less-studied entire regions. The high richness

at Hagia Sofía may be due to the prevalence of

areas suitable for adult foraging, as previously

noted by Arellano-Covarrubias et al. (2018)

for Riodinidae. At this site, inflorescences of

the introduced rambutan (Nephelium lappace-

um L.) and Indian almond trees (Terminalia

catappa L.) were major nectar sources. We sus-

pect that the differences in Theclinae diver-

sity between Hagia Sofía and Azulillo – similar

environments and sampling effort – are due to

the presence of these trees at the former locality.

The high species richness of the Loxicha

region on its Pacific slope is the result of its

biogeographic history, altitudinal-vegetational

heterogeneity, and its location within the Sierra

Madre del Sur. The Sierra Madre del Sur is

Fig. 10. Similarity in the composition of Lycaenidae species from the Pacific slope. Three groups of localities are recognized,

characterized by the altitude and the type of vegetation they present. Vegetation types: TDF, tropical deciduous forest; TSDF,

tropical subdeciduous forest; CF, cloud forest; OF, oak forest; OPCF, oak-pine – cloud forest. Regions: ATO, Atoyac; IXCA,

Ixcateopan; LOX, Loxicha; MAN, Manantlán; OMI, Omiltemi. Localities: Azuli, Azulillo; CLoxi, Candelaria Loxicha; Copal,

Copalita Siete Veneros; FAuro, Finca Aurora-Finca San Isidro; Pasio, La Pasionaria; Soled, La Soledad-Buenavista; Magda,

Magdalena El Lirio; PNHua, Parque Nacional Huatulco; PNHRC, Parque Nacional Huatulco Río Cacaluta; PluHi, Pluma

Hidalgo 4 km NW “La Curva”; PorRa, Portillo del Rayo-Finca El Encanto; Guajo, Puente Arroyo “El_Guajolote”; RhagS,

Rancho Hagía Sofía; RMoli, Río Molinos; SJPac, San José del Pacífico 1 km S; Faisa, El Faisanal; Parot, Las Parotas; NDelh,

Nueva Delhi; PLuga, Puente de los Lugardo; RSant, Río Santiago 4 km W; GBorr, Cañada W de la Gruta del Borrego; Potre,

Potrerillo; LPerr, Presa de la Perra; ROmil, Rancho Omiltemi; Ixcat, Ixcateopan de Cuauhtémoc; AguDu, Agua Dulce; Ahuac,

Ahuacapan; Caler, La Calera; Mazos, Los Mazos 9 mi SW Autlán; Plata, Platanarillo; Zenzo, Zenzontla.

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recognized as a region of high diversity due to

its complex geological history and altitudinal

gradient that generates ecosystem heterogene-

ity (Espinosa et al., 2016). This is reflected in

the Loxicha region, which includes five types of

vegetation. Therefore, the altitudinal gradient

and environmental heterogeneity of the Loxi-

cha region explain the high butterfly species

richness, particularly that of Lycaenidae.

Encounter rates: For the Loxicha region,

54 species (44 % of the fauna) were repre-

sented by 10 or fewer individuals. This result

is frequent in faunistic studies of hyperdiverse

groups and is pronounced in Theclinae (Arella-

no-Covarrubias et al., 2018; Lamas et al., 2021;

Luis-Martínez & Llorente-Bousquets, 1990;

Luis-Martínez et al., 1991; Vargas-Fernández

et al., 1994; Vargas-Fernández et al., 1999). The

significance of this low encounter rate for many

species is that it indicates a large number of

“unseen” species. Statistically, this significance

is reflected in large species richness estimate

intervals. In sharp contrast to the Theclinae,

most species of Polyommatinae were frequently

encountered, perhaps because they tend to

inhabit highly disturbed habitats.

Of the species previously recorded for the

Loxicha region but not sampled during field-

work, four were recorded from a single speci-

men in La Soledad – Buenavista during the

1970s and 1980s (Atlides inachus, Ministrymon

una, Nesiostrymon dodava, and Strephonota

syedra). Low encounter rates are a probable

explanation for the result, but we note the pos-

sibility that this locality was confused with La

Soledad in the Sierra de Juárez region on the

Gulf slope.

Phenology: The seasonal distribution of

Lycaenidae in the Loxicha region is similar to

that for Papilionidae, Pieridae, and Riodinidae

(Arellano-Covarrubias et al., 2018; Luis-Mar-

tínez et al., 2020; Luis-Martínez et al., 2022).

In all families, a marked decrease in richness

and abundance occurs at the transition from

the wet to the dry season (November). Alter-

natively, greatest richness and abundance occur

at the transition from the dry to the wet season

in May and June except in Nymphalidae, with

greatest richness in September and October

(Luis-Martínez et al., 2022).

At other Neotropical sites, Lycaenidae rich-

ness is also lowest at the transition from wet to

dry seasons. It may be greatest during the dry

season (Bates, 1864; Robbins & Small, 1981),

during the wet season (Prieto & Dahners,

2009), or at the transition from dry to wet sea-

sons (Lamas et al., 2021; Robbins et al., 1996).

Vegetation types: Lycaenidae distribu-

tion by vegetation type shows similarities with

butterfly families Papilionidae, Pieridae, and

Nymphalidae (Luis-Martínez et al., 2020; Luis-

Martínez et al., 2022). All are most diverse

in tropical subdeciduous forest and montane

cloud forest and are least diverse in oak-pine

forest. Tropical subdeciduous forest and mon-

tane cloud forest have the most exclusive and

characteristic Lycaenidae species, as is the case

with Papilionidae and Nymphalidae (Luis-Mar-

tínez et al., 2020; Luis-Martínez et al., 2022).

In contrast to these patterns of butterfly

diversity in the Sierra Madre del Sur, ferns

(Tejero-Díez et al., 2016), gymnosperms

(Contreras-Medina, 2016), Quercus (Valencia-

Ávalos & Morales-Saldaña, 2016), and Salvia

(Martínez-Gordillo et al., 2016) have greatest

richness in oak-pine forest and montane cloud

forest but lower richness in tropical subdecidu-

ous forest. In other words, in the Sierra Madre

del Sur, it would appear that plant diversity,

albeit a small sample of plants, need not beget

butterfly diversity.

Elevation: Species richness of Lycaenidae

in the Loxicha region is greatest at lower eleva-

tions, similar to Riodinidae, Papilionidae, Pieri-

dae, and Nymphalidae (Arellano-Covarrubias

et al., 2018; Luis-Martínez et al., 2020; Luis-

Martínez et al., 2022). More generally, but-

terfly richness decreases as elevation increases

(Álvarez-García et al., 2016; Lamas et al., 2021;

Luis-Martínez & Llorente-Bousquets, 1990;

Luis-Martínez & Llorente-Bousquets, 1993;

Luis-Martínez et al., 1991; Vargas-Fernández

20 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61cmxq06, enero-diciembre 2025 (Publicado Ago. 20, 2025)

et al., 1994; Vargas-Fernández et al., 1999). In

contrast, butterflies from Serra do Cipó, Brazil

(Pires et al., 2020) and Geometridae from Costa

Rica (Brehm et al., 2007) show greater diversity

at intermediate elevations. McCoy (1990) found

that short-term studies find higher diversity

at intermediate elevations, while longer stud-

ies, like this one for Lycaenidae, find greater

diversity at lower elevations. Given the broad

estimates of species richness (Fig. 7), definitive

conclusions would require more sampling.

Species composition similarity: The

three main community similarity groupings for

lycaenids in the Sierra Madre del Sur and the

Pacific slope are, as expected, correlated with

elevation and vegetation type (Fig. 10). How-

ever, the substantial variability of elevation and

vegetation type within each similarity grouping

is perhaps unexpected. In other words, eleva-

tion and vegetation type do not definitively

predict the lycaenid community that will occur

in a habitat. Rather, groupings appear to also be

explained by isolation of faunas at higher eleva-

tions, as found in other Papilionoidea (s.l.),

of mountainous systems (Llorente-Bousquets,

1984; Llorente-Bousquets & Escalante, 1994;

Llorente-Bousquets & Luis-Martínez, 1998).

This is the fourth publication on the lepi-

dopteran fauna of the Loxicha Region (Arella-

no-Covarrubias et al., 2018; Luis-Martínez et

al., 2020; Luis-Martínez et al., 2022), confirm-

ing that for the five Papilionoidea families ana-

lyzed, it is one of the most diverse regions in the

country. Systematic faunistic studies with ade-

quate sampling provide a foundation for inves-

tigating biological processes. They allow for the

recognition of disjunct distributions in moun-

tainous systems, and with that, processes of

speciation and endemism. They can also serve

as the basis for measuring changes in species

distribution due to climate change, especially

for stenotopic species that could lead to local

extinctions (e.g. Molina-Martínez et al., 2016).

Lycaenidae has historically received less

attention than other butterfly families in Mex-

ico. The geographic, seasonal, elevational, and

vegetational distribution of Lycaenidae in the

Loxicha region is the most comprehensive to

date for Mexican Lycaenidae. The high propor-

tion of rarely encountered species is a notable

characteristic of Lycaenidae, but many seasonal,

elevational, and vegetation-associated patterns

that we document are similar to those of other

butterfly families.

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.

See supplementary material

a48v73n1-suppl. 1

ACKNOWLEDGMENTS

Authors thank CONABIO, which in its

three decades of existence have supported the

Lepidoptera collection, granting resources for

infrastructure, fieldwork, and data repatriation

from specimens in USA collections (1991-

2000) and now included in the MARIPOSA

database. Also, we thank the research proj-

ect PAPIIT-IN212925, for financial support.

MTO thanks the support from CONAHCYT

(Estancias Posdoctorales por México para la

Formación y Consolidación de las y los Investi-

gadores por México; CVU131802). To the cura-

tors of the collections who granted access to

specimens, data and specialized literature dur-

ing the development of the MARIPOSA data-

base: Frederick Rindge † (American Museum

of Natural History, Nueva York), Jerry Powell

and John Chemsak (“Essig” Collection Depart-

ment of Entomological Sciences, University of

California, Berkeley campus, California), John

E. Rawlins ✝ (Carnegie Museum of Natural

History, Pittsburgh, Pennsylvania), David K.

Faulkner and John W. Brown (San Diego Natu-

ral History Museum, California), Paul Arnaud

Jr. and Norman Penny (California Academy of

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61cmxq06, enero-diciembre 2025 (Publicado Ago. 20, 2025)

Sciences, San Francisco, California), Brian V.

Brown and Brian Harris (Los Angeles County

Museum, California), Lee D. Miller ✝ (Allyn

de Entomology Museum, Sarasota, Florida),

George T. Austin ✝ (Nevada State Museum),

Philip Ackery and Dick Vane-Wright (British

Museum of Natural History), Harry Brailovsky

(Colección Entomológica del Instituto de

Biología, UNAM, Cd. de México), Thomas

Emmel ✝ and Jacqueline Miller ✝ (Collection

of the McGuire Center for Lepidoptera and

Biodiversity, Florida Museum of Natural His-

tory, University of Florida), Carmen Pozo de la

Tijera (Colección de Lepidoptera del Colegio

de la Frontera Sur (Chetumal)) and Mercedes

Luna (Colección de Lepidoptera del Museo

de Zoología de la Facultad de Estudios Supe-

riores Zaragoza, UNAM). Our great gratitude

to the residents of Oaxaca and owners of the

lands, like Armando Canavati (Rancho Hagia

Sofía, Santa María Huatulco), who supported

us in the fieldwork. To the managers of the

natural protected area Parque Nacional Huatul-

co for their support for the fieldwork during

five years. To Isabel Vargas Fernández, Jimena

Castro, and Uri García for their help in the

laboratory tasks, as well as Alejandra Sánchez

García, Andrew D. Warren, Blanca Claudia

Hernández Mejía, Ela Stephanie Esquivel Ruíz,

Jessica Hernández Jerónimo, John Kemner, José

Luis Salinas Gutiérrez, Marisol Esther Almaraz

Almaraz, and Sandra Nieves Uribe, who par-

ticipated in the sampling. Finally, Atzinameyali

Sánchez Castañeda, who prepared and labeled

the collected specimens for their incorporation

into the museum collection.

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