363

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Distribution, diversity, endemism, and ecology of Nymphalid butterflies

(Lepidoptera: Nymphalidae) in the Loxicha Region, Oaxaca, Mexico

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

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

Marysol Trujano-Ortega1, 2; https://orcid.org/0000-0001-8911-8504

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

Isabel Vargas-Fernández1; https://orcid.org/0000-0001-6524-7184

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

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

Autónoma de México, Av. Universidad 3000, Circuito Exterior S/N, C.P. 04510, Ciudad de México, México; alm@

ciencias.unam.mx, omaravalosh@ciencias.unam.mx, marysol_trujano@yahoo.com.mx, arellano.covarrubias@gmail.

com, ivf@ciencias.unam.mx, llorentebousquets@gmail.com (* Correspondence).

2. Facultad de Estudios Superiores Zaragoza, Universidad Nacional Autónoma de México (UNAM), Batalla 5 de Mayo

s/n, Ejercito de Oriente, Iztapalapa, 09230, Ciudad de México, México.

Received 26-X-2021. Corrected 14-III-2022. Accepted 27-V-2022.

ABSTRACT

Introduction: The Loxicha Region of Oaxaca, Mexico, has been historically important for the study of

Nymphalidae, second in the Papilionoidea for species richness. Describing the diversity patterns of this butterfly

clade in Loxicha can improve our understanding of the evolutionary history of the Sierra Madre del Sur, the

Mexican Pacific slope, and Mexico in general.

Objective: To describe the temporal and spatial patterns of Nymphalidae diversity along an elevational gradient

(80-2 600 m), and to compare Loxicha’s fauna with other regions in Mexico.

Methods: We obtained 28 756 records from 21 sites in the Loxicha Region, representing seven years of sam-

pling. We estimate and analyze the diversity, endemism, and distributional patterns for three elevational levels

and five vegetation types. We estimated species composition and similarity with other regions of the Pacific and

Atlantic slopes.

Results: We identified 189 taxa, including species and subspecies, from 85 genera and ten subfamilies of

Nymphalidae. Loxicha contains 46 % of the species in the family recognized for Mexico, including ten endemic

species and 56 endemic subspecies. Cloud forest and low elevations were the most diverse habitats for this

family. There is a clear divergence between the Atlantic and Pacific faunas, and the Sierra Madre del Sur has

two faunal components. High-elevation sites in Oaxaca, and in the neighboring state of Guerrero, have a distinc-

tive fauna, apparently isolated from low-elevation sites, revealing an archipelagic distribution for cloud forest

Nymphalidae.

Conclusions: The Loxicha Region is one of the richest areas for Nymphalidae in Mexico. Distribution on the

Pacific slope is determined by geographical history and ecological conditions, including elevation. Nymphalidae

can be used to test hypotheses of biogeographic regionalization in Mexico.

Key words: Brush-footed butterflies; elevational gradient; sampling efficiency; phenology; biogeographic

provinces; Van Someren-Rydon traps.

https://doi.org/10.15517/rev.biol.trop..v70i1.48821

INVERTEBRATE BIOLOGY

364 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Brush-footed butterflies (Nymphalidae)

represent the second most diverse family of

Papilionoidea worldwide, behind the skip-

pers (Hesperiidae). Because of the remarkable

diversity and local endemism featured within

Nymphalidae, the description and analysis of

spatio-temporal patterns of this group are a pri-

ority for advancing conservation and systemat-

ics. This is particularly true for Mexico, which is

a megadiverse country and biodiversity hotspot

where Nymphalidae constitute 25 % (413 spe-

cies) of the country’s Papilionoidea fauna sensu

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

Oaxaca is one of the most biodiverse states

in Mexico (Flores-Villela & García-Vázquez,

2014; García-Mendoza & Meave del Castillo,

2011; Navarro-Sigüenza et al., 2014), and this

pattern is reflected in Nymphalidae. Some 339

species or 82 % of the country’s nymphalids

are documented from Oaxaca, ranking it sec-

ond only to the state of Chiapas in nymphalid

species richness (Luis-Martínez et al., 2016).

This outstanding biodiversity in Oaxaca is

attributable to the convergence of five different

biogeographic provinces in the state, with the

most well-represented of these being the Sierra

Madre del Sur, Costa del Pacífico, and Golfo de

México (Morrone, 2005; Morrone et al., 2002).

The Loxicha Region of Oaxaca lies within

the Sierra Madre del Sur, and it is of special

faunal interest. Loxicha experienced two main

collecting periods in the 20th century, during

which numerous species of Nymphalidae were

described and named (Luis-Martínez et al.,

2020). The MARIPOSA database (Luis-Mar-

tínez et al., 2005), contains 1 248 records from

the Loxicha Region during that century, with

139 species of Nymphalidae being documented

by more than 30 collectors. Particularly notable

among those collectors were Eduardo Cecilio

Welling with 66 species (265 records), and

John Kemner with 78 species (402 records).

Modern description and analysis of the

Papilionoidea from the Pacific slope of the

Loxicha Region has been the core focus of an

ongoing project, begun in 2005, to analyze the

butterfly fauna of Oaxaca (Luis-Martínez et

al., 2016). To date, the families Papilionidae,

Pieridae, and Riodinidae of the Loxicha Region

have been analyzed (Arellano-Covarrubias et

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

present study aims to describe the temporal and

spatial patterns of Nymphalidae diversity in

the Loxicha Region. Based on a review of the

literature and historical records, coupled with

sampling from 2005-2014 along an elevational

gradient from 80–2 600 m, we document this

diversity and make a preliminary comparison

of nymphalids in Loxicha to other compara-

tively well-sampled regions to describe broad-

scale patterns.

Historical outline of the collection of

Papilionoidea from the Pacific slope and the

Loxicha Region: The historical collection and

classification of Mexican species of Papilionoi-

dea began at the end of the 19th century and

continued through the early 1980’s. Collectors

generally focused on the Southeastern states,

the Atlantic slope, and the states of Guerrero

and Morelos due to the high diversity of these

areas and to the presence of associated high-

ways (v. gr. Llorente-Bousquets et al., 1986;

Llorente-Bousquets & Luis-Martínez, 1993;

Llorente-Bousquets et al.,1998; Luis-Martínez

et al., 2000; Luis-Martínez et al., 2003; Michán

et al., 2004). During this period, sporadic col-

lections were also made in Northern Mexico,

mostly carried out by academic institutions of

the USA (v. gr. Comstock, 1953; Holland, 1995;

Spade et al., 1988; Stanford, 1998). The Pacific

slope of Mexico was often ignored by collec-

tors, resulting in relatively few publications

except for a butterfly checklist for the Cha-

mela Biological Research Station in the state

of Jalisco (Beutelspacher, 1982), notes on the

butterflies of some Pacific islands (Vázquez,

1958; Vázquez, 1959; Vázquez, 1960), and a

contribution on the fauna of the Pacific slope

of Jalisco (v. gr. Comstock & Vázquez, 1961).

Additionally, Members of the Mexican Soci-

ety of Lepidopterology published the results

of periodic collections along this slope (v. gr.

Maza & Maza, 1981; Maza & Maza, 1983;

Maza et al., 1982; Velázquez, 1976; Velázquez

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& Velázquez, 1975), with the main goal of

describing new taxa from unexplored areas.

The first systematic sampling of the Pacif-

ic slope extended from the coast (Puerto San

Blas) to the mountains (Sierra San Juan) in the

state of Nayarit during 1978-1980, spanning an

elevational gradient from sea level to 1 300 m.

This effort documented 135 species of Nym-

phalidae, marking the beginning of dedicated

research on the distribution of Papilionoidea

from the Pacific slope. A decade later, faunistic

studies were carried out in the Sierra de Atoyac

(Vargas-Fernández et al., 1994) and the Omilt-

emi region (Luis-Martínez & Llorente-Bous-

quets, 1993) of the Sierra Madre del Sur (SMS)

in the state of Guerrero. The former spanned

an elevational range of 300-3 100 m, and the

latter a range of 2 300-3 000 m (v. gr. Llorente-

Bousquets, 1984; Llorente-Bousquets et al.,

2004; Warren & Llorente-Bousquets, 1999).

Further studies on the faunistic composi-

tion of Papilionoidea from the Pacific slope

were completed during the last three decades

in Pedernales, state of Michoacán (Balcázar,

1993); Mismaloya, Jalisco and Bahía de Ban-

deras, Nayarit (Warren & Llorente-Bousquets,

1999); the Sierra de Manantlán, Jalisco-Colima

(Vargas-Fernández et al., 1999); and West-

ern Michoacán (Luis-Martínez, 1997; Luis-

Martínez, 1999; MARIPOSA database 2001).

Additional analyses of the alpha and beta

diversity of the Nymphalidae (Tapia-Sedeño,

2013) and the lepidopterofauna of the Loxicha

Region, Oaxaca (Arellano-Covarrubias et al.,

2018; Luis-Martínez et al., 2020) were also

released. Most of these studies focused on

the local, regional, or elevational distribution

and the phenology of Papilionoidea species,

under distinct ecological conditions and across

a wide environmental gradient. Some fau-

nal comparisons between different areas have

also been made (v. gr. Monteagudo-Sabaté &

Luis-Martínez, 2013; Monteagudo-Sabaté et al.,

2001; Monteagudo-Sabaté et al., 2014).

When analyzing the fauna of the Pacific

region and the Sierra Madre del Sur, the

diversity of the state of Morelos must be

considered. This state is located mainly in the

Eje Neovolcánico and Cuenca del Balsas bio-

geographic provinces (Morrone et al., 2002).

Michoacán and Guerrero also include portions

of these two provinces. More than half of

Guerrero lies in the Cuenca del Balsas, and the

state also encompasses a large portion of the

Sierra Madre del Sur, particularly its Pacific

slopes (Luna-Reyes et al., 2012). There are 450

species or subspecies of Papilionoidea sensu

lato recorded from Morelos, classified in 214

genera. The Nymphalidae is represented in

Morelos by 147 species or subspecies, of which

42 are endemic to Mexico (Luna-Reyes, 2020).

Besides these faunistic studies over the

past three decades, species lists have been

published for 10 of the 11 states in the Pacific

slope. Table 1 presents current data for the

number of species and endemics in these Mexi-

can states according to Llorente-Bousquets et

al. (2014) and Luis-Martínez et al. (2016).

MATERIALS AND METHODS

Faunistic inventory and sites: To update

the Papilionoidea species list for Oaxaca pub-

lished by Luis-Martínez et al. (2004), systemat-

ic sampling was begun in 2005 (Luis-Martínez

et al., 2016) which increased scientific knowl-

edge of the geographical and vegetational

distribution of the group. This sampling effort

focused on areas with the greatest diversity and

endemism. The Loxicha Region is one such

area, and thus was subject to 267 sampling

days over seven years (2005, 2007, 2008,

2011-2014), including 21 sampling sites that

span eight municipalities within the Pacific

slopes of the Sierra Madre del Sur (Table 2).

In 12 of these sites, sampling was system-

atic. A total of 16 collectors participated in the

sampling effort, with 13 to 65 sampling days

per site. Sites were grouped by elevational

levels and vegetation type for comparison

and analysis purposes. Vegetation classifica-

tion followed Rzedowski (1978) and Llorente-

Bousquets (1984), with four types recognized

in the Loxicha Region: tropical deciduous

forest (TDF), tropical sub-deciduous forest

(TSDF), cloud forest (low- and mid-elevation)

366 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

(CF), and oak-pine forest (OPF). Besides these,

a fifth vegetation type was recognized in some

sites at 2 000-2 400 m elevation, denominated

as oak-pine and high-elevation cloud forest

(OPCF). OPCF is similar to OPF but has inter-

mixed elements of CF. Because the OPCF can

be easily distinguished from both the OPF and

the CF, and is located at different elevations in

the region, this vegetation type was considered

a distinct category (Arellano-Covarrubias et al.,

2018; Luis-Martínez et al., 2020).

Taxonomic determination: Taxa were

identified by comparison with specimens in the

Lepidoptera Collection of the Zoology Muse-

um, Facultad de Ciencias, Universidad Nacio-

nal Autónoma de México (MZFC), together

with the taxonomic expertise of the authors and

reference to specialized literature. The list of

species follows the taxonomic order proposed

by Vargas-Fernández et al. (2016), and Llor-

ente-Bousquets, Luis-Martínez et al. (2006),

for the Satyrinae subfamily. All specimens

were deposited in the Lepidoptera Collection

of the MZFC, which is registered at the Secre-

taría de Medio Ambiente, Recursos Naturales

y Pesca (SEMARNAP) (DFE.IN.071.0798).

Specimens were collected under scientific

collecting permit FAUT-0148 issued by the

Secretaría de Medio Ambiente y Recursos

Naturales (SEMARNAT). Collecting data were

entered into the MARIPOSA database (Luis-

Martínez et al., 2005).

Nets and Van Someren-Rydon traps:

Sampling techniques included 4-6 aerial nets

set for eight hours and 10-20 Van Someren-

Rydon traps (Rydon, 1964) placed daily 1-2.5

m high along transects, with 50 m between

each trap. Traps used a mixture of water,

brown sugar, pineapple, and banana as bait.

Transects covered distinct microhabitats (open

and closed vegetation) to obtain a more rep-

resentative sample. The number of specimens

captured with traps in each locality is presented

in Table 2.

Species richness estimation and diver-

sity analysis by vegetation type and eleva-

tional level: A species accumulation curve was

estimated using data from all specimens col-

lected. Data were randomized (500 runs) with

TABLE 1

Species richness and endemism of Nymphalidae in Mexico and its Pacific-slope states

State spp. E % Reference

Baja California 41 0 0 Brown et al., 1992

Baja California Sur 43 2 5 Brown et al., 1992

Sonora 101 6 6 Bailowitz et al., 2017

Sinaloa 106 8 8

Nayarit 144 12 8 Llorente-Bousquets et al., 2004

Jalisco 176 18 10 Llorente-Bousquets, Luis-Martínez et al., 1996; Vargas-Fernández et

al., 1996; Vargas-Fernández et al., 1999; Warren et al., 1996

Colima 139 15 11 Llorente-Bousquets, Warren et al., 1996

Michoacán 168 15 9

Guerrero 216 22 10 Vargas-Fernández et al., 1994

Oaxaca 339 31 9 Luis-Martínez et al., 2016

Chiapas 351 26 7 Maza & Maza, 1993; Luis-Martínez et al., 2011

Mexico (country) 413 123 12

spp.: species, E: taxa endemic to the state/country, %: percentage of endemics relative to the total species richness of the

state/country.

Note. Chiapas endemism is reduced because species’ ranges extend towards Central America as a biogeographical and

ecological unit; the same occurs in Baja California.

367

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EstimateS 9.1 (Colwell, 2013) and fitted to the

Clench model (Soberón & Llorente-Bousquets,

1993) to estimate species richness. Species

richness was also estimated for each vegeta-

tion type (Table 2) and three elevational levels:

0-750 m, 750-1 800 m, and 1 800-2 850 m

(Luis-Martínez et al., 2020). For the elevational

analysis, the non-parametric estimation method

Chao1 (Colwell & Coddington, 1994) was

implemented in SPADE (Chao & Shen, 2010).

For each estimation, the 95 % confidence

interval (CI) was estimated using the bootstrap

technique. Significant differences between the

species richness of distinct vegetation types or

elevational levels were evaluated by compari-

son and overlap of the confidence intervals.

Additionally, the real diversity defined as

the exponential of the Shannon-Wiener index

(H’) was estimated for each elevational level

along with the 95 % CI, using the method pro-

posed by Chao and Shen (2003) implemented

in SPADE (Chao & Shen, 2010). The scale of

the real diversity (effective number of species)

is linear; therefore, the values of each eleva-

tional level can be compared directly using the

CI overlap (Jost, 2006).

Species composition similarity analy-

sis: Similarities were estimated in species

composition between the Loxicha Region and

other regions with roughly equivalent sampling

effort on the Pacific and Atlantic slopes of

TABLE 2

Sampling sites in the Loxicha Region, Oaxaca, Mexico

Site Elevation

(m)

Geographic location Vegetation

type SE/ R /T

Lat North Long West

a. Parque Nacional Huatulco, Río Cacaluta 80-100 15°47’07” 96°10’34” TDF 16/ 1 627 /781

b. Parque Nacional Huatulco 100 15°45’20” 96°09’19” TDF 24/ 1 869 /977

c. Azulillo 380-500 15°53’25” 96°29’27” TSDF 65/ 4 929 /1 755

d. Rancho Hagia Sofía 410 15°52’01” 96°21’55” TSDF 49/ 6 672 /3 430

e. Río Molinos* 530-700 15°56’10.8” 96°30’41.8” CF 4/ 71 /30

f. Copalita, Río Copalita* 600-1 200 15°56’48” 96°20’23.43” CF 21/ 1 200 /450

g. Magdalena, El Lirio 750-900 15°55’11” 96°23’34” CF 13/ 1 934 /789

h. Copalita, Los Plátanos* 900 CF 1/ 31 /6

i. Copalita, Siete Veneros* 940 CF 24/ 1 066 /457

j. San Mateo Piñas* 1 000 15°59’57.50” 96°20’4.47” CF 27/ 426 /132

k. Pluma Hidalgo, 4 km NW “La Curva” 1 100-1 200 15°56’23” 96°25’59” CF 41/ 2 243 /528

l. Finca Aurora-Finca San Isidro 1 100-1 250 15°56’30” 96°24’13” CF 25/ 1 013 /390

m. Copalita, Llano de Ocote* 1 200 CF 17/ 434 /229

n. Portillo del Rayo-Finca El Encanto 1 200-1 530 15°58’38” 96°31’11” CF 24/ 1 218 /622

o. La Soledad-Buenavista 1 470-1 550 15°58’18” 96°31’54” CF 31/ 1 690 /574

p. La Pasionaria 1 500-1 650 15°66’09” 96°25’08” CF 15/ 1 153 /420

q. Puente Arroyo “El Guajolote” 2 020-2 150 16°03’28” 96°30’18” OPF-OPCF 17/ 274 /53

r. San José del Pacífico, 1 km S 2 280-2 400 16°09’28” 96°29’21” OPF-OPCF 33/ 771 /126

s. Manzanal-Doncella* 2 700-2 800 16°07’47.9” 96°30’12” OPF 1/ 1 /-

t. Camino a San Agustín Loxicha* 2 700-2 800 16°07’41.57” 96°29’54.2” OPF 2/ 5 /-

u. Nevería-La Ciénega* 2 820-2 850 16°12’01” 96°20’56” OPF 2/ 19 /-

* Sites without systematic sampling; SE: sampling effort (days); R: records (specimens); T: specimens collected with Van

Someren-Rydon traps; Vegetation types: tropical deciduous forest (TDF), tropical sub-deciduous forest (TSDF), cloud forest

(low- and mid-elevation) (CF), oak-pine forest (OPF), and oak-pine and high-elevation cloud forest (OPCF). (Arellano-

Covarrubias et al., 2018; Llorente-Bousquets, 1984, Luis-Martínez et al., 2020; Rzedowski, 1978). Note Van Someren-

Rydon traps were not used at sites s, t, and u.

368 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Oaxaca and other states. Besides Loxicha, our

analysis included the Sierra de Juárez, Oaxaca

(Luis-Martínez et al., 1991), the Sierra Maza-

teca, Oaxaca (Álvarez-García et al., 2016),

the Sierra de Atoyac de Álvarez, Guerrero

(Vargas-Fernández et al., 1994), the Sierra de

Manantlán, Jalisco-Colima (Vargas-Fernández

et al., 1999) and the Sierra de San Juan, Nayarit

(MARIPOSA database). A distance matrix was

estimated with presence/absence data of the

species in each region using the Jaccard index

with the “vegan” package (Oksanen et al.,

2019) in R 4.0.0 (R Core Team, 2020). Using

the same method, the similarity among 77

sites distributed across Mexico was estimated,

including multiple biogeographic provinces

and both Pacific and Atlantic slopes.

Natural history: Some biological obser-

vations made during fieldwork are presented,

which include elevational migrations and spe-

cies aggregations among other data. These

observations are valuable for describing the

fine-scale spatial and temporal distribution of

some rare or hard-to-find species and could aid

future efforts to collect these species.

RESULTS

Species list: Based on a review of pub-

lished records (1950-2004) and a query of the

MARIPOSA database up to the year 2000, 139

taxa of Nymphalidae were historically recorded

from the Loxicha Region. Our subsequent

field sampling from 2005-2014 across 21 sites

in the region (Table 2) increased that list to

189 taxa, including 10 subfamilies, 23 tribes,

15 subtribes, and 85 genera. Our sampling

effort resulted in 28 756 records plus 1 248

records from historical collections archived

in the MARIPOSA database, for a total of

30 004 records of Nymphalidae for the Loxicha

Region. Data from publications and scientific

collections are mostly from the 400-1 400 m

elevational range, while records from our field

work are distributed from 80-2 850 m. All spe-

cies and subspecies historically recorded for

the region were collected during our fieldwork

except for Adelpha donysa ssp., Pedaliodes

dejecta ssp., and Actinote guatemalena guer-

rerensis J. Maza, 1982.

The Nymphalidae species documented in

the Loxicha Region constitute 46 % of Mexi-

can nymphalid species and 56 % of those

recorded from Oaxaca (Luis-Martínez et al.,

2016). A total of 10 species and 56 subspecies

endemic to Mexico are represented in the Loxi-

cha Region. These endemic taxa were binned

into four groups according to their degree of

endemism (Fig. 1): group 1, endemic to the

Loxicha Region (5 spp.); group 2, endemic

to the Sierra Madre del Sur (13 spp.); group

3, endemic to the Pacific slope (23 spp.); and

group 4, endemic to Mexico (15 spp.). The taxa

exclusive to the Loxicha Region are Memphis

wellingi (L. Miller & J. Miller, 1976); Cyllopsis

jacquelineae (L. Miller, 1974); Callicore texa

loxicha (R. G. Maza & J. Maza, 1983); Chlo-

syne gaudialis wellingi (L. Miller & Rotger,

1979), and Altinote stratonice oaxaca (J. Miller

& L. Miller, 1979).

Species list of Nymphalidae for the Loxi-

cha Region, Oaxaca. Taxa in bold text are

endemics, and superscript numbers correspond

to their degree of endemism: 1, endemic to

the Loxicha Region; 2, endemic to the Sierra

Madre del Sur; 3, endemic to the Pacific slope;

4, endemic to Mexico.

Family NYMPHALIDAE

Rafinesque, 1815

Subfamily Libytheinae Boisduval, 1833

1. Libytheana carinenta mexicana

Michener, 1943

Subfamily Danainae Boisduval, 1833

Tribe Euploeini Herrich-Schäffer, 1849

Subtribe Itunina Reuter, 1896

2. Anetia thirza thirza Geyer, [1833]

3. Lycorea halia atergatis Doubleday,

[1847]

4. Lycorea ilione albescens (Distant, 1876)

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Tribe Danaini Boisduval, 1833

Subtribe Danaina Boisduval, 1833

5. Danaus eresimus montezuma Talbot,

1943

6. Danaus gilippus thersippus (Bates,

1863)

7. Danaus plexippus plexippus (Linnaeus,

1758)

Subfamily Ithomiinae Godman & Salvin, 1879

Tribe Tithoreini Fox, 1940

8. Aeria eurimedia pacifica Godman &

Salvin, 1879

9. Tithorea harmonia hippothous Godman

& Salvin, 1879

10. Tithorea tarricina duenna Bates, 1864

Tribe Melinaeini Clark, 1947

11. Melinaea lilis flavicans Hoffmann,

1924 3

Tribe Mechanitini Bar, 1878

12. Mechanitis lysimnia utemaia Reakirt,

1866

13. Mechanitis menapis doryssus Bates,

1864

14. Mechanitis polymnia lycidice Bates,

1864

Tribe Oleriini Fox, 1940

15. Oleria paula (Weymer, 1883)

Tribe Dircennini D’Almeida, 1941

16. Dircenna klugii klugii (Geyer, 1837)

17. Episcada salvinia portilla J. Maza &

Lamas, 1978 3

18. Pteronymia artena praedicta J. Maza

& Lamas, 1982 2

19. Pteronymia cotytto cotytto (Guérin-

Méneville, [1844])

2. Pteronymia rufocincta (Salvin, 1869) 3

Tribe Godyridini D’Almeida, 1941

21. Greta annette moschion (Godman,

1901) 3

22. Greta morgane morgane (Geyer,

1837) 3

Subfamily Charaxinae Guenée, 1865

Tribe Anaeini Reuter, 1896

23. Hypna clytemnestra mexicana Hall,

1917

24. Consul electra electra (Westwood,

1850)

25. Consul excellens genini (Le Cerf, 1922)

26. Consul fabius cecrops (Doubleday,

[1849])

27. Phantos callidryas (R. Felder, 1869)

28. Siderone galanthis ssp.

29. Zaretis ellops (Ménétriés, 1855)

30. Anaea troglodyta aidea (Guérin-

Méneville, [1844])

31. Fountainea eurypyle glanzi (Rotger,

Escalante & Coronado, 1965) 3

32. Fountainea glycerium glycerium

(Doubleday, [1849])

33. Fountainea nobilis rayoensis (J. Maza

& Díaz, 1978) 3

34. Memphis forreri (Godman & Salvin,

1884)

35. Memphis perenna perenna (Godman &

Salvin, [1884])

36. Memphis pithyusa pithyusa (R. Felder,

1869)

37. Memphis wellingi L. Miller & J.

Miller, 1976 1

Tribe Preponini Rydon, 1971

38. Archaeoprepona amphimachus baroni

J. Maza, 1982 2

39. Archaeoprepona demophon

occidentalis Stoffel & Descimon, 1974

40. Archaeoprepona demophoon mexicana

Llorente, Descimon & K. Johnson,

1993 3

41. Archaeoprepona phaedra ssp. 2

42. Prepona laertes octavia Fruhstorfer,

1905

43. Prepona brooksiana ibarra

Beutelspacher, 1982 3

370 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Subfamily Morphinae Newman, 1834

Tribe Morphini Newman, 1834

Subtribe Morphina Newman, 1834

44. Morpho polyphemus Westwood, [1850]

45. Morpho helenor guerrerensis Le Moult

& Réal, 1962 3

Tribe Brassolini Boisduval, 1836

Subtribe Brassolina Boisduval, 1836

46. Caligo telamonius memnon (C. Felder &

R. Felder, 1867)

47. Caligo uranus Herrich-Schäffer, 1850

48. Opsiphanes boisduvallii Doubleday,

[1849]

49. Opsiphanes cassina fabricii (Boisduval,

1870)

50. Opsiphanes quiteria quirinus Godman

& Salvin, 1881

51. Opsiphanes tamarindi tamarindi C.

Felder & R. Felder, 1861

Subfamily Satyrinae Boisduval, 1833

52. Manataria hercyna maculata (Hopffer,

1874)

53. Oxeoschistus hilara ssp. 2

54. Oxeoschistus tauropolis ssp. 3

55. Pedaliodes dejecta ssp. 2

56. Cissia similis (Butler, 1867)

57. Cissia terrestris (Butler, 1867)

58. Cissia sp.

59. Cissia themis (Butler, 1867)

60. Cyllopsis clinas (Godman & Salvin,

1889) 2

61. Cyllopsis diazi L. Miller, 1974 4

62. Cyllopsis hedemanni hedemanni R.

Felder, 1869

63. Cyllopsis jacquelineae L. Miller, 1974 1

64. Cyllopsis nayarit (R. L. Chermock,

1947) 4

65. Cyllopsis pyracmon pyracmon (Butler,

1867)

66. Cyllopsis suivalenoides L. Miller, 1974

67. Euptychia fetna Butler, 1870

68. Hermeuptychia hermes (Fabricius, 1775)

69. Megisto rubricata pseudocleophes L.

Miller, 1976 4

70. Paramacera xicaque rubrosuffusa L.

Miller, 1972 2

71. Pindis squamistriga R. Felder, 1869

72. Taygetis kerea Butler, 1869

73. Taygetis mermeria griseomarginata L.

Miller, 1978 3

74. Taygetis uncinata Weymer, 1907 4

75. Taygetis virgilia (Cramer, 1776)

76. Taygetis weymeri Draudt, 1912

77. Gyrocheilus patrobas patrobas

(Hewitson, 1862) 4

Subfamily Apaturinae Boisduval, 1840

78. Asterocampa idyja argus (Bates, 1864)

79. Doxocopa laure laure (Drury, 1773)

80. Doxocopa pavon theodora (Lucas,

1857)

Subfamily Biblidinae Boisduval, 1833

Tribe Cyrestini Guenée, 1865

81. Marpesia chiron marius (Cramer, 1779)

82. Marpesia petreus ssp. nov.

83. Marpesia zerynthia dentigera

(Fruhstorfer, 1907)

Tribe Biblidini Boisduval, 1833

Subtribe Biblidina Boisduval, 1833

84. Biblis hyperia aganisa Boisduval, 1836

85. Mestra dorcas amymone (Ménétriés,

1857)

Subtribe Ageroniina Doubleday, [1847]

86. Hamadryas amphinome mazai Jenkins,

1983 3

87. Hamadryas atlantis lelaps (Godman &

Salvin, 1883) 3

88. Hamadryas februa ferentina (Godart,

[1824])

89. Hamadryas glauconome glauconome

(Bates, 1864)

90. Hamadryas guatemalena marmarice

(Fruhstorfer, 1916) 4

Subtribe Epicaliina Guenée, 1865

91. Eunica alcmena alcmena (Doubleday,

[1847])

92. Eunica monima (Stoll, 1782)

93. Eunica tatila tatila (Herrich-Schäffer,

[1855])

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94. Catonephele cortesi R. G. Maza,

1982 3

95. Catonephele numilia immaculata

Jenkins, 1985 2

96. Myscelia cyananthe cyananthe C.

Felder & R. Felder, 1867 4

97. Myscelia cyaniris alvaradia R. G.

Maza & Díaz, 1982 3

98. Myscelia ethusa ethusa (Doyère, [1840])

Subtribe Epiphilina Jenkins, 1987

99. Nica flavilla bachiana (R. G. Maza &

J. Maza, 1985) 2

100. Temenis laothoe quilapayunia R. G.

Maza & Turrent, 1985 3

101. Bolboneura sylphis beatrix R. G.

Maza, 1985 3

102. Epiphile adrasta escalantei Descimon

& Mast, 1979 4

103. Pyrrhogyra edocla paradisea R. G.

Maza & J. Maza, 1985 3

104. Pyrrhogyra neaerea hypsenor Godman

& Salvin, 1884

Subtribe Callicorina Orfila, 1952

105. Diaethria anna mixteca J. Maza, 1977

106. Diaethria astala asteroide R. G. Maza

& R. F. Maza, 1985 2

107. Callicore texa loxicha R. G. Maza & J.

Maza, 1983 1

108. Cyclogramma pandama (Doubleday,

[1848])

Subtribe Eubagina Burmeister, 1878

109. Dynamine dyonis Geyer, 1837

110. Dynamine postverta mexicana

D’Almeida, 1952

111. Dynamine theseus (C. Felder & R.

Felder, 1861)

Subfamily Limenitidinae Behr, 1864

Tribe Limenitidini Behr, 1864

Subtribe Limenitidina Behr, 1864

112. Adelpha barnesia leucas Fruhstorfer,

1915

113. Adelpha basiloides (Bates, 1865)

114. Adelpha bredowii Geyer, 1837

115. Adelpha diocles ssp. 3

116. Adelpha donysa ssp. 2

117. Adelpha fessonia fessonia (Hewitson,

1847)

118. Adelpha iphicleola iphicleola (Bates,

1864)

119. Adelpha iphiclus iphiclus (Linnaeus,

1758)

120. Adelpha leuceria leuceria (Druce, 1874)

121. Adelpha leucerioides ssp. 3

122. Adelpha lycorias melanthe (Bates, 1864)

123. Adelpha naxia naxia (C. Felder & R.

Felder, 1867)

124. Adelpha paraena massilia (C. Felder &

R. Felder, 1867)

125. Adelpha phylaca phylaca (Bates, 1866)

126. Adelpha pithys (Bates, 1864)

127. Adelpha serpa celerio (Bates, 1864)

Subfamily Nymphalinae Rafinesque, 1815

Tribe Coeini Scudder, 1893

128. Historis acheronta acheronta (Fabricius,

1775)

129. Historis odius dious Lamas, 1995

130. Pycina zamba zelys Godman & Salvin,

1884

Tribe Nymphalini Rafinesque, 1815

131. Colobura dirce dirce (Linnaeus, 1758)

132. Smyrna blomfildia datis Fruhstorfer,

1908

133. Smyrna karwinskii Geyer, [1833]

134. Hypanartia dione disjuncta Willmott, J.

Hall & Lamas, 2001

135. Hypanartia godmanii (Bates, 1864)

136. Hypanartia lethe (Fabricius, 1793)

137. Hypanartia trimaculata autumna

Willmott, J. Hall & Lamas, 2001

138. Nymphalis antiopa antiopa (Linnaeus,

1758)

139. Vanessa atalanta rubria (Fruhstorfer,

1909)

140. Vanessa cardui (Linnaeus, 1758)

141. Vanessa virginiensis (Drury, 1773)

Tribe Victorinini Scudder, 1893

142. Siproeta epaphus epaphus (Latreille,

[1813])

372 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

143. Siproeta stelenes biplagiata (Fruhstorfer,

1907)

144. Anartia fatima fatima (Fabricius, 1793)

145. Anartia jatrophae luteipicta Fruhstorfer,

1907

Tribe Junoniini Reuter, 1896

146. Junonia coenia Hübner, [1822]

147. Junonia evarete nigrosuffusa Barnes &

McDunnough, 1916

148. Junonia genoveva ssp. nov.

Tribe Melitaeini Herrich-Schäffer, 1843

Subtribe Nova

149. Chlosyne cynisca (Godman & Salvin,

1882) 4

150. Chlosyne erodyle ssp.

151. Chlosyne gaudialis wellingi L. Miller

& Rotger, 1979 1

152. Chlosyne hippodrome hippodrome

(Geyer, 1837)

153. Chlosyne janais janais (Drury, 1782)

154. Chlosyne lacinia lacinia (Geyer, 1837)

155. Chlosyne marina marina (Geyer, 1837)

156. Chlosyne melanarge (Bates, 1864)

157. Chlosyne theona theona (Ménétriés,

1855)

158. Microtia elva elva Bates, 1864

Subtribe Phyciodina Higgins, 1981

159. Phyciodes graphica graphica (R. Felder,

1869)

160. Phyciodes mylitta thebais Godman &

Salvin, 1878

161. Phyciodes pallescens (R. Felder,

1869) 4

162. Phyciodes phaon phaon (Edwards,

1864)

163. Phyciodes tharos tharos (Drury, 1773)

164. Tegosa guatemalena (Bates, 1864)

165. Anthanassa ardys ardys (Hewitson,

1864) 4

166. Anthanassa argentea (Godman &

Salvin, 1882)

167. Anthanassa atronia (Bates, 1866)

168. Anthanassa frisia tulcis (Bates, 1864)

169. Anthanassa nebulosa alexon (Godman

& Salvin, 1889) 4

170. Anthanassa otanes oaxaca

Beutelspacher, 1990 4

171. Anthanassa ptolyca amator (Hall,

1929) 4

172. Anthanassa sitalces cortes (Hall,

1917) 4

173. Anthanassa texana texana (Edwards,

1863)

174. Eresia phillyra phillyra Hewitson, 1852

Subfamilia Heliconiinae Swainson, 1822

Tribe Acraeini Boisduval, 1833

175. Altinote stratonice oaxaca (J. Miller &

L. Miller, 1979) 1

176. Actinote guatemalena guerrerensis J.

Maza, 1982 2

Tribe Heliconiini Swainson, 1822

Subtribe Heliconiina Swainson, 1822

177. Agraulis vanillae incarnata (Riley,

1926)

178. Dione juno huascuma (Reakirt, 1866)

179. Dione moneta poeyii Butler, 1873

180. Dryas iulia moderata (Riley, 1926)

181. Dryadula phaetusa (Linnaeus, 1758)

182. Eueides aliphera gracilis Stichel, 1903

183. Eueides isabella eva (Fabricius, 1793)

184. Heliconius charithonia vazquezae W. P.

Comstock & F. M. Brown, 1950

185. Heliconius erato cruentus Lamas,

1998 3

186. Heliconius hortense Guérin-Méneville,

[1844]

187. Heliconius ismenius telchinia

Doubleday, 1847

Tribe Argynnini Swainson, 1833

Subtribe Euptoietina Simonsen, 2006

188. Euptoieta claudia daunius (Herbst,

1798)

189. Euptoieta hegesia meridiania Stichel,

1938

Alpha diversity: Considering only the

data obtained during our fieldwork, 186 Nym-

phalidae species were collected. When species

from the literature review and historically col-

lected specimens were added, the list reached

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Fig. 1. Nymphalidae taxa endemic to Mexico that occur in the Loxicha Region, Oaxaca. Numbers correspond to names

available in our species list. Dorsal view: left side of the specimen; Ventral view: right side.

374 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

189 species. The Clench model estimated 188

species (Fig. 2) while Chao1 estimated 193

species. Therefore, the Clench model underes-

timated the region’s species richness, while the

Chao1 estimator was potentially more reliable.

According to Chao1, our list is 98 % complete.

Across the 11 states of Mexico’s Pacific

slope from Baja California to Chiapas (Table

1), the nymphalid species richness of the

Loxicha Region alone (189 spp.) surpasses

all states except for Guerrero (216 spp.) and

Chiapas (351 spp.). The high species richness

in Guerrero is attributable to the biogeographic

provinces Planicie Costera del Pacífico and

Sierra Madre del Sur occupying most of the

state, with the SMS having substantial faunal

endemism. In Chiapas, the high species rich-

ness can be explained by the convergence of

Atlantic and Pacific slopes in the state, as well

as the presence of tropical evergreen forest, a

vegetation type that supports more than 50 %

of Mexican species of Papilionoidea (Salinas-

Gutiérrez et al., 2004).

Species composition similarity across

regions: We compared nymphalid faunal rich-

ness across eight regions with differing ele-

vational gradients, comprising six from the

Pacific slope and two from the Atlantic slope

(Table 3). All regions have roughly similar

sampling effort, and species estimation meth-

ods indicate that more than 90 % of the species

have been recorded from each, thus validat-

ing the comparison. The Loxicha Region is

the most diverse for Nymphalidae on the

Pacific slope, and the second most diverse in

Oaxaca, behind only the Sierra de Juárez (259

spp.) of the Atlantic slope. Identical patterns

exist for Papilionidae and Pieridae (Luis-Mar-

tínez et al., 2020).

Similarity analysis shows two main groups

(Fig. 3), corresponding to the arrangement of

mountain ranges in Mexico and the conse-

quent isolation of the lepidopterofauna. The

first group comprises the Sierra Mazateca and

the Sierra de Juárez, on the Atlantic slope of

Oaxaca. The second group is composed of six

regions on the Pacific slope or in the Amacuzac-

Cuenca del Balsas, which are divided into two

subgroups with a similarity value of more than

50 %. The first subgroup contains the regions

of the Southern Sierra Madre del Sur: Atoyac

and Loxicha, which share 66 % of their species

(142 spp.). The second subgroup corresponds

to the regions located North of the Costa del

Pacífico, in the Western Eje Neovolcánico (San

Juan-Manantlán), the Northern Sierra Madre

del Sur (SMS-Michoacán), and Morelos which

includes parts of two biogeographic provinces:

Cuenca del Balsas and Eje Neovolcánico.

Fig. 2. Species accumulation curve of Nymphalidae in the Loxicha Region, Oaxaca, Mexico fitted to the Clench model. The

asymptote (dashed line) corresponds to 188 estimated species.

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Species composition distances between

Loxicha and other regions are correlated with

geographic distances, but also with biogeo-

graphic regionalization (Fig. 3). The two

regions with the greatest faunistic distance

are those from the Atlantic slope (the Sierra

de Juárez and Sierra Mazateca), even though

they are the closest geographically. The same

pattern was observed with Morelos, which

belongs to the Cuenca del Balsas: although

Morelos is closer to Loxicha than regions

like San Juan and Manantlán, these two have

lower faunistic distances. Considering only the

four regions in the SMS, faunistic distances

TABLE 3

Species richness of Nymphalidae across eight regions in Mexico

Region spp. Elevation (m) Slope

1. Sierra de Juárez, Oaxaca 259 100-3 100 Atlantic

2. Sierra Mazateca, Oaxaca 185 100-2 200 Atlantic

3. Loxicha Region, Oaxaca 189 80-2 600 Pacific

4. Sierra de Atoyac, Guerrero 170 300-3 100 Pacific

5. Morelos (state) 147 1 000-3 000 Pacific -CB

6. Michoacán, SMS portion 134 800-1 800 Pacific

7. Sierra de Manantlán, Jalisco-Colima 144 250-1 750 Pacific

8. Sierra de San Juan, Nayarit 135 0-1 350 Pacific

1: Luis-Martínez et al., 1991; 2: Álvarez-García et al., 2016; 3: this article; 4: Vargas-Fernández et al., 1994; 5: Luna-Reyes

et al., 2012 and Luna-Reyes, 2020; 6 and 8: MARIPOSA database; 7: Vargas-Fernández et al., 1999. CB: Cuenca del Balsas.

Fig. 3. Similarity of Nymphalidae species composition across eight regions in Mexico. The distance matrix was estimated

with the Jaccard index. For full names of the regions see Table 3.

376 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

are correlated with geographic distances, with

Atoyac being the closest and most similar to

Loxicha while also being the farthest and most

distinct relative to San Juan.

Species richness by site: The MARI-

POSA database contains records of Nymphali-

dae species from more than 5 000 sites across

Mexico. However, only 40 of those sites have

over 100 species recorded, eight of which are

on the Pacific slope while the other 32 are dis-

tributed elsewhere in Mexico (Table 4). From

the Loxicha Region, only Azulillo and Rancho

Hagia Sofía exceed 100 species, placing them

in sixth and fourth place at a national scale,

respectively. Areas like Acahuizotla, Guerrero

(158 spp.) and Candelaria Loxicha, Oaxaca

(138 spp.) also have high species richness but

are not specific sites, instead being extended

areas that include several sites with wide

variation in elevation and vegetation type.

These extended or historical “sites” served as

geographic references for decades prior to this

study, where collected specimens from other

sites were gathered and labeled. Additional

such historical “sites” across Mexico include

Xalapa, Catemaco, and Presidio in the state of

Veracruz (Luis-Martínez et al., 1996), Chilte-

pec, Oaxaca (Luis-Martínez et al., 1991), and

some others on the Atlantic slope.

Of the 40 sites with more than 100 spe-

cies of Nymphalidae (Table 4), 16 have been

sampled by members of the MZFC, and 30

are located in 11 of the regions with high-

est diversity of Papilionoidea in Mexico. Of

these high-diversity regions, nine were rec-

ognized by Luis-Martínez et al. (2003): 1,

Orizaba-Córdoba-Fortín de las Flores corridor;

TABLE 4

Sites with more than 100 species of Nymphalidae along the Atlantic and Pacific slopes of Mexico

State R Site spp. State R Site spp.

Atlantic slope

Veracruz 3 Xalapa* 193 San Luis Potosí Tamazunchale 109

Veracruz 1 Córdoba 180 Chiapas Santa Rosa 108

Veracruz 1 Presidio 170 Veracruz 3 Parque Francisco Javier Clavijero1107

Chiapas San Antonio Buena Vista 162 Oaxaca 4 Cerro Armadillo1106

Oaxaca 4 San José Chiltepec 156 Veracruz Cuetzalapan 104

Oaxaca 4 Metates1152 Veracruz 2 Popoctépetl 104

Veracruz 3 Barranca de Cayoapa*1149 Oaxaca 4 Soyolapan El Bajo1103

Puebla 10 Tequezquitla 145 Oaxaca Matías Romero 102

Veracruz 1 Fortín de las Flores*1146 Oaxaca La Gringa, Sta. María Chimalapa1102

Veracruz 2 Catemaco 142 Veracruz 1 Orizaba 102

Veracruz 2 Dos Amates 138 Veracruz 2 Tapalapan 102

Veracruz 1 Presidio, Ixhuatlán del Café* 134

Veracruz 3 Teocelo1132 Pacific slope

Puebla 10 Barranca de Patla* 132 Guerrero 8 Acahuizotla* 158

Oaxaca 4 La Esperanza1131 Oaxaca 11 Candelaria Loxicha* 138

Chiapas Santa Rosa, Comitán 130 Oaxaca 11 Rancho Hagia Sofía1121

Chiapas Zona Arqueológica Yaxchilán1129 Chiapas San Jerónimo, Tacaná 117

Oaxaca 4 Puerto Eligio1110 Jalisco 6 La Calera1115

Veracruz 2 Laguna de Catemaco 117 Oaxaca 11 Azulillo1109

Tabasco Cerro del Coconá1115 Guerrero 5 Río Santiago1106

Oaxaca 4 Naranjal Chiltepec 115 Guerrero 5 El Faisanal1105

*: extended or historical “sites” that combine records from multiple distinct sites under a single name (see text); 1: sites

collected by MZFC members; R: sites that lie in the most diverse regions of Mexico (see text).

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2, Tuxtlas; 3, Xalapa-Coatepec-Teocelo cor-

ridor; 4, Sierra de Juárez; 5, Sierra de Atoyac;

6, Sierra de Manantlán; 7, Mismaloya-Bahía de

Banderas corridor; 8, Acahuizotla; and 9, Sierra

de San Juan. Recently, two more high-diversity

regions were recognized: 10, La Sierra Norte

de Puebla (MARIPOSA database), and 11,

Loxicha (Arellano-Covarrubias et al., 2018;

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

2021; data presented here).

Species composition similarity across

sites: Of the 77 sites included in the spe-

cies composition similarity analysis (Table

5), 62 are on the Pacific slope and 15 on

the Atlantic slope, 44 have been sampled by

members of the MZFC, 32 are located between

0-750 m, 27 between 800-1 750 m, and six

above 1 800 m elevation. Five vegetation types

are included: TDF, TSDF, OPF, OF, and CF.

The latter includes three elevation-delimited

subtypes: low-CF (750-1 199 m), medium-

CF (1 200-1 800 m), and high-CF (> 2 000

m). Additionally, we recognize six ecotones:

OF-CF, OPF-CF, TDF-TSDF, TDF-TSDF-CF,

TSDF-CF and TSDF-OPF. Twenty-four sites

are in the CF, and seven more exist in an eco-

tone that includes CF. Therefore, CF is the most

well-represented vegetation type across the

sites (48 % of the sites).

The similarity analysis showed two groups

corresponding to the sites of the Atlantic and

TABLE 5

Sites included in the species composition similarity analysis for Mexican Nymphalidae

State Site Spp. Abbreviated name Elevation Vegetation type

Atlantic slope

San Luis Potosí Tamazunchale 109 SLP_Tam_350 350 TEGF

Puebla Tequezquitla 145 PUE_Teq_650 650 TSDF-CF

Puebla Barranca de Patla 132 PUE_Bpa_600 600 TSDF-CF

Veracruz Barranca de Cayoapa1149 VER_Bca_600 600 CF

Veracruz Catemaco 142 VER_Cat_250 250 TEGF

Veracruz Córdoba 180 VER_Cor_900 900 CF

Veracruz Fortín de las Flores1146 VER_FFl_900 900 CF

Veracruz Xalapa 193 VER_Jal_1350 1 350 CF

Veracruz Teocelo1132 VER_Teo_1200 1 200 CF

Oaxaca San José Chiltepec 156 OAX_Chi_100 100 TEGF

Oaxaca La Esperanza1131 OAX_Esp_1750 1 750 CF

Oaxaca Metates1152 OAX_Met_900 900 CF

Oaxaca Puerto Eligio1110 OAX_Pel_650 650 TEGF

Chiapas San Antonio Buena Vista 162 CHIS_SAB_1350 1 350 CF

Chiapas Santa Rosa, Comitán 130 CHI_SRC_1800 1 800 CF

Pacific slope

Nayarit Compostela141 NAY_Com_800 800 TDF

Nayarit Jumatán178 NAY_Jum_300 300 TDF

Nayarit La Bajada161 N_Baj_250 250 TSDF

Nayarit La Yerba, Tepeltite172 NAY_Yer_900 900 CF

Nayarit Mecatán147 NAY_Mec_300 300 TSDF

Nayarit Palapita166 NAY_Pal_650 650 TSDF

Nayarit Pintadeño123 NAY_Pin_750 750 OF

Nayarit San Blas145 NAY_SBl_50 50 MAN

Nayarit Singayta159 NAY_Sin_50 50 OP

Nayarit Venustiano Carranza169 NAY_Vca_1250 1 250 CF

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State Site Spp. Abbreviated name Elevation Vegetation type

Jalisco Ahuacapán185 JAL_Ahu_900 900 TDF

Jalisco Estación de Biología, UNAM, Chamela 50 JAL_Cha_100 100 TDF

Jalisco La Calera1115 JAL_Cal_650 650 TDF

Jalisco Los Mazos176 JAL_Maz_1600 1 600 CF

Jalisco Puerto Vallarta 49 JAL_Pva_50 50 TDF

Jalisco Zenzontla177 JAL_Zen_800 800 TDF

Colima Agua Dulce186 COL_Agu_250 250 TSDF

Colima Platanarillo188 COL_Pla_350 350 TSDF

Michoacán Arteaga 70 MICH_Art_900 900 TDF

Michoacán Chiquihuitillo 75 MICH_Chi_260 260 TDF

Michoacán Los Chorros del Varal172 MICH_Cho_900 900 TSDF

Michoacán P.H. Cupatitzio 83 MICH_CUP_1000 1 000 TSDF-PF

Michoacán Rancho “El Zorrillo” 92 MICH_Rzo_750 750 TDF

Michoacán Rancho “El Zorrillo”, Cañada Húmeda 70 MICH_RzC_750 750 TDF-TSDF

Morelos 3 km al NE de Chiautla 41 MOR_Shia_1000 1 000 TDF

Morelos Km 1.5 Carretera Tetecala-Coatlalco 56 MOR_TC_1100 1 100 TDF

Morelos Cañón de Lobos 51 MOR_CL_1200 1 200 TDF

Morelos 1.5 Km al E de Palo Grande 57 MOR_PG_1200 1 200 TDF

Morelos Sierra de Huautla 58 MOR_SH_1000 1 000 TDF-TSDF

Guerrero Agua Salada MOR_AS_760 760 TDF

Guerrero Coapango 74 GRO_Coa_1330 1 330 TDF

Guerrero Las Vías 72 GRO_LV_1200 1 200 TDF

Guerrero Cascada de las Granadas 81 GRO_CdeG_1370 1 370 TDF

Guerrero Los Amates 65 GRO_Ama_980 980 TDF

Guerrero Quetzalapa 66 GRO_Que_850 850 TDF

Guerrero Piedras Negras 54 GRO_PN_1300 1 300 TDF

Guerrero Palmillas 57 MOR_Pal_1050 1 050 TDF

Guerrero Acahuizotla 158 GRO_Aca_900 900 TDF-TSDF-CF

Guerrero El Faisanal194 GRO_Fai_1250 1 250 TSDF-CF

Guerrero El Iris124 GRO_Iris_2000 2 000 OF-CF

Guerrero La Golondrina145 GRO_LGo_1800 1 800 CF

Guerrero Las Parotas178 GRO_Par_350 350 TSDF

Guerrero Los Retrocesos 71 GRO_Ret_1600 1 600 CF

Guerrero Nueva Delhi182 GRO_Nde_1350 1 350 CF

Guerrero Puente de Los Lugardo194 GRO_PLL_800 800 TSDF

Guerrero Puerto del Gallo138 GRO_Pga_2350 2 350 BPE-CF

Guerrero Río Santiago, 4 km W1106 GRO_Rsa_680 680 TSDF

Oaxaca Azulillo1109 OAX_Azu_380 380 TSDF

Oaxaca Copalita, Río Copalita 58 OAX_Cop_600 600 CF

Oaxaca Copalita, Siete Veneros 54 OAX_Ven_940 940 CF

Oaxaca Finca Aurora-Finca San Isidro163 OAX_FAS_1100 1 100 CF

Oaxaca La Pasionaria172 OAX_Pas_1500 1 500 CF

Oaxaca La Soledad-Buenavista170 OAX_Sbu_1470 1 470 CF

Oaxaca Magdalena, El Lirio195 OAX_Mag_750 750 CF

Oaxaca Parque Nacional Huatulco168 OAX_PNH_100 100 TDF

Oaxaca Parque Nacional Huatulco, Río Cacaluta174 OAX_PNHC_80 80 TDF

Oaxaca Pluma Hidalgo, 4 km NW “La Curva” 1 95 OAX_Phi_1100 1 100 CF

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Pacific slopes, respectively (Fig. 4). Sites of

the Atlantic slope, in the low part of the phe-

nogram (1), are associated with the vegetation

and environmental conditions of the Gulf of

Mexico (East of Chiapas). This group has 163

exclusive taxa, six of which are distributed in

85 % of the sites of the group: Archaeoprepona

demophon centralis (Fruhstorfer, 1905), Cato-

nephele mexicana Jenkins & R. G. Maza, 1985,

Fountainea eurypyle confusa (A. Hall, 1929),

State Site Spp. Abbreviated name Elevation Vegetation type

Oaxaca Portillo del Rayo-Finca El Encanto188 OAX_PRa_1200 1 200 CF

Oaxaca Puente Arroyo “El Guajolote”174 OAX_Gua_2020 2 020 OPF

Oaxaca Rancho Hagia Sofía1121 OAX_Rha_410 410 TSDF

Oaxaca San José del Pacífico, 1 km S161 OAX_Pac_2280 2 280 OPF-CF

Chiapas San Jerónimo, Tacaná 117 CHIS_Sje_750 750 CF

1: sites collected by MZFC members; CF, cloud forest; MAN, mangrove; OF, oak forest; OP, Orbygnia palmar; OPF, oak-

pine forest; TDF, tropical deciduous forest; TEGF, tropical evergreen forest; TSDF, tropical sub-deciduous forest.

Fig. 4. Nymphalidae species composition similarity across 77 sites in Mexico (15 in the Atlantic and 62 in the Pacific

slopes). The distance matrix was estimated with the Jaccard index. Groups: 1. Atlantic slope; 2. Pacific slope; 3. Sierra de

Atoyac high-lands; 4. Costa del Pacífico, Northern Sierra Madre del Sur, and Western Eje Neovolcánico; 5. Sierra Madre

del Sur (Guerrero-Oaxaca); 6. Cuenca del Balsas (Guerrero-Morelos).

380 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Hamadryas amphinome mexicana (Lucas,

1853), Morpho helenor montezuma Guenée,

1859, and Taygetis thamyra (Cramer, 1779).

The second group comprises the sites in the

Pacific slope (2), divided into four subgroups.

In the first subgroup are the sites of the high-

lands of the Sierra de Atoyac de Álvarez (La

Golondrina, El Iris, and Puerto del Gallo) (3).

No exclusive species were recorded for these

three sites. Interestingly, although these sites

are geographically close to the sites of Balsas

and SMS, they segregate from other sites of the

Pacific because of their high-elevation species

assemblage (1 800-2 350 m). In the second

subgroup are the sites of the biogeographic

provinces Costa del Pacífico, Northern Sierra

Madre del Sur, and Western Eje Neovolcánico

(4). Six species are exclusive to this subgroup:

Bolboneura sylphis sylphis (Guérin-Méneville,

1844), Chlosyne rosita montana A. Hall, 1924,

Cyllopsis pallens L. Miller, 1974, Fountainea

halice tehuana (A. Hall, 1917), Polygonia

interrogationis (Fabricius, 1798), and Texola

elada hepburni (Godman, 1901), but all are

restricted to just 4-12 % of the sites in the

subgroup. A third subgroup is formed by the

sites in the Balsas province of Guerrero and

Morelos (6). These sites support mostly TDF

and lie at elevations from 760-2 280 m. Of

the three exclusive species to Balsas, Zisch-

kaia lupita (Reakirt, [1867]) is in 79 % of the

sites of this subgroup, while Chlosyne cyneas

cynisca (Godman & Salvin, 1882) and Cyl-

lopsis pertepida pertepida (Dyar, 1912) are

documented only in 7 % of the sites. The fourth

subgroup includes the Loxicha Region and is

composed of the sites of Oaxaca and Guerrero

in the SMS (5). Exclusive species recorded in

at least 80 % of the sites of this subgroup are

Archaeoprepona amphimachus baroni, Cato-

nephele numilia immaculata, Diaethria astala

asteroide, and D. anna mixteca. This subgroup

is divided into two sets of sites: those with low-

elevation CF, and those with TSDF between

350-900 m. However, the high-elevation site

El Guajolote (2 202 m) is similar to these sites,

possibly because of its geographical proximity.

The other set of sites are those within the SMS

with mid-elevation CF, between 600-1 600 m.

A clear distinction exists between the sites of

Oaxaca and those of Guerrero. These results

show that the position of the sites in the pheno-

gram corresponds to their geographic location

but also to their respective elevations (Fig. 4).

Diversity of the Loxicha Region relative

to other regions of the Sierra Madre del Sur

and Costa del Pacífico: The Mexican Pacific

slope, particularly in Guerrero and Oaxaca,

stands out for its remarkable diversity and

endemism at both the species and genus level.

In this area, only two faunistic studies exist that

cover a complete elevational gradient, and that

have an inventory completeness level higher

than 95 % according to the species richness

estimators. The first study was carried out in

the Sierra de Atoyac, Guerrero, from 300-3

100 m (Vargas-Fernández et al., 1994) and the

second one is the present study from 80-2 850

m (Table 6). Including both inventories, the

diversity of Nymphalidae in the area is 214

species including 11 subfamilies and 84 genera.

Of these 214 species, 88 % occur in the Loxi-

cha Region and 78 % in the Sierra de Atoyac.

Species richness is higher in Loxicha for nine

of the 11 subfamilies, with the greatest differ-

ence being in Nymphalinae (11 sspp.); only

Satyrinae has a higher species richness in the

Sierra de Atoyac, with nine subspecies.

Considering both transects, 13 species

endemic to Mexico were recorded: Pteronymia

rufocincta, Cyllopsis caballeroi Beutelspacher,

1982, C. clinas, C. diazi, C. jacquelineae,

C. nayarit, C. perplexa L. Miller, 1974,

Paramacera copiosa L. Miller, 1972, Tay-

getis uncinata, Catonephele cortesi, Chlosyne

cynisca, Chlosyne eumeda (Godman & Salvin,

1894), and Phyciodes pallescens. At the sub-

species level there are 66 endemics, consti-

tuting 31 % of the Nymphalidae recorded in

these areas. At a national scale, Loxicha and

the Sierra de Atoyac contain 43 % of the spe-

cies or subspecies of Nymphalidae endemic

to Mexico. Of these endemics, 56 are present

in the Loxicha Region and 55 in the Sierra

de Atoyac, with 64 % of the endemics shared

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between the two (Llorente-Bousquets, Trujano-

Ortega et al., 2006; Luis-Martínez et al., 2003;

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

al., 2021). As with the endemics of the Loxicha

Region, endemics of the Sierra de Atoyac were

grouped into four categories: 1, endemic to the

Sierra de Atoyac (3 sspp: Drucina championi

ssp., Eunica malvina almae Vargas, Llorente &

Luis, 1996, and Eueides isabella nigricornis R.

G. Maza, 1982); 2, endemic to the Sierra Madre

del Sur (17 sspp.); 3, endemic to the Pacific

slope (23 sspp.); and 4, endemic to Mexico (12

sspp.). These numbers are very similar to those

of the Loxicha Region, with differences of at

most three species. In both regions, Nymphali-

nae has the highest species richness and Biblid-

inae has most of the endemic taxa. Libytheinae,

Danainae, and Apaturinae have the lowest rich-

ness, a common pattern in America (Lamas,

2004; Pelham, 2008). Additionally, in Loxicha

these three subfamilies have no endemic taxa.

Phenology, elevational patterns, and

exclusivity of species in the transect

Phenology: Temporal distribution of the

species richness and abundance of Nymphali-

dae is influenced by season (Fig. 5). Richness

and abundance both reach their maximum

values in October (the end of the rainy season),

after which both parameters suddenly drop to

their lowpoints in December. The increase in

species richness is more or less steady, with

just a few more species in the rainy season

compared with the dry season; in contrast,

abundance has a steep increase from the dry to

the rainy season. April shows a slight increase

in species richness but a decrease in abundance,

suggesting an increase of rare species during

this month, compared with other months.

Elevational diversity patterns: Species

richness and diversity (eH) of Nymphalidae

decrease abruptly at elevations above 1 800

m, with just 33 % of the total species assem-

blage compared with 79 % at low elevations

and 82 % at mid elevations (Fig. 6). Although

observed species richness is higher at the

750-1 800 m level (151 spp.) than at the 0-750

m level (146 spp.), diversity estimates show no

significant difference between these two levels.

However, diversity is higher at low elevations,

which may be because of a higher dominance

of Hermeuptychia hermes and Cissia similis at

mid elevations which reduces overall diversity.

Elevational abundance patterns: Abun-

dance is highest at low elevations with 58 %

TABLE 6

Species richness and endemism of Nymphalidae subfamilies for two regions of the Pacific slope

of the Sierra Madre del Sur, Mexico (modified from Luis-Martínez et al., 2021)

Subfamily Loxicha Region, Oaxaca Sierra de Atoyac, Guerrero

Genera Species Subspecies Genera Species Subspecies

Libytheinae 1 1/0 1/0 1 1/0 1/0

Danainae 3 6/0 6/0 3 5/0 5/0

Ithomiinae 9 2/1 15/6 7 2/1 12/8

Charaxinae 10 4/1 21/8 9 3/0 16/6

Morphinae 3 3/0 8/1 3 2/0 7/1

Satyrinae 12 16/5 26/12 12 20/8 35/16

Apaturinae 2 3/0 3/0 1 1/0 1/0

Biblidinae 16 5/1 31/15 14 6/1 29/14

Limenitidinae 1 3/0 16/3 1 2/0 13/2

Nymphalinae 16 12/2 47/8 15 12/1 36/5

Heliconiinae 9 2/0 15/3 7 19/0 12/3

TOTAL

Richness/Endemism 82 57/10 189/56 73 73/11 167/55

382 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

of the total (15 861 records), decreases at mid

elevations with 39 % (10 520 records), and is

just 3 % (796 records) at high elevations. Each

level has dominant or characteristic species,

but the overall proportions of those dominant

species are highest in the mid and high eleva-

tions. The most abundant species in each level

belong to the Satyrinae subfamily. At low ele-

vations, Cissia similis (607 records) and Her-

meuptychia hermes (869 records) account for

5.1 and 5.4 % of the specimens, respectively;

at mid elevations, H. hermes is also the most

abundant species with 1 273 records (12 %);

at high elevations, 22 % of the specimens (177

records) are Paramacera xicaque rubrosuffusa

(Satyrinae) and 15 % (122 records) are Antha-

nassa a. ardys (Nymphalinae).

A total of 35 species were collected

from all three elevational levels (Appendix

1), but these species showed the following

three distinct distribution patterns. Pattern A:

Dione moneta poeyii, Paramacera xicaque

Fig. 6. Observed (black) and estimated (white) Nymphalidae species richness and diversity (exponential of Shannon-Wiener

index) across three elevational levels in the Loxicha Region, Oaxaca, Mexico.

Fig. 5. Phenology of Nymphalidae abundance and species richness in the Loxicha Region, Oaxaca, Mexico based on seven

years of sampling data. February was not sampled during this study.

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rubrosuffusa, and Vanessa virginiensis mainly

occur at high elevations with just one to three

records from low elevations. Pattern B: 19

spp. (v. gr. Anartia f. fatima, Cissia similis,

Smyrna blomfildia datis) with the opposite

pattern, being abundant at low elevations and

decreasing at mid elevations, with just a few

specimens at high elevations. Pattern C: 13

species (v. gr. Anthanassa a. ardys, Chlosyne h.

hippodrome, Hermeuptychia hermes) that are

abundant at mid elevations with fewer records

at both low and high elevations. Most of the

taxa (104, 56 %) are present in two elevational

levels, mainly in the low and mid elevations

with just 10 species in both the mid and high

elevations (v. gr. Anthanassa atronia, A. otanes

oaxaca) (see Appendix 1). Remarkably, eight

species Adelpha f. fessonia, Anartia jatrophae

luteipicta, Chlosyne m. marina, Danaus eresi-

mus montezuma, Doxocopa l. laure, Phyciodes

g. graphica, P. mylitta thebais, and Pteronymia

artena praedicta were collected only in the low

and high elevations, implying that they are also

present in the mid elevations. Finally, 45 spe-

cies are restricted to a single elevational level

(Table 7). Of these, 17 species are exclusive

to 0-750 m elevation, with the most abundant

being Bolboneura sylphis beatrix, Hamadryas

g. glauconome, and Hypna clytemnestra mexi-

cana; 20 species are exclusive to mid eleva-

tions where Anthanassa argentea and Cyllopsis

diazi comprised most of the records; and eight

species are distributed only in the highest

elevations where Adelpha bredowii, Nymphalis

a. antiopa, and Phyciodes t. tharos were the

most abundant.

Diversity patterns by vegetation type:

The vegetation type with the most observed

Nymphalidae species is the CF (147 species);

TABLE 7

Exclusive species of each elevational level (m)

Taxa 0-750 750-1 800 1 800-2 850

Danainae

Lycorea ilione albescens 9

Danaus p. plexippus 1

Ithomiinae

Episcada salvinia portilla 22

Pteronymia c. cotytto 9

Charaxinae

Hypna clytemnestra mexicana 159

Memphis p. perenna 33

Memphis wellingi 1

Archaeoprepona phaedra ssp. 2

Prepona brooksiana ibarra 2

Morphinae

Caligo uranus 1

Opsiphanes quiteria quirinus 1

Opsiphanes t. tamarindi 6

Satyrinae

Oxeoschistus hilara ssp. 5

Oxeoschistus tauropolis ssp. 2

Cyllopsis clinas 1

Cyllopsis diazi 124

Cyllopsis h. hedemanni 87

Cyllopsis nayarit 5

Megisto rubricata pseudocleophes 2

384 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

however, according to estimations the species

richness in the TSDF, the CF, and the OPCF

are not significantly different (Fig. 7). Interest-

ingly, the OPCF has just 9 % of the abundance

of both the TSDF and the CF. Below these three

types, the TDF supports 47 % of the species in

the region with 90 estimated taxa. The species

richness in the OPF is much lower than the

other vegetation types, with no rare species,

so the total richness estimate is also low (six

spp.). Estimations indicate that all vegetation

types were adequately sampled. The differ-

ence between estimated and observed species

is higher in the OPCF, suggesting either lower

sampling effort or greater numbers of rare spe-

cies in this vegetation type.

Abundance patterns by vegetation type:

The TSDF and the CF present the highest and

most similar abundance of Nymphalidae, with

11 703 (43 %) and 10 903 (40 %) records,

respectively. Abundance in other vegetation

types is distinctly different, with just 3 496

records (13 %) in the TDF and 1 028 (4 %) in

the OPCF. This family is even scarcer in the

OPF, with less than 1 % of the overall abun-

dance (47 records) in the Loxicha Region.

Dominant species in most vegetation types

mainly belong to the Satyrinae. Dominance

is low in most vegetation types, except in the

OPF where Nymphalis a. antiopa (24 speci-

mens) and Paramacera xicaque rubrosuffusa

(17 specimens) comprise 87 % of the records.

Paramacera xicaque rubrosuffusa is also the

most abundant species in the OPCF with

15 % of the recorded specimens, followed

closely by Anthanassa a. ardys with 14 % of

the abundance in this vegetation. The most

abundant species in the CF and the TSDF is

also a member of the Satyrinae: Hermeupty-

chia hermes, with 12 and 7 % of the records,

respectively. The TDF is the only vegetation

type where Satyrinae are not dominant, but is

also the vegetation where dominance is lowest,

with Microtia e. elva and Hamadryas g. glau-

conome comprising only 9 % and 8 % of the

records, respectively.

Most species (80 %) were collected in

multiple vegetation types, although some have

just one or two records in the vegetation at the

highest or lowest elevations. The only species

found in all five vegetation types was Smyrna

blomfildia datis, with one record in the OPF.

Thirty-seven species are exclusive to a veg-

etation type (Table 8), half of which are rare

species with one or two records. Fig. 8, Fig. 9,

Fig. 10, Fig. 11, and Fig. 12 show the eleva-

tional distribution by vegetation type of these

Fig. 7. Observed (black) and estimated (white) species richness by vegetation types of Nymphalidae in the Loxicha Region,

Oaxaca, Mexico. TDF = tropical deciduous forest, TSDF = tropical sub-deciduous forest, CF = cloud forest (low- and mid-

elevation), OPCF = oak-pine and high-elevation cloud forest, OPF = oak-pine forest.

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TABLE 8

Nymphalidae species exclusive to each vegetation type in the Loxicha Region, Oaxaca, Mexico

Taxa TDF TSDF CF OPCF OPF

Danainae

Lycorea ilione albescens 9

Danaus p. plexippus 1

Ithomiinae

Dircenna k. klugii 134

Episcada salvinia portilla 22

Pteronymia c. cotytto 9

Charaxinae

Memphis perenna perenna 33

Memphis wellingi 1

Archaeoprepona phaedra ssp. 2

Prepona brooksiana ibarra 2

Morphinae

Caligo uranus 1

Opsiphanes quiteria quirinus 1

Opsiphanes t. tamarindi 6

Satyrinae

Oxeoschistus tauropolis ssp. 2

Cyllopsis clinas 1

Cyllopsis h. hedemanni 87

Megisto rubricata pseudocleophes 2

Taygetis virgilia 1

Gyrocheilus p. patrobas 4

Apaturinae

Asterocampa idyja argus 18

Biblidinae

Marpesia zerynthia dentigera 18

Myscelia c. cyananthe 11

Bolboneura sylphis beatrix 211

Pyrrhogyra edocla paradisea 1

Limenitidinae

Adelpha barnesia leucas 3

Adelpha bredowii 15

Adelpha diocles ssp. 1

Nymphalinae

Pycina zamba zelys 4

Smyrna karwinskii 2

Hypanartia trimaculata autumna 42

Nymphalis a. antiopa 24

Vanessa cardui 1

Junonia genoveva ssp. nov. 5

Chlosyne cyneas cynisca 1

Phyciodes pallescens 1

Phyciodes p. phaon 24

Phyciodes t. tharos 19

Anthanassa texana texana 8

Heliconiinae

Dryadula phaetusa 4

Heliconius ismenius telchinia 1

Vegetation types: TDF = tropical deciduous forest, TSDF = tropical sub-deciduous forest, CF = cloud forest (low- and mid-

elevation), OPCF = oak-pine and high-elevation cloud forest, OPF = oak-pine forest. For each species listed, the habitat

preference value is 90-100 %.

386 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Fig. 8. Elevational profiles and characteristic species of Limenitidinae, Danainae, and Ithomiinae for each vegetation type in

the Loxicha Region, Oaxaca, Mexico. TDF = tropical deciduous forest, TSDF = tropical sub-deciduous forest, CF = cloud

forest (low- and mid-elevation), OPCF = oak-pine and high-elevation cloud forest.

Fig. 9. Elevational profiles and characteristic species of Charaxinae and Morphinae for each vegetation type in the Loxicha

Region, Oaxaca, Mexico. TDF = tropical deciduous forest, TSDF = tropical sub-deciduous forest, CF = cloud forest (low-

and mid-elevation), OPCF = oak-pine and high-elevation cloud forest.

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Fig. 10. Elevational profiles and characteristic species of Satyrinae and Apaturinae for each vegetation type in the Loxicha

Region, Oaxaca, Mexico. TDF = tropical deciduous forest, TSDF = tropical sub-deciduous forest, CF = cloud forest (low-

and mid-elevation), OPCF = oak-pine and high-elevation cloud forest.

Fig. 11. Elevational profiles and characteristic species of Biblidinae for each vegetation type in the Loxicha Region, Oaxaca,

Mexico. TDF = tropical deciduous forest, TSDF = tropical sub-deciduous forest, CF = cloud forest (low- and mid-elevation),

OPCF = oak-pine and high-elevation cloud forest.

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37 species (Table 8); all species with 95 % or

more of their records in a single vegetation type

were also included.

Efficiency and efficacy of Van Someren-

Rydon traps: During our fieldwork, 94 species

of seven subfamilies of Nymphalidae were col-

lected with Van Someren-Rydon traps (Appen-

dix 2). These taxa represent 50 % of the species

recorded and 41 % of the specimens. Some spe-

cies were captured with both traps and aerial

nets, but abundance was higher in traps (11

543; 73 %). Only 18 species (173 specimens)

were collected exclusively in traps. Biblidinae

was the subfamily with most species recorded

in traps with 24 spp., followed by Satyrinae

and Charaxinae with 23 and 21 species, respec-

tively; these three subfamilies account for 74

% of the specimens collected in traps (Fig.

13). In terms of abundance, Satyrinae was the

subfamily with the most records (40 % of the

specimens), followed by Nymphalinae (21 %).

Comparing the relative abundance and species

richness solely of trapped specimens for each

subfamily, two patterns can be recognized: A.

Satyrinae and Nymphalinae had greater abun-

dance than richness; B. Charaxinae, Biblidinae,

and Morphinae presented a relatively higher

richness than specimens, but still lower abun-

dance than subfamilies in pattern A (Fig. 13).

The efficiency and efficacy of the traps

at each elevational level (0-750 m, 750-1 800

m, and 1 800-2 850 m) were evaluated based

on the species and specimens captured rela-

tive to the totals. The percentage of speci-

mens collected with traps decreased at higher

elevations (Appendix 2; Fig. 14), and this was

more evident for Apaturinae and Nymphali-

nae. However, traps reduce to almost 40 % of

their efficiency at the higher elevations. On

the other hand, efficacy measured as the % of

recorded species of each subfamily, remained

constant along the elevational gradient. Traps

captured 100 % of the species of Charaxinae

and Morphinae at all levels (Fig. 15). Efficacy

was lower at higher elevations for Apaturinae

Fig. 12. Elevational profiles and characteristic species of Nymphalinae and Heliconiinae for each vegetation type in the

Loxicha Region, Oaxaca, Mexico. TDF = tropical deciduous forest, TSDF = tropical sub-deciduous forest, CF = cloud forest

(low- and mid-elevation), OPCF = oak-pine and high-elevation cloud forest.

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and Biblidinae, and slightly lower for Satyri-

nae. Finally, Van Someren-Rydon traps are less

effective in capturing species of Nymphalinae,

since percentages are lower than 40 % in all

elevational levels, although this efficacy stays

constant along the gradient. Of note, 16 species

of Limenitidinae were recorded in the Loxicha

Region but only two were collected in traps,

with an efficiency of 19 specimens out of the

298 collected of those species: Adelpha basi-

loides (5/227) and Adelpha i. iphiclus (14/71).

Natural history remarks on selected

taxa of the Loxicha Region: During our

fieldwork, some nymphalid species were par-

ticularly abundant or showed special behaviors.

We summarize that natural history data for each

relevant species below.

Hypna clytemnestra mexicana. Found in

the understory of the TDF, mainly in humid

and dark microhabitats. Easily collected with

traps in two sites: Parque Nacional Huatul-

co, and Río Cacaluta within Parque Nacional

Fig. 13. Relative abundance (black) and species richness (grey) of seven subfamilies of Nymphalidae collected in Van

Someren-Rydon traps in the Loxicha Region, Oaxaca, Mexico.

Fig. 14. Percentage of specimens of each Nymphalidae subfamily captured in Van Someren-Rydon traps by elevational

level, relative to all specimens captured in that level, in the Loxicha Region, Oaxaca, Mexico.

390 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Huatulco. More abundant in the rainy season

(July–November).

Consul excellens genini. Recorded and

collected in OPF, mostly on humid cliffs where

elements of CF are present. Attracted to fer-

mented fruit. Recorded mainly in San José

del Pacífico.

Chlosyne gaudialis wellingi. Flight faster

than other species of this genus, and flies

unusually high in search of flowers three to

four meters above ground level, in sunny open

places. Unusually low abundance relative to

other species of Chlosyne, which tend to be

very abundant.

Pycina zamba zelys. In Mexico, this taxon

has been recorded at just a few sites and with

low abundance. Only four specimens were col-

lected, the first records for the Pacific slope. All

specimens were collected on a wet wall along

a road where a stream formed puddles. Many

other species were also collected in this micro-

habitat: Pycina zamba zelys specimens were

licking the wet wall five or six meters above

ground level, as were specimens of Hypanar-

tia dione disjuncta, H. godmanii, H. lethe, H.

trimaculata autumna, Historis a. acheronta,

Epiphile adrasta escalantei, Diaethria anna

mixteca, D. astala asteroide, Marpesia chiron

marius, Siproeta e. epaphus, and S. stelenes

biplagiata, among other common taxa that

exploit this special microhabitat.

Altinote stratonice oaxaca. This species

is associated with low elevation CF from

1 000-2 500 m, making large elevational migra-

tions at the end of the summer. Present in open

spaces, with a slow flight, visiting bushes with

white composite flowers.

Archaeoprepona demophoon mexicana.

An elusive species, mainly occurring in the

forest canopy but sometimes in the understory

associated with open spaces like roads or river-

banks. Along rivers it can be found feeding on

minerals dissolved in wet sand and shows simi-

lar behavior along roads with some humidity

on the ground. Feeds mostly on decomposing

fruit and exudates of certain trees. Rapid fliers

that quickly flee when disturbed, often into the

forest canopy. Males are territorial and chase

away other males.

Bolboneura sylphis beatrix. Mostly along-

side roads or streams. If near rivers, it is likely

to be found on moist sand feeding on dissolved

minerals; if along roads, likely feeding on small

flowers. Flight is slow and halting, generally

about 50 cm above ground level. Groups of

more than 10 specimens can be found along-

side streams. When resting usually perches on

leaves. Recorded only in TDF.

Fig. 15. Percentage of species of each Nymphalidae subfamily recorded in Van Someren-Rydon traps by elevational level,

relative to all species recorded in that level, in the Loxicha Region, Oaxaca, Mexico.

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Heliconius erato cruentus. Found mainly

in open spaces and along roads and small paths.

Flight slow and halting, always less than one

meter above ground level. Often feeds at road-

side flowers, but sometimes also attracted to

decomposing fruit. Gregarious, forming groups

to overnight in protected places.

Microtia elva elva. Frequently found along

open roads with some shade. Flight low and

halting, at most 50 cm above ground level.

Feeds mostly on small composite flowers along

roadsides. When disturbed, usually flies just a

few meters away.

DISCUSSION

More than 60 % of Mexico’s surface is

covered by mountains, but faunal surveys of

Lepidoptera across elevational gradients are

scarce, except for those done by members of

the MZFC (Arellano-Covarrubias et al., 2018;

Luis-Martínez & Llorente-Bousquets, 1990;

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

2020; Vargas-Fernández et al., 1994; Vargas-

Fernández et al., 1999). This type of analysis

provides important insight into various bio-

logical processes. Such insights include the

recognition of disjunct distributions across

mountain archipelagos due to speciation and

endemism processes, and the ability to detect

changes of species distribution due to climate

change, especially in stenotopic species vulner-

able to extinction.

Of all elevational gradients in Mexico,

the Loxicha Region is the most systematically

sampled area for Papilionoidea sensu lato,

particularly for the Nymphalidae (Table 2).

Many experienced collectors have sampled

this region for decades (Álvarez-García et al.,

2016; Arellano-Covarrubias et al., 2018; Luis-

Martínez et al., 2000; Luis-Martínez et al., 2003;

Luis-Martínez et al., 2020; Michán et al., 2004;

Pozo et al., 2008). Estimates of Nymphalidae

species richness in the Loxicha Region show

that it has the highest percentage of inventory

completeness for any region in Mexico (Var-

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

et al., 1999). Furthermore, it has the highest

species richness of any region on the Pacific

slope (Table 1 and Table 3) and the second

richest in the country, behind only the Sierra de

Juárez (Table 3).

Sampling in the Loxicha Region began in

1960, based on historical specimens that we

checked and on publications that described a

dozen taxa from those specimens. However,

even with Loxicha’s great species richness and

endemism, the present work is the first sys-

tematic study that provides a detailed regional

description of the diversity and distributional

patterns of Nymphalidae, mirroring similar

works published previously for Riodinidae

(Arellano-Covarrubias et al., 2018), Papilioni-

dae, and Pieridae (Luis-Martínez et al., 2020).

Species richness of Nymphalidae by site:

Comparison of the diversity on both the Pacific

and Atlantic slopes confirms that the latter has

higher species richness (v. gr. Andresen, 2008;

Ceballos, 1995; Flores-Contreras & Luna-

Reyes, 2017; González-Ramírez et al., 2017;

Luis-Martínez et al. 2003; Salinas-Gutiérrez

et al. 2004). Of sites with 100 or more species

of Nymphalidae, only 20 % are on the Pacific

slope. Moreover, among these Pacific slope

“sites” are Acahuizotla, Guerrero and Cande-

laria Loxicha, Oaxaca, which are larger areas

that historically encompassed a group of sites

with distinct elevations and vegetation types

but labeled under a single site name. These

areas were used as reference points for collec-

tors for three decades (1950-1980) of the 20th

century. Therefore, actual species richness of

these reference sites is lower than cited, further

reducing to 15 % the Pacific slope sites with

over 100 species. In the last 40 years, more

than 10 faunistic studies on Papilionoidea

were done on the Pacific slope by members of

the MZFC (Llorente-Bousquets et al., 1996;

Llorente-Bousquets et al., 2004; Luis-Martínez

& Llorente-Bousquets, 1993; Luis-Martínez,

1997; Luis-Martínez, 1999; Luis-Martínez,

2001; Vargas-Fernández et al., 1994; Vargas-

Fernández et al., 1996; Vargas-Fernández et

al., 1999; Warren & Llorente-Bousquets, 1999;

Warren et al., 1996; Warren et al.,1998), but only

392 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

six sites exceed 100 species. Of these, only four

are in Sierra Madre del Sur (two in the Sierra

de Atoyac and two in the Loxicha Region),

which is the mountain range with the greatest

diversity on the Pacific slope despite lack-

ing tropical evergreen forest vegetation. San

Jerónimo, Tacaná, Chiapas is the only Mexican

site with more than 100 species that lacks a

systematic faunistic study; data from this site

were obtained from national and international

specimen collections (MARIPOSA database).

Species composition similarity across

regions: The similarity analysis reflects how

Lepidoptera are associated with mountain

archipelagos, as described by Llorente-Bous-

quets (1984) and Halffter (1987). In Mexico,

the states of Guerrero and Oaxaca support the

most faunistically similar mountain habitat

“islands” (Llorente-Bousquets, 1984). These

two isolated regions each have exclusive or

endemic species, yet they share at least 50 %

of their species, forming geographic and genea-

logic relations between them. Other areas are

disjunct, each with its own biogeographic histo-

ry based on patterns of biotic provinces defined

by Morrone et al. (2002) and Morrone (2005).

The distinction between the diurnal Lepidop-

tera on the Atlantic and Pacific slopes had been

further revealed using panbiogeographic meth-

ods (Llorente-Bousquets, Trujano-Ortega et al.,

2006; Luis-Martínez et al., 2006; Oñate-Ocaña

et al., 2006; Vargas-Fernández et al., 2006).

The faunistic similarity of Nymphalidae

shows a division in the Pacific slope and SMS

in two parts, a Northwest section (Nayarit,

Jalisco, Michoacán) and a Southeast section

(Guerrero, Oaxaca), with the latter including

the Loxicha Region. This division of the SMS

in two subprovinces was proposed by Santiago-

Alvarado et al. (2016) based on data from 32

species endemic to the SMS. No distribution

was consistent with the SMS as a whole.

Later, Morrone (2017) named and described

three subprovinces in the SMS: the West sub-

province with the Jalisciense and Jalisciense-

Manantlán districts, the Central subprovince

with the Michoacán district, and the Oriental

subprovince comprising the Guerrerense and

Tierras Altas de Oaxaca districts. Although

Nymphalidae data do not reflect the separation

of these three subprovinces, when analyzing

the data at a site-level scale, Nymphalidae

of the Southwest SMS are shown to be more

similar to the Balsas province fauna than to the

fauna of the Northeast SMS.

Species composition similarity of sites:

Diurnal Lepidoptera associated with the CF at

all elevational levels are an excellent model for

the study of complex dispersion, vicariance,

and speciation patterns, due to the disjunct

distribution and isolation of this vegetation

community (Rzedowski, 1978). Generally, CF

fauna distribution patterns are consistent with

the geographic barriers that delimit each of

the mountain ranges of Mexico. An integrated

phylogeographic analysis that included birds,

mammals, and plants concluded that the evo-

lution of the biota of the CF in Mesoamerica

resulted from a complex combination of dis-

tinct histories of range expansion, isolation,

and biotic differentiation (Ornelas et al., 2013).

A similar process seems to have occurred

with the Papilionoidea sensu lato and particu-

larly with Nymphalidae, based on distribution

patterns resolved by multiple studies (Llor-

ente-Bousquets, 1984; Llorente-Bousquets &

Escalante, 1994; Llorente-Bousquets & Luis-

Martínez, 1998).

Nymphalidae assemblages are closely

related to vegetation type and elevation but

are also influenced by geographic history and

distance. Sites are grouped in six sets accord-

ing to their similarity (Fig. 4). The first one

corresponds to the Atlantic slope, while the

other subgroups represent the diverse faunas

of the Pacific slope distributed in distinct phys-

iographic units and biotic provinces. These

results concur with the data published in fau-

nistic studies and generic reviews, describing

the distribution of fauna in three elevational

levels of the CF based on the proposal of

Llorente-Bousquets (1984). Sites on the Pacific

slopes (Fig. 4, subgroups 3 to 6) are grouped

similarly to the elevational gradients (Fig. 3).

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The Sierra Madre del Sur (Guerrero and Oaxa-

ca portion) (Fig. 4, subgroup 5) is more similar

to the Cuenca del Balsas (Fig. 4, subgroup

6), than to the sites in the Costa del Pacífico,

Northern Sierra Madre del Sur, and Western

Eje Neovolcánico provinces (Fig. 4, subgroup

4). Notably, the sites above 1 800 m in the

Sierra de Atoyac de Álvarez (Fig. 4, subgroup

3, La Golondrina, El Iris, and Puerto del Gallo)

are separated from subgroups 4 and 6, and

from the sites above 1 800 m in the Loxicha

Region (Puente Arroyo “El Guajolote” and San

José del Pacífico, 1 km S), even though these

two regions are geographically close. Puente

Arroyo “El Guajolote” is part of the subgroup

of low elevation Guerrero-Oaxaca sites below

900 m, while San José del Pacífico is separated

from other sites of the Sierra Madre del Sur,

placed near the sites of the Cuenca del Balsas,

as the most distinct site of this group. These

similarity patterns of the high-elevation sites

of Atoyac and Loxicha suggest that these are

isolated communities, with a distinctive fauna,

different from nearby low-elevation sites as

well as from other high-elevation sites.

In general, the faunistic composition of

Nymphalidae agrees with the biogeographic

history of Mexico and its proposed provinces

(Morrone, 2005; Morrone et al., 2002). First,

there is a clear distinction between the faunas

of the Atlantic and the Pacific. At a finer scale,

a division of the Sierra Madre del Sur into a

Northern and Southern portion was recognized.

Within the Southern portion of SMS, sites are

grouped by elevation and vegetation type, with

high elevation sites separated from the rest.

Therefore, distribution patterns of Nymphali-

dae in Mexico are produced at large scales by

historical elements and at smaller scales by

ecological elements.

Phenology: In the Loxicha Region, the

temporal distribution of Nymphalidae species

richness and abundance is correlated direct-

ly with the summer-associated rainy season

(Fig. 5), which is common in most of the

regions in Mexico (Álvarez-García et al., 2016;

Luis-Martínez & Llorente-Bousquets, 1990;

Luis-Martínez & Llorente-Bousquets, 1993;

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

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

Loxicha Region, nymphalid species richness

and abundance are higher from July to October.

This pattern is echoed in the Sierra de Juárez

(Oaxaca), the Sierra de Atoyac (Guerrero),

the Sierra de Manantlán (Jalisco-Colima), and

the Sierra de San Juan (Nayarit), although the

month with the highest species richness differs

among the regions. In Loxicha, October had

the highest richness. In comparison, September

is the month with most species recorded in the

Sierras de Juárez, Manantlán, and San Juan,

while in the Sierra de Atoyac most species

were recorded in July. As rainfall decreases so

too does species richness, with April and May

being both the driest months and having among

the lowest species richness in all regions.

Diversity and abundance by vegetation

type: Stenoic species of Nymphalidae have a

wider distribution across different vegetation

types (Table 8) than the stenoic species of

Papilionidae, Pieridae, and Riodinidae, which

mostly occur in the TSDF and CF (Arellano-

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

2020). The elevational profiles (Fig. 8, Fig.

9, Fig. 10, Fig. 11, and Fig. 12) show that the

stenoic species of both Satyrinae and Nym-

phalinae are characteristic of each vegetation

type. In contrast, Charaxinae and Biblidinae

lack stenoic species in the OPCF but are repre-

sented in the lower-elevation vegetation types.

The TDF is the vegetation type with the fewest

stenoic species, with only 11 characteristic spe-

cies of four subfamilies: Charaxinae, 3 spp.;

Satyrinae, 1 sp.; Apaturinae, 1 sp.; Biblidinae,

5 spp.; and Nymphalinae, 1 sp. Comparing

with other families, Riodinidae has a single

characteristic species of the TDF (Arellano-

Covarrubias et al., 2018). In Papilionidae only

the TSDF has a significant number of stenoic

species with five of the 10 species recorded,

while the number of stenoic species of Pieridae

does not differ among vegetation types (Luis-

Martínez et al., 2020).

394 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

The observed species richness and abun-

dance of Nymphalidae show that the OPCF is

the most diverse vegetation type. Although the

estimated species richness for the OPCF is not

significantly different from that of the CF and

the TSDF, the OPCF abundance is lower, with

nine exclusive species. Despite being originally

considered a combination of two vegetation

types (Luis-Martínez et al., 2020), these nym-

phalid results support the recognition of the

OPCF as a distinct ecological unit. We suggest

the analysis of other taxonomic groups to fur-

ther characterize this vegetation type. In com-

parison, the CF presented the highest number

of exclusive species with 16 taxa, although the

strong dominance of some species decreases

its diversity to a value lower than that of the

OPCF. Considering the diversity and exclusive

species of Nymphalidae that they support, we

recommend that the OPCF, CF, and the TSDF

be considered conservation priorities.

Van Someren-Rydon traps: Fruit-feeding

butterflies constitute approximately 50 % of

all Nymphalidae species (Santos et al., 2011).

Similarly, other authors have reported 40-55

% of fruit-feeding Nymphalidae species from

tropical forests, with some variation in the

composition among the subfamilies (Daily &

Ehrlich, 1995; DeVries & Walla, 2001; DeVries

et al., 1999; Freire-Jr. et al., 2021a; Pinheiro

& Ortiz, 1992; Uehara-Prado et al., 2004;

Vargas et al., 1994; Vargas et al., 1999). In the

last two decades, this guild has been used in

analyses of forest fragmentation, conservation,

monitoring, community structure, and ecology,

because these species are more easily sampled

(Freire Jr., Oliveira, et al., 2021; Freitas et al.,

2003; González-Valdivia et al., 2016; Pozo,

2006; Pozo et al., 2005; Pozo et al., 2009). The

growing importance of this guild underscores

the relevance of the Van Someren-Rydon trap

efficiency and efficacy data presented here.

During our fieldwork, all species of

Charaxinae, Morphinae, and Apaturinae were

collected in traps, along with most species of

Satyrinae (92 %) and Biblidinae (77 %). In

contrast, only two species of Limenitidinae

were collected in traps (13 %). Apparently,

Limenitidinae are only occasional visitors to

traps, probably looking for water as some

Ithomiinae also do, since neither of these

two subfamilies belong to the fruit-feeding

guild (Meave del Castillo & Luis-Martínez,

2000). Traps are especially useful for collect-

ing Charaxinae because these species frequent

open spaces above the forest canopy, while

Satyrinae frequent the understory less than one

meter above ground (Freire Jr., Ribeiro, et al.,

2021). Across all Nymphalidae, 19 % of the

taxa were collected exclusively with this tech-

nique, and for 69 % of the species documented,

75 % of the specimens were trapped. We thus

emphasize that Van Someren-Rydon traps are

critical for accurately documenting the species

richness, abundance, and elevational and veg-

etational distributions of Nymphalidae.

Ethical statement: The authors declare

that they all agree with this publication and

made significant contributions; that there is no

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

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are

fully and clearly stated in the acknowledge-

ments section. A signed document has been

filed in the journal archives.

ACKNOWLEDGMENTS

JLB and ALM thank the projects DGAPA

PAPIIT IN220521, PAPIME PE202820,

CONACyT 284966, and CONABIO JF018 for

financial support. MTO thanks the Postdoctoral

Research Fellowship CONACyT 284966. AAC

thanks the Posgrado en Ciencias Biológicas de

la Universidad Nacional Autónoma de México.

We thank the Departmento de Biología Evo-

lutiva of the Facultad de Ciencias, Universi-

dad Nacional Autónoma de México (UNAM),

México; Armando Canavati, owner of Rancho

Hagia Sofía; and the authorities of the Parque

Nacional Huatulco for their support during this

project. Thanks to Sandra Nieves, Blanca Clau-

dia Hernández and other colleagues and stu-

dents who participated in the field work, and to

395

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Adriana Atzinameyali Sánchez Castañeda for

processing the collected material. The authors

thank Adam G. Clause for English language

review, and the peer reviewers for their com-

ments on the manuscript.

RESUMEN

Distribución, diversidad, endemismo y ecología de las

mariposas Nymphalidae (Lepidoptera: Nymphalidae)

en la Región Loxicha, Oaxaca, México

Introducción: La Región Loxicha de Oaxaca, México, ha

sido históricamente importante para el estudio de Nympha-

lidae, segunda en riqueza de especies en Papilionoidea.

Describir los patrones de diversidad de este taxón de mari-

posas en Loxicha puede mejorar nuestra comprensión de la

historia evolutiva de la Sierra Madre del Sur, la vertiente

del Pacífico mexicano y México en general. Objetivo:

Describir los patrones temporales y espaciales de la diver-

sidad de Nymphalidae a lo largo de un gradiente altitudinal

(80-2 600 m), y comparar la fauna de Loxicha con otras

regiones de México.

Métodos: Obtuvimos 28 756 registros de 21 sitios de la

Región Loxicha, que representan siete años de muestreo.

Estimamos y analizamos la diversidad, el endemismo y

los patrones de distribución para tres niveles altitudinales

y cinco tipos de vegetación. Estimamos la composición de

especies y la similitud con otras regiones de las vertientes

del Pacífico y Atlántico de México.

Resultados: Identificamos 189 taxones, incluyendo espe-

cies y subespecies, de 85 géneros y diez subfamilias de

Nymphalidae. Loxicha contiene 46 % de las especies de la

familia reconocidas para México, incluidas diez especies

endémicas y 56 subespecies endémicas. El bosque mesófilo

y las elevaciones bajas fueron los hábitats más diversos

para esta familia. Existe una clara divergencia entre las

faunas del Atlántico y del Pacífico, y la Sierra Madre del

Sur tiene dos componentes faunísticos. Los sitios de eleva-

ciones altas en Oaxaca, y en el estado vecino de Guerrero,

tienen una fauna distintiva, aparentemente aislada de los

sitios de elevaciones bajas, lo que revela una distribución

archipelágica para los Nymphalidae del bosque mesófilo.

Conclusiones: La Región Loxicha es una de las zonas más

diversas para Nymphalidae en México. La distribución en

la vertiente del Pacífico está determinada por la historia

geográfica y las condiciones ecológicas, incluida la eleva-

ción. Nymphalidae puede usarse para probar hipótesis de

regionalización biogeográfica en México.

Palabras clave: mariposas ninfálidas; gradiente de eleva-

ción; eficiencia de muestreo; fenología; provincias biogeo-

gráficas; trampas Van Someren-Rydon.

REFERENCES

Álvarez-García, H., Ibarra, A., & Escalante, P. (2016).

Riqueza y distribución altitudinal de las mariposas de

la Sierra Mazateca, Oaxaca (Lepidoptera: Papilionoi-

dea). Acta Zoológica Mexicana (Nueva Serie), 32(3),

323–347. https://doi.org/10.21829/azm.2016.323967

Andresen, E. (2008). Dung beetle assemblages in primary

forest and disturbed habitats in a tropical dry forest

landscape in western Mexico. Journal of Insect Con-

servation, 12(6), 639–650. https://doi.org/10.1007/

s10841-007-9100-y

Arellano-Covarrubias, A., Llorente-Bousquets, J., & Luis-

Martínez, A. (2018). Distribución y fenología de la

familia Riodinidae (Lepidoptera: Papilionoidea) en el

bosque tropical subcaducifolio de Oaxaca, México.

Revista de Biología Tropical, 66(2), 503–558. http://

dx.doi.org/10.15517/rbt.v66i2.33378

Bailowitz, R., Brock, J., & Danforth, D. (2017). Annotated

checklist of the butterflies (Lepidoptera) of Sono-

ra, Mexico. Dugesiana, 24(2), 125–147. https://doi.

org/10.32870/dugesiana.v24i2.6639

Balcázar, M. (1993). Butterflies of Pedernales, Michoacán,

Mexico, with notes on seasonality and faunistic affi-

nities (Lepidoptera: Papilionoidea and Hesperioidea).

Tropical Lepidoptera, 4(2), 93–105.

Beutelspacher, C. (1982). Lepidópteros de Chamela, Jalis-

co, México I. Rhopalocera. Anales del Instituto de

Biología (Zoología), UNAM, 52(1), 371–388.

Brown, J. W., Real, H. G., & Faulkner, D. K. (1992).

Butterflies of Baja California. Faunal survey, natu-

ral history, conservation biology. The Lepidoptera

Research Foundation, Inc.

Ceballos, G. (1995). Vertebrate diversity, ecology and con-

servation in neotropical dry forests. In S. H. Bullock,

H. A. Mooney, & E. Medina (Eds.), Seasonally dry

tropical forests (pp. 195–220). Cambridge University

Press.

Chao, A., & Shen, T. J. (2003). Nonparametric estima-

tion of Shannon’s index of diversity when there

are unseen species in sample. Environmental and

Ecological Statistics, 10(4), 429–443. https://doi.

org/10.1023/A:1026096204727

Chao, A., & Shen, T. J. (2010). Program SPADE (Species

Prediction and Diversity Estimation). User guide.

Available at http://chao.stat.nthu.edu.tw

Colwell, R. K. (2013). ESTIMATES: Statistical Estimation

of Species Richness and Shared Species from Sam-

ples (Version 9.1). Available at http://purl.oclc.org/

estimates.

Colwell, R. K., & Coddington, J. A. (1994). Estimating

terrestrial biodiversity through extrapolation. Philo-

sophical Transactions of the Royal Society of London.

396 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Series B: Biological Sciences, 345(1311), 101–118.

https://doi.org/10.1098/rstb.1994.0091

Comstock, J. A. (1953). Collecting butterflies in the coastal

area of Mexico near Manzanillo, Colima, with notes

on the life history of a rare skipper. Bulletin of the

Southern California Academy of Sciences, 52(2),

43–45.

Comstock, J. A. & Vázquez, L. (1961). Estudios de los

ciclos biológicos en lepidópteros mexicanos. Anales

del Instituto de Biología, UNAM, 31(1/2), 349–448.

Daily, G. C., & Ehrlich, P. R. (1995). Preservation of biodi-

versity in small rainforest patches: rapid evaluations

using butterfly trapping. Biodiversity Conservation,

4, 35–55. http://dx.doi.org/10.1007/BF00115313

DeVries, P. J., & Walla, T. R. (2001). Species diversity and

community structure in neotropical fruit-feeding but-

terflies. Biological Journal of the Linnean Society, 74,

1–15. http://dx.doi.org/10.1111/j.1095-8312.2001.

tb01372.x

DeVries, P. J., Walla, T. R., & Grenney, H. F. (1999). Spe-

cies diversity in spatial and temporal dimensions of

fruit-feeding butterflies from two Ecuadorian rainfo-

rests. Biological Journal of the Linnean Society, 68,

333–353. https://doi.org/10.1111/j.1095-8312.1999.

tb01175.x

Flores-Contreras, I., & Luna-Reyes, M. (2017). Diversidad

y distribución de cinco familias de Papilionoidea

(Lepidoptera) de las selvas altas en la provincia

biogeográfica del Golfo de México. Acta Zoológica

Mexicana, 33(2), 211–230. https://doi.org/10.21829/

azm.2017.3321062

Flores-Villela, O., & García-Vázquez, U. O. (2014). Biodi-

versidad de reptiles en México. Revista Mexicana de

Biodiversidad, Suplemento Biodiversidad de México,

85, S467–S475. https://doi.org/10.7550/rmb.43236

Freire-Jr., G., Oliveira, H., Silva, T., Pamela, H., Dias, J.,

Santos, J., Marini-Filho, O., Freitas, A., & Diniz, I.

(2021a). Good Things Come in Larger Packages: Size

Matters in Neotropical Fruit-Feeding Butterfly Dis-

persal [artículo]. MDPI in 1st International Electro-

nic Conference on Biological Diversity, Ecology and

Evolution session Biodiversity Conservation. https://

doi.org/10.20944/preprints202111.0113.v1

Freire-Jr., G., Ribeiro, D. B., de Carvalho Santos, A.,

Silva, T., Dias, J. P., Rodrigues, H. P., & Diniz, I.

R. (2021b). Horizontal and vertical variation in

the structure of fruit-feeding butterfly (Nymphali-

dae) assemblages in the Brazilian Cerrado. Insect

Conservation and Diversity, 14, 1–10. https://doi.

org/10.1111/icad.12547

Freitas, A. V. L., Francini, R. B., & Brown, K. S. (2003).

Insetos como indicadores ambientais. In L. Cullen,

R. Rudran, & C. Valladares-Padua (Eds.), Métodos

de estudos em biologia da conservação & manejo da

vida silvestre (pp.125–152). Editora UFPR.

García-Mendoza, A. J., & Meave del Castillo, J. A. (2011).

Diversidad florística de Oaxaca: de musgos a angios-

permas (colecciones y lista de especies). Universidad

Nacional Autónoma de México-Comisión Nacional

para el Conocimiento y Uso de la Biodiversidad.

González-Ramírez, M., Zaragoza-Caballero, S., & Pérez-

Hernández, C. X. (2017). Análisis de la diversidad

de Coleoptera en el bosque tropical caducifolio en

Acahuizotla, Guerrero, México. Revista Mexica-

na de Biodiversidad, 88(2), 381–388. https://doi.

org/10.1016/j.rmb.2017.03.008

González-Valdivia, N. A., Pozo, C., Ochoa-Gaona, S.,

Ferguson, B. G., Cambranis, E., Lara, O., Pérez-

Hernández, I., Ponce-Mendoza, A., & Kampichler, C.

(2016). Nymphalidae frugívoras (Lepidoptera: Papi-

lionoidea) asociadas a un ecomosaico agropecuario y

de bosque tropical lluvioso en un paisaje del sureste

de México. Revista Mexicana de Biodiversidad, 87,

451–464. https://doi.org/10.1016/j.rmb.2016.04.003

Halffter, G. (1987). Biogeography of the montane entomo-

fauna of Mexico and Central America. Annual Review

of Entomology, 32, 95–114. https://doi.org/10.1146/

annurev.en.32.010187.000523

Holland, R. W. (1995). Distribution of selected Anthocha-

ris, Euchloe and Pontia (Pieridae) in New Mexico,

Texas, Chihuahua and Sonora. Journal of the Lepi-

dopterists’ Society, 49(2), 119–135.

Jost, L. (2006). Entropy and diversity. Oikos, 113(2), 363–375.

https://doi.org/10.1111/j.2006.0030-1299.14714.x

Lamas, G. (2004). Checklist: Part 4A. Hesperioidea - Papi-

lionoidea. In J. B. Heppner (Ed.), Atlas of Neotropical

Lepidoptera. Vol. 5A. Association for Tropical Lepi-

doptera, Scientific Publishers.

Llorente-Bousquets, J. (1984). Sinopsis sistemática y

biogeográfica de los Dismorphiinae de México con

especial referencia al género Enantia Hübner (Lepi-

doptera: Pieridae). Folia Entomológica Mexicana,

58, 3–206.

Llorente-Bousquets, J., Garcés, A., & Luis-Martínez, A.

(1986). Las mariposas de JalapaTeocelo, Veracruz (El

Paisaje Teoceleño IV). Teocelo, 4, 14–37.

Llorente-Bousquets, J., & Escalante, P. (1994). Insular bio-

geography of submontane humid forests in México. In

S. P. Darwin, & A. L. Welden (Eds.), Biogeography of

Mesoamerica (pp. 139–146). University of Louisiana.

Llorente-Bousquets, J., & Luis-Martínez, A. (1993). A

conservation oriented analysis of Mexican butterflies:

the Papilionidae (Lepidoptera: Papilionoidea). In T. P.

Ramammorthy, J. Fa, R. Bye, & A. Lot (Eds.), The bio-

logical diversity of Mexico: origins and distributions

(pp. 147–177). Oxford University Press.

397

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Llorente-Bousquets, J., & Luis-Martínez, A. (1998). Análisis

conservacionista de las mariposas mexicanas: Papi-

lionidae (Lepidoptera: Papilionoidea). In T. Ramam-

morthy, R. Bye, A. Lot, & J. Fa (Eds.), Diversidad

Biológica de México: orígenes y distribución (pp.

149–178). Instituto de Biología, Universidad Nacional

Autónoma de México (UNAM), México.

Llorente-Bousquets, J., Luis-Martínez A., & Vargas-Fer-

nández, I. (2006). Apéndice general de Papilionoidea:

Lista sistemática, distribución estatal y provincias

biogeográficas. In J. J. Morrone, & J. Llorente-

Bousquets (Eds.), Componentes bióticos principales

de la entomofauna mexicana (Vol. II, pp. 945–1009).

Las Prensas de Ciencias, Universidad Nacional Autó-

noma de México (UNAM), México.

Llorente-Bousquets, J., Luis-Martínez, A., Vargas-Fernán-

dez, I., & Warren, A. D. (1996). Lista de las mariposas

del estado de Jalisco, México. Revista Sociedad Mexi-

cana Historia Natural, 46, 35–48.

Llorente-Bousquets, J., Luis-Martínez, A., Vargas-Fernán-

dez, I., & Warren, A. D. (2004). Butterflies of the state

of Nayarit, Mexico. Journal of the Lepidopterists’

Society, 58(4), 203–222.

Llorente-Bousquets, J., Trujano-Ortega, M., Luis Martí-

nez, A., & Vargas-Fernández, I. (2006). Patrones

de distribución de la familia Pieridae (Lepidoptera).

In J. J. Morrone, & J. Llorente-Bousquets (Eds.),

Componentes bióticos principales de la entomofau-

na mexicana (Vol. II, pp. 715–770). Las Prensas de

Ciencias, Universidad Nacional Autónoma de Méxi-

co (UNAM), México.

Llorente-Bousquets, J., Vargas-Fernández, I., Luis-Martí-

nez, A., Trujano-Ortega, M., Hernández-Mejía, B. C.,

& Warren, A. D. (2014). Biodiversidad de Lepidop-

tera en México. Revista Mexicana de Biodiversidad,

Suplemento Biodiversidad de México, 85, 353–371.

https://doi.org/10.7550/rmb.31830

Llorente-Bousquets, J., Warren, A. D., Vargas-Fernández,

I., & Luis-Martínez, A. (1996). Mariposas diur-

nas de Colima. Dugesiana, 3(2), 1-18. https://doi.

org/10.32870/dugesiana.v28i2.7165

Luis-Martínez, A. (1997). Papilionoidea (Lepidoptera:

Insecta) del estado de Michoacán: Fase I. Informe

final SNIB-CONABIO proyecto No. B150. UNAM,

México. http://www.conabio.gob.mx/institucion/pro-

yectos/resultados/InfB150.pdf

Luis-Martínez, A. (1999). Papilionoidea (Lepidoptera:

Insecta) del estado de Michoacán: Fase II. Informe

final SNIB-CONABIO proyecto No. H209. UNAM,

México. http://www.conabio.gob.mx/institucion/cgi-

bin/datos.cgi?Letras=H&Numero=209.pdf

Luis-Martínez, A. (2001). Papilionoidea (Lepidoptera:

Insecta) del estado de Michoacán: Fase III. Informe

final SNIB-CONABIO proyecto No. R256. UNAM,

México. http://www.conabio.gob.mx/institucion/pro-

yectos/resultados/Inf%20R256.pdf

Luis-Martínez, A., Ávalos-Hernández, O., Trujano-Ortega,

M., Arellano-Covarrubias, A., Vargas-Fernández, I.,

& Llorente-Bousquets, J. (2021). Riqueza y Ende-

mismo de Papilionoidea (Lepidoptera) de la región

Loxicha en el estado de Oaxaca, México. Dugesiana,

28(2), 233–246. https://doi.org/10.32870/dugesiana.

v28i2.7165

Luis-Martínez, A., Hernández-Mejía, B., Trujano-

Ortega, M., Warren, A. D., Salinas-Gutiérrez, J.,

Ávalos-Hernández, O., Vargas-Fernández, I., & Llo-

rente-Bousquets, J. (2016). Avances faunísticos en los

Papilionoidea (Lepidoptera) sensu lato de Oaxaca,

México. Southwestern Entomologist, 41(1), 171–224.

https://doi.org/10.3958/059.041.0119

Luis-Martínez, A., & Llorente-Bousquets, J. (1990). Maripo-

sas en el Valle de México: Introducción e Historia. 1.

Distribución local y estacional de los Papilionoidea de

la Cañada de los Dínamos, Magdalena Contreras, D. F.,

México. Folia Entomológica Mexicana, 78, 95–198.

Luis-Martínez, A., & Llorente-Bousquets, J. (1993). Mari-

posas. In I. Luna & J. Llorente (Eds.), Historia

Natural del Parque Ecológico Estatal Omiltemi,

Chilpacingo, Guerrero, México (pp. 307–385). Facul-

tad de Ciencias, Universidad Nacional Autónoma de

México (UNAM), México.

Luis-Martínez, A., Llorente-Bousquets, J., & Vargas-

Fernández, I. (2005). MARIPOSA database. (Una

megabase de datos de mariposas y la regionalización

biogeográfica de México). En J. Llorente-Bousquets,

& J. J. Morrone (Eds.), Regionalización geográfica

en Iberoamérica y tópicos afines: Primeras Jorna-

das Biogeográficas de la Red Iberoamericana de

Biogeografía y Entomología Sistemática (RIBES

XII.X-CYTED) (pp. 269-294). Las Prensas de Cien-

cias, Universidad Nacional Autónoma de México

(UNAM), México.

Luis-Martínez, A., Llorente-Bousquets, J., Vargas-Fernán-

dez, I., & Gutiérrez, A. L. (2000). Síntesis preliminar

del Conocimiento de los Papilionoidea (Lepidoptera:

Insecta) de México. In F. Martín, J. Morrone, & A.

Melic (Eds.), Monografías Tercer Milenio (Boletín

Sociedad Entomológica Aragonesa), 1, 275–285.

Luis-Martínez, A., Llorente-Bousquets, J., Vargas-Fernán-

dez, I., & Warren, A. D. (2003). Biodiversity and

Biogeography of mexican butterflies (Lepidoptera:

Papilionoidea and Hesperioidea). Proceedings of

the Entomological Society of Washington, 105(1),

209–224.

Luis-Martínez, A., Llorente-Bousquets, J., Warren, A. D.,

& Vargas-Fernández, I. (2004). Los lepidópteros:

Papilionoideos y Hesperoideos. In A. J. García-Men-

doza, M. J. Ordóñez, & M. A. Briones-Salas (Eds.),

Biodiversidad de Oaxaca (pp. 331–356). Instituto

de Biología, Universidad Nacional Autónoma de

398 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

México (UNAM)-Fondo Oaxaqueño para la Conser-

vación de la Naturaleza-WWF.

Luis-Martínez, A., Salinas-Gutiérrez, J. L., & Llorente-

Bousquets, J. (2011). Papilionoidea y Hesperioidea

(Lepidoptera: Rhopalocera). In F. Álvarez (Coord.).

Chiapas: Estudios sobre su diversidad biológica (pp.

363–392). Instituto de Biología, Universidad Nacio-

nal Autónoma de México (UNAM), México

Luis-Martínez, A., Sánchez García, A., Ávalos-Hernández,

O., Salinas-Gutiérrez, J. L., Trujano-Ortega, M.,

Arellano-Covarrubias, A., & Llorente-Bousquets, J.

(2020). Distribution and diversity of Papilionidae

and Pieridae (Lepidoptera: Papilionoidea) in Loxicha

Region, Oaxaca, Mexico. Revista de Biología Tropi-

cal, 68(1), 139–155. https://doi.org/10.15517/RBT.

V68I1.37587

Luis-Martínez, A., Trujano-Ortega, M., Llorente-Bousquets,

J., & Vargas-Fernández, I. (2006). Patrones de distri-

bución de las subfamilias Danainae, Apaturinae, Bibli-

dinae y Heliconiinae (Lepidoptera: Nymphalidae).

In J. J. Morrone, & J. Llorente-Bousquets (Eds.),

Componentes bióticos principales de la entomofau-

na mexicana (Vol. II, pp. 771–866). Las Prensas de

Ciencias, Universidad Nacional Autónoma de Méxi-

co (UNAM), México.

Luis-Martínez, A., Vargas-Fernández, I., & Llorente-Bous-

quets, J. (1991). Lepidopterofauna de Oaxaca I.

Distribución y Fenología de los Papilionoidea de la

Sierra de Juárez. Publicaciones Especiales del Museo

de Zoología, UNAM, 3, 1–121.

Luis-Martínez, A., Vargas-Fernández, I., & Llorente-Bous-

quets, J. (1996). Síntesis de los Papilionoidea (Rho-

palocera: Lepidoptera) del estado de Veracruz. Folia

Entomológica Mexicana, 93, 91–133.

Luna-Reyes, M. (2020). Mariposas diurnas (Lepidoptera).

La biodiversidad en Morelos: Estudio de Estado 2

(pp. 203–209). Comisión Nacional para el Conoci-

miento y Uso de la Biodiversidad (CONABIO) y

Gobierno del Estado de Morelos, México.

Luna-Reyes, M., Luis-Martínez, A., Vargas-Fernández, I.,

& Llorente-Bousquets, J. (2012). Mariposas del esta-

do de Morelos, México (Lepidoptera: Papilionoidea).

Revista Mexicana de Biodiversidad, 83, 623–666.

https://doi.org/10.7550/rmb.27987

Maza, J., & Maza, R. G. (1981). Notas sobre los Ithomiidae

de Mexico. IV: Zona Loxicha, Pacífico de Oaxaca.

Boletín informativo. Sociedad mexicana de Lepidop-

terología, 7(1), 3–10.

Maza, J., Maza, R. G., & Maza, R. F. (1982). Lepidópteros

nuevos del Estado de Guerrero, México. (Papilionoi-

dea). Revista de la Sociedad mexicana de Lepidopte-

rología, 7(1), 2–14.

Maza, R. G., & Maza, J. E. (1983). Descripción de nuevas

subespecies del género Callicore Hübner de México

y Guatemala (Nymphalidae: Nymphalinae). Revista

de la Sociedad mexicana de Lepidopterología, 8(1),

3–11.

Maza, R. G., & Maza, J. E. (1993). Mariposas de Chiapas.

Espejo de Obsidiana.

Meave del Castillo, J. A. & Luis-Martínez, A. (2000).

Caracterización biológica del Monumento Natural

Yaxchilán como un elemento fundamental para el

diseño de su plan rector de manejo. Hoja de cálculo

SNIB-CONABIO proyecto No. M099. Universidad

Nacional Autónoma de México, México.

Michán, L., Llorente-Bousquets, J., Luis-Martínez, A., &

Castro, D. J. (2004). Breve historia de la Taxonomía

de Lepidoptera en México durante el siglo XX. In

J. Llorente, J. J. Morrone, O. Yáñez, & I. Vargas

(Eds.), Biodiversidad, Taxonomía y Biogeografía

de Artrópodos de México: Hacia una Síntesis de su

Conocimiento (Vol. IV, pp. 5–42). Facultad de Cien-

cias, Universidad Nacional Autónoma de México

(UNAM), México.

Monteagudo-Sabaté, D., & Luis-Martínez, A. (2013).

Patrones de riqueza altitudinal de Papilionidae, Pie-

ridae y Nymphalidae (Lepidoptera: Rhopalocera)

en áreas montañosas de México. Revista Biología

Tropical, 61(3), 1509–1520.

Monteagudo-Sabaté, D., Luis-Martínez, A., Vargas-Fer-

nández, I., & Llorente-Bousquets, J. (2001). Patrones

altitudinales de la diversidad de mariposas en la

Sierra Madre del Sur (México). (Lepidoptera: Papi-

lionoidea). SHILAP Revista de Lepidopterología,

29(115), 207–237.

Monteagudo-Sabaté, D., Salinas-Gutiérrez, J. L., & Luis-

Martínez, A. (2014). Relaciones biogeográficas de

Papilionidae, Pieridae y Nymphalidae (Lepidoptera:

Papilionoidea) en el bosque mesófilo de montaña de

México. Entomotropica, 29(1), 29–37.

Morrone, J. J. (2005). Hacía una síntesis biogeográfica de

México. Revista Mexicana de Biodiversidad, 76(2),

207–252.

Morrone, J. J. (2017) Biogeographic regionalization of

the Sierra Madre del Sur province, Mexico. Revista

Mexicana de Biodiversidad, 88, 710–714. https://doi.

org/10.1016/j.rmb.2017.07.012

Morrone, J. J., Espinosa-Organista, D., & Llorente-Bous-

quets, J. (2002). Mexican biogeographic provinces:

preliminary scheme, general characterizations, and

synonymie. Acta Zoológica Mexicana (Nueva Serie),

85, 83–108.

Navarro-Sigüenza, A., Rebón-Gallardo, M. F., Gordillo-

Martínez, A., Peterson, A. T., Berlanga-García, H.,

& Sánchez-González. L. A. (2014). Biodiversidad de

Aves en México. Revista Mexicana de Biodiversidad,

Suplemento Biodiversidad de México, 85, S476–

S495. https://doi.org/10.7550/rmb.41882

399

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Oksanen, J., Blanchet, F., Friendly, M., Kindt, R., Legen-

dre, P., McGlinn, D., Minchin, P. R., O’Hara, R. B.,

Simpson, G. L., Solymos, P., Henry, M., Stevens,

H., Szoecs, E., & Wagner, H. (2019). Vegan: Com-

munity Ecology (Version 2.5-6, R package). https://

CRAN.R-project.org/package=vegan

Oñate-Ocaña, L., Trujano-Ortega, M., Llorente-Bousquets,

J., Luis Martínez, A., & Vargas-Fernández, I. (2006).

Patrones de distribución de la familia Papilionidae

(Lepidoptera). In J. J. Morrone, & J. Llorente-Bous-

quets (Eds.), Componentes bióticos principales de la

entomofauna mexicana (Vol. II, pp. 661–714). Las

Prensas de Ciencias, Universidad Nacional Autóno-

ma de México (UNAM), México.

Ornelas, J. F., Sosa, V., Soltis, D. E., Daza, J. M., González,

C., Soltis, P. S., & Ruiz-Sánchez, E. (2013). Compa-

rative phylogeographic analyses illustrate the com-

plex evolutionary history of threatened cloud forests

of northern Mesoamerica. PloS One, 8(2), e56283.

https://doi.org/10.1371/journal.pone.0056283

Pelham J. P. (2008). A catalogue of the butterflies of the

United States and Canada with a complete biblio-

graphy of the descriptive and systematic literature.

Journal of Research on the Lepidoptera, 40, 1–652.

Pinheiro, C. E. G., & Ortiz, J. V. C. (1992). Communities

of fruit-feeding butterflies along a vegetation gra-

dient in central Brazil. Journal of Biogeography, 19,

505–511. http://dx.doi.org/10.2307/2845769

Pozo, C. (2006). Los Rhopalocera de la región de Calak-

mul, Campeche: métodos de estudio, fenología y

su uso y monitoreo como indicadores de distur-

bio. Universidad Nacional Autónoma de México.

http://132.248.9.195/pd2006/0604448/Index.html

Pozo, C., Llorente-Bousquets, J., Luis-Martínez, A., Var-

gas-Fernández, I., & Salas, N. (2005). Reflexiones

acerca de los métodos de muestreo para mariposas

en las comparaciones biogeográficas. In J. Llorente,

& J. Morrone (Eds.), Regionalización geográfica

en Iberoamérica y tópicos afines: primeras jorna-

das biogeográficas de la Red Iberoamericana de

Biogeografía y Entomología Sistemática (RIBES

XII.I. –CYTED) (pp. 203–215). Universidad Nacional

Autónoma de México.

Pozo, C., Luis-Martínez, A., Llorente-Bousquets, J., Salas-

Suárez, N., Maya-Martínez, A., Vargas-Fernández,

I., & Warren, A. D. (2008). Seasonality and pheno-

logy of the butterflies (Lepidoptera: Papilionoidea

and Hesperioidea) of Mexico’s Calakmul region.

Florida Entomologist, 91, 407–422. https://doi.

org/10.1653/0015-4040(2008)91[407:SAPOTB]2.0.

CO;2

Pozo, C., Salas-Suárez, N., Prado-Cuéllar, B., & May-Uc,

E. (2009). Riqueza de mariposas diurnas (Lepidop-

tera: Rhopalocera) en el Santuario del Manatí y una

propuesta para su uso en el monitoreo de ambientes

terrestres del área. In J. Espinoza-Avalos, G. A.

Islebe, & H. A. Hernández-Arana (Eds.), El sistema

ecológico de la bahía de Chetumal/Corozal: costa

occidental del Mar Caribe. El Colegio de la Frontera

Sur, México.

R Core Team (2020). R: A language and environment for

statistical computing. R Foundation for Statistical

Computing, Vienna, Austria. https://www.R-project.

org/

Rydon, A. (1964). Notes on the use of butterfly traps in east

Africa. Journal Lepidopterists’ Society, 18, 51–58.

Rzedowski, J. (1978). Vegetación de México. Limusa,

México.

Salinas-Gutiérrez, J. L., Luis-Martínez, A., & Llorente-

Bousquets, J. (2004). Papilionoidea of the evergreen

tropical forests of Mexico. Journal of the Lepidopte-

rists’ Society, 58, 125–142.

Santiago-Alvarado, M., Montaño-Arias, G., & Espino-

sa, D. (2016). Áreas de endemismo de la Sierra

Madre del Sur. In I. Luna-Vega, D. Espinosa, & R.

Contreras-Medina (Eds.), Biodiversidad de la Sierra

Madre del Sur (pp. 431–448). Universidad Nacional

Autónoma de México (UNAM), México.

Santos, J. P., Iserhard, C. A., Teixeira, M. O., & Roma-

nowski, H. P. (2011). Fruit-feeding butterflies guide

of subtropical Atlantic Forest and Araucaria moist

forest in the State of Rio Grande do Sul, Brazil. Biota

Neotropica, 11, 253–274.

Soberón, J., & Llorente-Bousquets, J. (1993). The use of

species accumulation functions for the prediction of

species richness. Conservation Biology, 7(3), 480–488.

https://doi.org/10.1046/j.1523-1739.1993.07030480.x

Spade, P., Hamilton A. T., & Brown, J. (1988). The biology

of seven troidine swallowtail butterflies (Papilioni-

dae) in Colima, Mexico. Journal of Research on the

Lepidoptera, 26(1/4), 13–26.

Stanford, R. E. (1998). Collecting Lepidoptera in Sonora

and Sinaloa, Mexico, 26-31 December 1954. News of

the Lepidopterists’ Society, 40(3), 44.

Tapia-Sedeño, A. (2013). Análisis de los componentes alfa

y beta de la familia Nymphalidae (Papilionoidea:

Lepidoptera) en el estado de Michoacán (Tesis de

Licenciatura). Universidad Nacional Autónoma de

México (UNAM), México. http://132.248.9.41:8880/

jspui/handle/DGB_UNAM/TES01000704303

Uehara-Prado, M., Freitas, A. V. L., Francini, R. B., &

Brown, K. S. (2004). Guia das borboletas frugívoras

da Reserva Estadual do Morro Grande e região de

Caucaia do Alto, Cotia (São Paulo). Biota Neotropi-

ca, 4(1), 1-25.

Vargas-Fernández, I., Llorente-Bousquets, J., & Luis-Mar-

tínez, A. (1994). Listado Lepidopterofaunístico de la

Sierra de Atoyac de Álvarez en el estado de Guerrero:

400 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

notas acerca de su distribución local y estacional (Rho-

palocera: Papilionoidea). Folia Entomológica Mexica-

na, 86, 41–178.

Vargas-Fernández, I., Llorente-Bousquets, J., & Luis-Mar-

tínez, A. (1999). Distribución de los Papilionoidea

(Lepidoptera: Rhopalocera) de la Sierra de Manantlán

(250–1 650 m) en los estados de Jalisco y Colima.

Publicaciones Especiales del Museo de Zoología,

UNAM, 11, 1153.

Vargas-Fernández, I., Llorente-Bousquets, J., & Luis-Mar-

tínez, A. (2016). Adiciones a la serie Papilionoidea

de México: Distribución Geográfica e Ilustración.

Las Prensas de Ciencias, Universidad Nacional Autó-

noma de México (UNAM), México.

Vargas-Fernández, I., Luis-Martínez, A., Llorente-Bous-

quets, J., & Warren, A. D. (1996). Butterflies of the

state of Jalisco. Journal of the Lepidopterists’ Society,

50(2), 97–138.

Vargas-Fernández, I., Trujano-Ortega, M., Llorente-Bous-

quets, J., & Luis Martínez, A. (2006). Patrones de

distribución de las subfamilias Ithomiinae, Morphinae

y Charaxinae (Lepidoptera: Nymphalidae). In J. J.

Morrone, & J. Llorente-Bousquets (Eds.), Com-

ponentes bióticos principales de la entomofauna

mexicana (Vol. II, pp. 867–944). Las Prensas de

Ciencias, Universidad Nacional Autónoma de Méxi-

co (UNAM), México.

Vázquez, L. (1958). Notas sobre lepidópteros de las Islas

Revillagigedo. I. Anales del Instituto de Biología,

UNAM, 28(1/2), 301–307.

Vázquez, L. (1959). Notas sobre lepidópteros de las Islas

Revillagigedo. II. Anales del Instituto de Biología,

UNAM, 29(1/2), 349–353.

Vázquez, L. (1960). La isla Socorro. X. Observaciones

sobre los artrópodos. Monografías del Instituto de

Geofísica, Universidad Nacional Autónoma de Méxi-

co, 2, 217–234.

Velázquez, C. A. (1976). Reporte de un viaje de colecta a

los Estados de Michoacán, Jalisco, Colima y Oaxaca.

Boletín informativo. Sociedad Mexicana de Lepidop-

terología, 2(4), 6.

Velázquez, N. V., & Velázquez, C. A. (1975). Viaje de

colecta a Jalisco y Colima. Boletín informativo.

Sociedad Mexicana de Lepidopterología, 1(4), 6–7.

Warren, A. D., & Llorente-Bousquets, J. (1999). Butterflies

of Mismaloya, Jalisco and adjacent parts of Banderas

Bay and Southern Nayarit, México. Dugesiana, 6(1),

17–32. https://doi.org/10.32870/dugesiana.v6i1.3904

Warren, A. D., Vargas-Fernández, I., Luis-Martínez, A.,

& Llorente-Bousquets, J. (1996). Mariposas diur-

nas de Jalisco. Dugesiana, 3(1), 1–20. https://doi.

org/10.32870/dugesiana.v3i1.3795

Warren, A. D., Vargas-Fernández, I., Luis-Martínez, A., &

Llorente-Bousquets, J. (1998). Butterflies of the state

of Colima, México. Journal of the Lepidopterists’

Society, 52(1), 32–64.

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APPENDIX 1

Nymphalidae of the Loxicha Region across a vegetational-elevational gradient

Monthly vegetational and elevational distribution of 189 species of Nymphalidae in the Loxicha Region, Oaxaca,

Mexico. Records were obtained from 267 sampling days over seven years (2005, 2007, 2008, 2011–2014) in 21 sites from

eight municipalities. The first column (Sp.) is the species identifier number corresponding to the list of species in the Results

section. The next 11 columns are the months of the year excluding February from which no records are available. Column 13

(Total) is the total number of records for each species. The next five columns are the specimens recorded in each vegetation

type: TDF, tropical deciduous forest; TSDF, tropical subdeciduous forest; CF, cloud forest; OPCF, oak-pine forest with ele-

ments of high-elevation cloud forest; OPF, oak-pine forest. The next two columns (100 and >70 %) present the elevational

range where 100 % and more than 70% of the specimens of each species were recorded, respectively. These two columns

show the elevational range with the highest probability for finding the species, although some taxa undergo seasonal eleva-

tional migrations. The last column (%) refers to the percentage of specimens collected in the >70 % elevational range. The

last two rows correspond to the total number of specimens (TAb) and species (TSp) of each column. Species 55. Pedaliodes

dejecta ssp., 116. Adelpha donysa ssp., and 176. Actinote guatemalena guerrerensis were not recorded during fieldwork and

hence are not included in this table.

Sp. J M A M J J A S O N D Total TDF TSDF CF OPCF OPF 100 >70% %

1 10 6 8 3 2 29 10 18 1 80-900 80-500 97

2 1 9 10 3 7 500-2400 1200-2400 92

33 8 1 5 1 3 3 8 1 33 5 21 7 290-1650 100-500 81

4 1 1 7 9 9 290-1650 1100-1650 92

520 1 1 6 13 29 4 2 36 112 65 40 6 1 80-2400 80-410 93

6 223794311 2 43 17 21 5 80-900 80-500 88

7 1 1 1 1500-2400 1500-2400 100

81 2 3 9 25 11 1 2 54 53 1 100-2150 350-500 99

910 3 1 4 10 32 48 76 57 13 4 258 243 12 3 290-2150 350-500 93

10 2 2 2 1 3 3 31 6 8 58 1 57 350-1650 750-1250 69

11 4 1 3 4 12 9 33 14 19 350-1530 410-1250 97

12 10 3 1 3 2 12 9 43 30 6 5 124 51 73 350-1530 350-900 79

13 2 2 20 18 1 43 22 21 290-1650 350-1300 89

14 3 14 3 13 10 40 113 5 201 3 150 47 1 80-2150 350-900 94

15 63 15 7 6 11 18 58 115 119 12 3 427 86 341 350-1650 350-1300 98

16 2 7 21 91 21 142 142 290-1650 1100-1650 89

17 1 17 4 22 22 1200-1650 1200-1650 100

18 1 2 3 2 1 600-2400 600-850 67

19 1 8 9 9 350-500 350-500 100

20 3 4 2 8 3 18 76 35 14 4 167 1 165 1 410-1650 1100-1650 69

21 1 9 8 1 28 47 43 4 1200-2400 1200-1650 92

22 3 8 8 9 22 69 86 80 8 293 74 215 4 350-2150 350-940 74

23 5 3 33 8 75 17 18 159 157 2 80-410 80-100 99

24 1 1 2 3 3 3 1 14 4 10 410-1650 410-1650 100

25 1 1 1 8 11 10 1 1200-2400 1500-1650 82

26 10 6 1 9 16 9 53 30 2 136 1 117 15 3 410-2150 350-500 87

27 3 1 4 2 1 15 5 31 7 22 1 1 80-2150 350-500 72

28 8 1 2 1 3 4 19 13 3 3 80-900 80-900 100

29 31 48 10 42 14 21 7 145 51 1 370 104 244 22 80-900 80-500 95

30 8 4 26 52 80 9 13 4 196 116 14 59 7 100-2400 80-940 88

31 1 6 4 1 4 4 4 28 2 54 11 8 35 80-1650 80-1650 100

32 6 11 87 37 13 75 29 258 100 1 157 80-1650 80-1650 100

33 6 1 2 16 19 38 9 5 1 97 2 95 80-1550 940-1550 88

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Sp. J M A M J J A S O N D Total TDF TSDF CF OPCF OPF 100 >70% %

34 4 12 12 3 7 33 12 19 1 103 87 15 1 80-900 100-500 99

35 4 1 1 2 1 5 18 1 33 33 1200-1650 1200-1550 95

36 3 1 1 11 4 6 17 8 51 19 18 13 1 80-2400 100-500 73

37 1 1 1 410-500 410-500 100

38 711 5 4 5 19 25 31 26 7 1 141 86 52 3 410-2150 350-940 93

39 7 6 10 4 8 7 5 7 2 5 1 62 6 26 28 2 100-2150 350-940 77

40 3 7 1 4 10 31 5 20 20 1 1 103 12 82 8 1 80-2150 350-500 80

41 2 2 2 410-1530 410-1530 100

42 2 3 1 5 3 4 1 19 3 14 1 1 80-2150 350-500 77

43 1 1 2 2 1470-1550 1470-1550 100

44 8 19 17 5 15 38 57 2 1 162 55 75 32 80-1550 80-500 81

45 2436139118 138 72 4 387 1 214 171 1 80-2150 350-900 86

46 1 1 1 11 6 6 8 34 26 8 290-1300 350-900 95

47 1 1 1 500-900 500-900 100

48 3 8 9 4 1 2 27 10 11 6 80-1530 80-500 80

49 311385611 4 1 43 3 38 2 80-2150 350-500 90

50 1 1 1 350-500 350-500 100

51 3 1 1 1 6 6 350-500 350-500 100

52 2 2 1 1 350-1300 350-1300 100

53 2 1 2 5 3 2 1470-2150 1470-2150 100

54 1 1 2 2 500-1550 1200-1550 75

56 66 85 56 155 34 50 435 63 1 2 947 148 645 151 3 80-2400 350-700 70

57 53 1 12 17 44 5 17 1 150 54 96 410-1650 350-1300 86

58 6 42 6 13 58 55 66 103 87 41 4 481 4 153 324 80-1650 350-1530 98

59 1 3 32 215 89 5 9 354 18 308 28 80-1650 350-500 86

60 1 1 1 2020-2150 2020-2150 100

61 8 4 3 2 8 35 52 11 1 124 121 3 1100-2150 1100-2150 100

63 20 1 4 2 7 2 36 5 31 350-1650 350-1650 100

64 1 1 2 1 5 4 1 1100-1530 1100-1530 100

62 4 1 1 5 12 30 10 30 2 95 95 444-1650 1100-1550 77

65 1 6 1 1 3 12 3 9 1200-2400 2280-2400 75

66 1200-1530 1200-1530 100

67 2 4 51 18 62 34 62 61 294 22 270 2 410-2400 750-1550 73

68 194 100 46 91 346 269 292 339 496 81 13 2267 43 791 1430 3 80-2400 350-1650 98

69 2 2 2 2280-2400 2280-2400 100

70 4 30 58 13 3 4 68 180 1 1 161 17 2020-2800 2280-2400 88

71 3 2 1 1 4 1 12 4 8 290-1650 290-1650 100

72 3 2 4 12 2 6 6 1 36 35 1 240-2150 350-500 95

73 1 1 1 4 5 6 2 20 18 1 1 350-2150 350-500 93

74 35 21 12 13 28 102 62 98 66 437 16 399 20 2 80-2150 350-500 91

75 1 1 1 410-500 410-500 100

76 3 2 1 12 3 37 9 16 2 1 86 71 15 350-1530 350-1530 100

77 4 4 4 2280-2400 2280-2400 100

78 1 9 3 5 18 18 1100-1650 1100-1650 100

79 5 2 1 3 1 12 8 1 3 80-2400 80-2400 100

80 3 3 1 2 1 4 12 14 1 1 42 3 35 2 2 80-2150 350-500 86

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Sp. J M A M J J A S O N D Total TDF TSDF CF OPCF OPF 100 >70% %

81 6 3 1 25 71 54 24 21 39 1 245 24 206 11 4 80-2150 350-500 84

82 12 5 6 40 17 5 7 10 1 103 47 40 15 1 2280-2400 100-500 85

83 2 1 5 2 5 2 1 18 18 500-1650 1100-1650 76

84 4 8 3 7 45 41 23 56 26 213 6 9 198 100-1550 80-900 86

85 1 2 3 1 2 410-850 410-850 100

86 5 5 3 19 7 63 20 107 20 249 14 208 26 1 80-2150 350-500 85

87 6 3 5 13 1 28 26 2 80-410 80-410 100

88 21 196 41 68 53 91 16 56 28 570 158 305 107 80-1650 80-500 83

89 11 90 26 32 105 18 1 4 5 292 276 16 80-500 80-100 94

90 4 6 2 38 40 38 10 91 46 6 281 86 175 19 1 80-2150 80-500 95

91 6 2 3 100 29 5 1 146 119 22 5 80-1530 80-100 77

92 6 4 1 3 35 46 16 12 8 2 1 134 30 71 30 3 80-2150 350-500 54

93 1 2 1 4 8 3 5 350-900 750-900 63

94 7 6 1 11 14 9 11 2 61 2 26 31 2 100-2150 80-940 91

95 2 4 2 28 55 53 13 157 71 86 290-1650 350-1300 93

96 2 3 5 1 11 11 80-500 80-100 92

97 2 8 15 23 2 1 4 55 41 14 350-1650 350-900 98

98 3 3 1 3 10 3 7 80-410 80-410 100

99 16 4 2 11 59 48 99 84 2 2 327 17 295 14 1 80-2150 350-500 94

100 7 5 4 17 11 22 24 56 50 196 11 179 4 2 80-2150 350-500 92

101 3 2 2 174 24 3 3 211 211 80-100 80-100 100

102 9 6 1 1 1 6 7 6 34 12 83 10 72 1 350-2150 1000-1650 75

103 1 1 1 500-900 500-900 100

104 6 12 5 27 55 60 22 23 37 1 1 249 30 217 1 1 80-2150 80-500 98

105 71 17 4 11 21 109 171 179 338 108 1029 55 972 2 410-2150 1100-1650 80

106 8 7 1 2 24 20 17 59 8 146 11 135 410-1650 1100-1650 73

107 2 2 10 14 29 20 77 4 58 15 80-2150 350-500 80

108 1 3 1 3 8 3 20 8 47 46 1 940-2400 1470-1650 75

109 3 22 25 22 34 16 122 1 2 119 410-1250 600-940 95

110 11 2 4 27 51 45 70 116 2 328 73 232 22 1 80-2150 350-500 71

111 1 3 2 16 22 1 9 12 80-940 410-940 96

112 3 3 3 410-500 410-500 100

113 6 3 4 2 7 17 14 117 49 8 227 201 5 21 350-2150 350-500 90

114 5 4 1 2 3 15 15 2280-2400 2280-2400 100

115 1 1 1 2280-2400 2280-2400 100

116

117 5 1 1 1 8 5 2 1 100-2400 100-500 89

118 5 4 1 4 15 6 23 52 40 7 157 14 84 44 15 100-2400 80-900 70

119 1 1 5 3 16 45 71 26 44 1 410-2150 410-940 99

120 1 4 2 7 6 1 500-2150 500-1530 95

121 12 3 4 14 11 10 54 53 1 600-2150 1100-1650 95

122 2 3 5 2 3 410-1300 410-900 94

123 2 1 5 4 12 2 8 2 80-2150 80-500 91

124 5 1 3 2 5 4 3 6 29 4 17 3 5 100-2150 100-900 86

125 2 3 6 4 4 8 27 16 6 5 290-2150 400-900 88

126 3 2 8 2 6 11 32 1 30 1 350-2150 1100-1650 82

127 4 2 5 4 5 20 18 2 290-1530 290-500 87

404 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Sp. J M A M J J A S O N D Total TDF TSDF CF OPCF OPF 100 >70% %

128 3 1 8 3 6 2 2 1 26 8 13 4 1 80-2150 80-2150 100

129 1 1 2 36 7 17 11 25 7 2 109 28 57 24 80-1300 350-940 73

130 4 4 4 1470-1550 1470-1550 100

131 33 15 3 19 18 20 34 180 41 10 6 379 6 317 55 1 80-2150 350-500 83

132 77 41 9 134 113 157 148 173 185 182 10 1229 191 340 693 4 1 80-2400 80-940 72

133 4 4 4 1200 1200 100

134 14 8 3 16 2 11 11 25 10 100 1 79 20 350-2400 1100-1650 83

135 3 3 1 1 3 6 8 24 11 1 61 3 57 1 350-2400 1100-1650 77

136 23 12 4 6 9 12 32 57 99 21 1 276 8 242 26 350-2400 1100-1650 80

137 5 2 13 14 9 43 43 444-1650 1100-1650 88

138 10 3 2 1 8 24 24 1500-2400 2280-2400 75

139 3 1 4 1 3 1500-2400 2280-2400 75

140 2 2 2 940 940 100

141 1 5 4 22 8 1 21 1 63 1 12 49 1 80-2400 2280-2400 75

142 11 1 2 22 11 17 24 5 93 22 70 1 350-2150 350-980 65

143 71 54 6 12 85 100 30 123 114 4 599 188 283 124 4 80-2400 350-500 72

144 88 29 7 13 37 171 104 203 165 88 9 914 34 548 328 4 100-2400 350-940 88

145 2 2 5 2 7 24 42 16 24 2 100-2400 80-500 95

146 2 1 1 1 1 2 1 9 4 1 4 350-2400 350-2400 100

147 3 1 6 1 11 9 2 350-1530 350-500 82

148 5 5 5 410-500 410-500 100

149 1 1 1 2280-2400 2280-2400 100

150 1 3 1 4 12 10 3 33 22 89 21 65 3 350-2400 350-940 65

151 6 2 1 5 17 3 34 68 1 66 1 350-1650 1100-1650 84

152 29 7 26 48 81 185 124 176 195 35 10 916 1 343 562 10 80-2400 350-1300 88

153 53 30 11 14 49 44 48 62 78 38 427 6 41 380 80-1650 1000-1650 69

154 1 1 8 15 20 59 40 42 14 200 11 89 100 80-1650 410-1650 95

155 14 1 4 19 1 18 410-2400 2280-2400 90

156 2 15 9 15 17 76 61 1 196 22 167 4 3 80-2400 350-500 85

157 3 5 6 51 104 47 47 12 275 2 162 109 2 80-2400 350-500 66

158 5 1 87 130 23 106 31 1 384 317 66 1 80-1300 80-100 82

159 1 1 4 6 1 5 80-2400 2280-2400 83

160 1 9 7 1 9 27 1 26 350-2400 2280-2400 87

161 1 1 1 1100-1300 1100-1300 100

162 1 1 20 2 24 24 500-2400 2280-2400 88

163 1 9 2 1 1 5 19 19 2280-2400 2280-2400 100

164 167 91 17 8 32 24 63 100 74 51 2 629 109 519 1 290-2150 750-1650 78

165 42 21 25 110 104 62 99 30 122 24 639 3 493 143 410-2400 1100-2400 98

166 9 20 10 10 58 1 41 87 26 6 268 267 1 444-2150 1100-1300 92

167 7 60 29 1 6 103 100 3 1100-2400 1100-1650 98

168 85 3 7 29 44 35 14 96 146 13 8 480 239 187 41 13 80-2400 80-500 88

169 25 22 11 12 30 14 32 44 23 7 220 1 213 6 235-2400 1000-1650 95

170 7 29 1 50 49 17 15 42 46 13 269 237 32 940-2400 1000-1650 87

171 31 14 6 12 3 1 2 15 84 2 79 3 290-2400 1000-1650 91

172 14 3 3 19 39 1 38 1200-2450 2280-2400 88

173 1 1 6 8 8 2280-2400 2280-2400 89

174 4 9 1 10 4 31 33 49 13 1 155 15 140 290-1650 530-1650 86

405

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Sp. J M A M J J A S O N D Total TDF TSDF CF OPCF OPF 100 >70% %

175 1 2 1 3 3 7 98 1 116 1 67 48 350-2400 1100-2400 96

176

177 5 1 2 6 18 24 5 6 3 70 47 14 4 5 100-2400 80-500 87

178 2 16 2 3 31 18 5 13 1 1 92 1 51 23 17 80-2400 350-500 57

179 2 4 10 14 1 1 4 5 64 4 109 3 15 87 4 80-2400 1470-2400 89

180 59 21 8 25 44 111 93 159 135 36 18 709 30 503 172 4 80-2400 350-940 86

181 3 1 4 4 410-500 410-500 100

182 6 3 3 14 12 74 9 11 132 124 4 4 290-2150 350-500 92

183 3 2 1 3 7 28 44 1 40 3 410-1550 410 81

184 54 35 15 23 11 54 81 64 142 35 7 521 48 259 193 21 80-2400 80-940 81

185 97 22 19 33 61 113 68 112 123 23 20 691 32 643 15 1 80-2150 80-500 98

186 13 7 5 6 17 18 8 28 25 127 125 2 600-2400 1100-1650 93

187 1 1 1 1200-1530 1200-1530 100

188 1 1 2 1 1 80-410 80-410 100

189 12 9 9 20 24 61 44 18 41 1 239 42 187 3 7 80-2400 350-500 79

TAb 1914 1432 610 1746 2688 3938 3443 5570 6002 1256 163 28756 3496 11699 12487 1051 23

TSp 109 106 96 120 120 126 135 146 159 91 40 189 79 36 147 108 4

APPENDIX 2

Nymphalidae of the Loxicha Region captured in Van Someren-Rydon traps

Monthly and elevational distribution of Nymphalidae species captured with Van Someren-Rydon traps in the Loxicha

Region, Oaxaca, Mexico. The list includes 94 species of seven of the 11 subfamilies of Nymphalidae, representing 50 %

of all species recorded and 41 % of all specimens collected. The first column (Sp.) is the species identifier number cor-

responding to the list of species in the Results section. The next 11 columns are the months of the year excluding February

from which no records are available. Column 13 (VSRT) is the number of records of each species obtained in Van Someren-

Rydon traps. Column 14 (Total) is the total number of specimens captured during our study. The next column (%) shows

the percentage of specimens captured with traps out of the total number of specimens. The last three columns represent the

number of specimens collected in traps at each elevational level (m): 1, 0–750; 2, 750–1 800, and 3, 1 800–2 850. The last

two rows correspond to the total number of specimens (TAb) and species (TSp) of each column.

Sp. J M A M J J A S O N D VSRT Total % 1 2 3

23 5 3 33 4 69 17 18 149 159 93.71 149

24 1 1 2 2 3 1 10 14 71.43 4 6

25 1 1 1 8 11 11 100 10 1

26 10 6 6 13 8 39 25 2 109 136 80.15 94 14 1

27 3 1 4 2 1 15 3 29 31 93.55 27 1 1

28 8 1 2 1 3 4 19 19 100 16 3

29 30 47 9 38 13 20 7 135 48 1 348 370 94.05 327 21

30 8 4 26 48 68 9 13 4 180 196 91.84 114 59 7

31 1 5 2 1 3 4 3 27 2 48 54 88.89 14 34

32 6 11 82 34 12 72 25 242 258 93.80 96 146

33 6 1 2 14 17 36 9 5 1 91 97 93.81 2 89

34 411 12 3 7 31 11 15 1 95 103 92.23 94 1

35 4 1 1 2 1 5 18 1 33 33 100 33

36 3 1 1 11 4 6 17 8 51 51 100 37 13 1

37 1 1 1 100 1

38 711 5 4 4 17 25 30 26 7 1 137 141 97.16 84 50 3

406 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 363-407, January-December 2022 (Published Jun. 07, 2022)

Sp. J M A M J J A S O N D VSRT Total % 1 2 3

39 7 5 10 4 8 7 5 7 2 4 1 60 62 96.77 31 27 2

40 3 6 1 4 10 28 5 19 20 1 1 98 103 95.15 90 7 1

41 2 2 2 100 2

42 2 3 1 5 3 4 1 19 19 100 17 1 1

43 1 1 2 2 100 2

44 8 13 15 5 14 31 41 2 1 130 162 80.25 100 30

45 2 4 2 6 1 36 106 122 66 4 349 387 90.18 192 156 1

46 1 1 1 10 3 6 8 30 34 88.24 24 6

47 1 1 1 100 1

48 2 7 7 4 1 21 27 77.78 17 4

49 3 1 1 3 7 5 5 10 4 1 40 43 93.02 38 2

50 1 1 1 100 1

51 3 1 1 1 6 6 100 6

52 2 2 2 100 1 1

53 2 1 2 5 5 100 3 2

54 1 1 2 2 100 2

56 60 71 44 138 28 42 356 50 1 1 791 947 83.53 672 117 2

57 53 1 10 17 44 3 15 1 144 150 96 53 91

58 6 41 6 9 48 49 60 84 65 37 4 409 481 85.03 132 277

59 1 3 27 182 81 4 9 307 354 86.72 283 24

60 1 1 1 100 1

61 7 2 2 8 27 52 11 1110 124 88.71 108 2

62 4 1 1 5 7 20 10 30 2 80 95 84.21 80

63 13 4 2 2 21 36 58.33 1 20

64 1 1 1 1 4 5 80 4

65 6 1 2 9 12 75 2 7

67 2 3 43 13 54 24 51 52 242 294 82.31 18 222 2

68 171 84 41 86 305 228 267 303 443 76 12 2016 2267 88.93 770 1245 1

70 10 48 1 1 18 78 180 43.33 78

71 3 2 1 1 4 1 12 12 100 4 8

72 3 2 1 5 2 3 6 1 23 36 63.89 22 1

73 1 1 1 3 4 5 2 17 20 85 16 1

74 34 12 11 8 19 89 57 71 53 354 437 81.01 336 16 2

75 1 1 1 100 1

76 3 2 1 10 3 33 9 15 1 1 78 86 90.70 66 12

77 4 4 4 100 4

78 1 7 2 3 13 18 72.22 13

79 1 2 1 3 1 8 12 66.67 5 3

80 1 3 1 3 11 11 1 1 32 42 76.19 31 1

81 15 10 25 245 10.20 24 1

84 3 3 2 7 33 31 15 46 21 161 231 69.70 12 149

86 5 5 3 13 3 47 19 87 15 197 249 79.12 173 23 1

87 3 3 3 9 18 28 64.29 18

88 19 154 30 50 39 71 14 47 22 446 570 78.25 359 87

89 11 66 19 27 84 16 1 4 4 232 292 79.45 232

90 4 6 2 38 35 28 8 66 38 2 227 281 80.78 216 11

91 14 5 1 20 145 13.79 20

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Sp. J M A M J J A S O N D VSRT Total % 1 2 3

92 4 2 35 32 10 4 8 1 96 134 71.64 86 9 1

93 1 2 3 8 37.50 3

94 7 4 1 11 14 8 11 1 57 61 93.44 28 27 2

95 2 4 2 22 45 46 13 134 157 85.35 57 77

96 1 5 1 7 11 63.64 7

97 2 4 14 20 2 1 4 47 55 85.45 39 8

98 2 2 1 1 6 10 60 6

99 12 2 1 11 43 33 78 66 2 1 249 327 76.15 236 13

100 7 5 4 13 10 20 18 39 34 150 196 76.53 144 4 2

101 1 9 10 211 4.74 10

102 4 6 1 1 1 1 4 2 10 8 38 83 45.78 1 36 1

104 3 1 1 6 3 4 18 249 7.23 18

105 13 1 22 69 105 1029 10.20 6 99

106 2 2 146 1.37 2

107 1 3 1 5 77 6.49 3 2

108 3 2 4 9 47 19.15 8 1

113 1 4 5 227 2.20 5

119 3 3 8 14 71 19.72 13 1

128 3 1 7 3 5 1 2 1 23 26 88.46 18 4 1

129 1 1 2 33 7 15 10 24 7 2 102 109 93.58 78 24

131 30 12 3 17 11 20 34 146 34 8 6 321 379 84.70 277 43 1

132 76 41 9 133 113 148 148 173 185 182 10 1218 1229 99.10 525 688 5

134 10 6 3 13 2 5 6 8 9 62 100 62 1 49 12

135 3 2 1 1 2 4 7 18 6 1 45 61 73.77 3 41 1

136 21 6 5 6 10 20 56 45 169 276 61.23 2 155 12

137 4 1 8 12 8 33 43 76.74 33

138 10 3 1 5 19 24 79.17 19

139 1 1 4 25 1

141 6 1 1 1 9 63 14.29 5 4

142 1 2 3 3 10 4 23 93 24.73 8 14 1

143 56 48 5 10 60 77 24 88 83 4 455 599 75.96 357 96 2

TAb 731 746 290 912 1080 1736 1357 2409 2061 463 49 11543 15824 72.95 6609 4709 191

TSp 49 53 43 57 57 63 69 68 76 38 20 94 94 74 69 42