An updated list of butterflies (Lepidoptera, Papilionoidea) of two Guatemalan seasonally dry forests

Abstract

Guatemala has a great diversity of butterflies, although there have been few intensive surveys on Lepidoptera in the country so far. We present an updated list of 218 species in 149 genera, 19 subfamilies, and six families of butterflies sampled at two seasonally dry forests in the Salamá and Motagua valleys in central and eastern Guatemala, by integrating new data from field surveys conducted in 2014–2021 into our previously published data (Yoshimoto et al. 2018, 2019), with Amblyscirteselissaelissa Godman, 1900, Repensflorus (Godman, 1900), and Niconiadesnikko Hayward, 1948 (Hesperiidae: Hesperiinae) as new country records. We collected a hairstreak species, Chalybshassan (Stoll, 1790) (Lycaenidae: Theclinae), at the Motagua Valley site, representing the second record for Guatemala since the early 20^th century, after we rediscovered it at the Salamá Valley site in 2011 and 2012 (Yoshimoto and Salinas-Gutiérrez 2015). Nymphalidae and Hesperiidae had larger numbers of species than the other four families at both sites. In Pieridae and Nymphalidae, species composition was similar between the sites, whereas in Lycaenidae, Riodinidae, and Papilionidae it differed more greatly between the sites. These results confirm the relatively high lepidopteran diversity of Guatemalan dry forests, noteworthy for the small areas that comprise the study sites, and represent marked similarities and differences in butterfly fauna and phenology within these forests.

Introduction

Neotropical seasonally dry forests are rich in flora and fauna (Pennington et al. 2006; Dirzo et al. 2011), although their ecosystems have been deteriorating because of various anthropogenic disturbances, such as deforestation due to agricultural expansion (e.g., Chazdon et al. 2011). Dry forests in Guatemala also harbor high lepidopteran diversity as well; our previous studies documented more than 150 and 100 butterfly species at the two small forest reserves in central and eastern Guatemala, respectively (Yoshimoto et al. 2018, 2019). These species lists, however, are still incomplete, and obviously, more species remain to be sampled at these sites. Moreover, we detected marked seasonal patterns in butterfly species richness and several conspicuous differences in the lepidopteran fauna between the two sites (Yoshimoto et al. 2019). Thus, it was apparent that additional field surveys were needed to make quantitative between-site comparisons of species composition, in order to enhance our understanding of butterfly fauna and phenology of these forests.

In Guatemala, approximately 400 species of Hesperiidae and nearly 700 species of the remaining families of Papilionoidea have been reported (Austin et al. 1998; Barrios et al. 2006; Salinas-Gutiérrez et al. 2009, 2012; Salinas-Gutiérrez 2013). Despite such high lepidopteran diversity, Guatemala’s butterfly fauna has been studied less intensively compared to neighboring countries; for example, in Mexico, exhaustive species lists for the whole country and for several states have been published (de la Maza et al. 1989, 1991; Luis-Martínez et al. 2011, 2016; Llorente-Bousquets et al. 2014), whereas Guatemala has had little research on Lepidoptera and few published inventories since the 20^th century (but see Austin et al. 1996 and Yoshimoto et al. 2021). Continued field surveys in various parts of Guatemala are thus important to fill a gap in our knowledge of the Neotropical butterfly fauna, which will in turn contribute to biodiversity conservation in the country.

Here, we present an updated and integrated list of papilionoid species (including Hesperiidae; van Nieukerken et al. 2011) for the same dry forest sites where we conducted our previous studies (Yoshimoto et al. 2018, 2019), by adding the new data from subsequent field surveys performed in 2014–2021, correcting identification errors, and modifying some of the species names based on taxonomic changes. Additionally, we examine between-site differences in butterfly fauna by comparing species composition at the family level, and identify seasonal patterns at the species level.

Materials and methods

This study was conducted at the Los Cerritos Municipal Park (hereafter, Los Cerritos; Fig. 1a) in the Salamá Valley (a subwatershed of the Chixoy region) of Baja Verapaz Department in central Guatemala (15°05'N, 90°18'W, 960–1160 m a.s.l., 69 ha), and at the Heloderma Natural Reserve (hereafter, Heloderma Reserve; Fig. 1b) in the Motagua Valley of Zacapa Department in eastern Guatemala (14°53'N, 89°47'W, 510–790 m a.s.l., 58 ha). The rainfall patterns are similar between the two areas, in which the rainy season usually begins in late May and ends in October; these six months were accordingly defined as the rainy season and the remaining months (November-April) as the dry season. This climatic trait fits the definition of seasonally dry tropical forests (4–6 months with rainfall being < 100 mm; Dirzo et al. 2011); see fig. 1 in Yoshimoto et al. (2018) and fig. 3 in Yoshimoto et al. (2019) for detailed precipitation information of each area.

Figure 1.

a Forest landscape of Los Cerritos Municipal Park and bHeloderma Natural Reserve.

The vegetation of both regions is characterized by an abundance of various aculeate plants such as cacti (Cactaceae). The most dominant species is a columnar cactus Stenocereuspruinosus (Otto) Buxb., with Pilosocereusleucocephalus (Poselg.) Byles & G. D. Rowley and Pereskialychnidiflora DC., also being abundant at both sites. On the other hand, there exist some marked differences in flora and in forest landscape. Heloderma Reserve has a dense forest with many arboreal species such as Bucidamacrostachya Standl. (Combretaceae), Lysilomadivaricatum (Jacq.) J. F. Macbr., Leucaenacollinsii Britton & Rose (both Mimosaceae), and Burseraexcelsa (Kunth) Engl. (Burseraceae), all of which can grow taller than the columnar cactus (Ariano-Sánchez and Salazar 2015; D. Ariano-Sánchez, pers. comm.; Fig. 1b). By contrast, none of these species have been reported from Los Cerritos (M. R. Álvarez, pers. comm.), where there are fewer high arboreal species and abundant shrubs and herbaceous plants (thus commonly called a spiny bush or scrub), thereby the columnar cactus being prominent in its forest landscape (Fig. 1a).

Field surveys were conducted on 21 days from July 2014 to August 2021 at Los Cerritos, and on 19 days from October 2017 to November 2021 at Heloderma Reserve. We collected adult butterflies with an insect net or photographed them in the daytime (09:00–17:00) at each site and in neighboring areas (a small garden at the foot of Los Cerritos and on a farm road adjacent to Heloderma Reserve). The individuals collected were mounted as voucher specimens and were deposited at the Colección de Artrópodos, Laboratorio de Entomología Sistemática, Universidad del Valle de Guatemala. All the individuals collected or photographed were identified to species or subspecies according to Warren et al. (2017). We did not include data for specimens that were not identified to species, except for Calephelis spp. (Riodinidae) and Bolla sp. (Hesperiidae: Pyrginae); see the footnotes of the Appendix 1 for the rationales for the inclusion of these data. We added all these data to our previous data (Yoshimoto et al. 2018, 2019), corrected identification errors, and modified scientific names of some of the species based on taxonomic changes, in order to compile an updated and integrated species list of the two sites. Note that the sampling methods were partially different in the previous surveys; only netting was done at Los Cerritos from January 2011 to November 2012, whereas at Heloderma Reserve between February 2016 and March 2017, data were obtained through netting, photographing, and observation; see Yoshimoto et al. (2018, 2019) for detailed information on the sampling methods for each site.

The site-level estimated species richness was calculated by using the Chao II index (Chao et al. 2005; Gotelli and Colwell 2011), after pooling the data across observation dates for each month for each site. The total estimated species richness was similarly obtained after pooling these data across both sites. The Jaccard dissimilarity index was used to quantify the between-site similarity in species composition; this index was calculated for all data and for each of the six families (Papilionidae, Pieridae, Lycaenidae, Riodinidae, Nymphalidae, and Hesperiidae). All the analyses were performed using R 4.1.2. (R Development Core Team 2021) with the package Vegan (Oksanen et al. 2020).

Results

By integrating our previous data (Yoshimoto et al. 2018, 2019), a total of 218 species (including one unidentified taxon and 107 subspecies) in 149 genera from 19 subfamilies of six families were recorded at the two sites (Appendix 1). Hesperiidae was the richest family (71 species), followed by Nymphalidae, Lycaenidae, Pieridae, Riodinidae, and Papilionidae (66, 36, 20, 16, and 9 species, respectively). Los Cerritos had 166 species in 117 genera, and Heloderma Reserve had 139 species in 107 genera (Appendix 1), 16 and 41 species of which had been newly recorded in the subsequent surveys, respectively (Fig. 2). The estimated species richness (mean±SE) of each site based on the Chao II index is 216.27±16.35 and 187.35±17.74, respectively, indicating that approximately 76.8% and 74.2% of the species inhabiting each site were sampled. The total estimated species richness for both sites is 272.80±17.85 (79.9%).

Figure 2.

Six of the species that were newly recorded in the present study at Los Cerritos or Heloderma Reserve a, bHeraclidesrumiko Shiraiwa & Grishin, 2014 (Papilionidae) cArchaeopreponademophoncentralis (Fruhstorfer, 1905) dCaligotelamoniusmemnon (C. Felder & R. Felder, 1867) (both Nymphalidae) eAbaeisnicippe (Cramer, 1779) (Pieridae) fLeptotescassiuscassidula (Boisduval, 1870) (Lycaenidae) gPirunaaea (Dyar, 1912) (Hesperiidae) a–d, gHeloderma Reserve e, f Los Cerritos. Note that P.aea had already been collected and identified to genus (Piruna sp.1) by Yoshimoto et al. (2019).

We detected identification errors for 20 individuals (identified as 11 species in our previous studies) and determined them to represent 13 species in this study; ten individuals from Los Cerritos and ten from Heloderma Reserve have been determined to number eight and six different species, respectively, with one species, Cissiathemis, shared between sites (Table 1). Additionally, Yoshimoto et al. (2019) incorrectly listed Piruna (Hesperiidae: Heteropterinae) in the subfamily Hesperiinae.

Table 1.

Butterfly species that were sampled at Los Cerritos and Heloderma Reserve (abbreviated as LC and HR, respectively) and were misidentified in Yoshimoto et al. (2018, 2019). Corrected species names are shown in bold.

Family	Species		Sampling month, year, and site
	Correct identification	Previous identification
Papilionidae	Heraclidesrumiko Shiraiwa & Grishin, 2014	Heraclidescresphontes (Cramer, 1777) A	Oct 2016 HR*
Pieridae	Abaeisnicippe (Cramer, 1779)	Pyrisitiaproterpia (Fabricius, 1775) A	Jul 2016 HR
Lycaenidae	Strymonmegarus (Godart, [1824])	Strymonmelinusfranki W. D. Field, 1938 A	Oct 2016 HR
Nymphalidae	Anthanassatulcis (H. Bates, 1864)	Anthanassadracaenaphlegias (Godman, 1901) B	May 2011 LC
Nymphalidae	Chlosyneerodyleerodyle (H. Bates, 1864)	Chlosynelacinialacinia (Geyer, 1837) B	Oct 2011 LC, Jul 2012 LC
Nymphalidae	Chlosynerositarosita A. Hall, 1924	Chlosynelacinialacinia (Geyer, 1837) B	Sep 2011 LC
Nymphalidae	Cissiasimilis (A. Butler, 1867)	Cissiapompilia (C. Felder & R. Felder, 1867) B	May 2012 LC, Jun 2012 LC
Nymphalidae	Cissiathemis (A. Butler, 1867)	Cissiapompilia (C. Felder & R. Felder, 1867) A, B	Aug 2011 LC, Aug 2016 HR**, Oct 2016 HR
Hesperiidae	Urbanusviterboana (Ehrmann, 1907)	Urbanusproteusproteus (Linnaeus, 1758) B	Nov 2011 LC
Hesperiidae	Heliopetesmacairamacaira (Reakirt, [1867])	Heliopyrgusdomicelladomicella (Erichson, [1849]) B	Jul 2012 LC
Hesperiidae	Amblyscirteselissaelissa Godman, 1900	Piruna sp.1 A	Aug 2016 HR, Sep 2016 HR
Hesperiidae	Copaeodesaurantiaca (Hewitson, 1868)	Ancyloxyphaarene (W. H. Edwads, 1871) B	Mar 2011 LC
Hesperiidae	Cymaenestrebius (Mabille, 1891)	Cymaenestripunctustheogenis (Capronnier, 1874) A	Sep 2016 HR**

^A Listed in Yoshimoto et al. (2019). ^B Listed in Yoshimoto et al. (2018). *Specimen not collected (recorded only by photographing; see Fig. 2a, b for its images). **Two individuals collected.

The following three skipper species (Hesperiidae: Hesperiinae) were recorded for the first time in Guatemala:

• Amblyscirteselissaelissa Godman, 1900. Reserva Heloderma, Cabañas, Zacapa, GUATEMALA. Three specimens: 30-08-2016, J442; 26-09-2016, J478; 01-06-2018, J769. Collected by Jiichiro Yoshimoto. Identified by Andrew D. Warren. Note that the two individuals (J442 and J478) were misidentified as Piruna sp.1 in Yoshimoto et al. (2019), as shown in Table 1. The specimens were deposited in the Colección de Artrópodos, Laboratorio de Entomología Sistemática, Universidad del Valle de Guatemala, and are being cataloged (Fig. 3c). Distribution: Southwestern Mexico (Warren et al. 2017).

• Repensflorus (Godman, 1900). Reserva Heloderma, Cabañas, Zacapa, GUATEMALA. One specimen: 23-10-2018, J800. Collected by Jiichiro Yoshimoto. Identified by Andrew D. Warren. The specimen was deposited as above and is being cataloged (Fig. 3d). Distribution: Eastern and Western Mexico, Belize, and Nicaragua (Warren et al. 2017).

• Niconiadesnikko Hayward, 1948. Los Cerritos, Salamá, Baja Verapaz, GUATEMALA. One specimen: 16-11-2020, J1024. Collected and identified by Jiichiro Yoshimoto. The specimen was deposited as above and is being cataloged (Fig. 3e). Distribution: Eastern Mexico to Ecuador, Southern Brazil, and Paraguay (Warren et al. 2017).

Figure 3.

One species of hairstreak (Lycaenidae: Theclinae) aChalybshassan (Stoll, 1790), one species of skipperling (Hesperiidae: Heteropterinae) bPirunaaea (Dyar, 1912), and three species of grass-skippers (Hesperiidae: Hesperiinae) cAmblyscirteselissaelissa Godman, 1900 dRepensflorus (Godman, 1900), and eNiconiadesnikko Hayward, 1948. The three grass-skipper species were newly recorded for Guatemala. Dorsal and ventral views, respectively, are shown at the left and right in each photograph.

Eighty-six species were shared between Los Cerritos and Heloderma Reserve (Table 2), which amounts to 51.8% and 61.9% of the species sampled at each site (the Jaccard dissimilarity index is 0.606). At both sites, species richness of Nymphalidae and Hesperiidae was greater than that of the other four families, although family-level species richness differed greatly between the sites (Table 2; Fig. 4). In particular, the proportion of Lycaenidae was much higher at Los Cerritos (18.7%) than at Heloderma Reserve (8.6%), which was mainly due to differences in the subfamily Theclinae (26 and 7 species, respectively: Appendix 1).

Table 2.

Species richness for six families at Los Cerritos and Heloderma Reserve, and comparisons of species composition at the family level between the sites, based on the number of shared species and the Jaccard dissimilarity index.

Family	Total No. species		No. shared species	Jaccard index
	Los Cerritos	Heloderma Reserve
Papilionidae	7	4	2	0.778
Pieridae	16	17	13	0.350
Lycaenidae	31	12	7	0.806
Riodinidae	10	9	3	0.813
Nymphalidae	57	46	37	0.439
Hesperiidae	45	51*	24	0.662*

*The data for Bolla sp. were included in the species count but excluded from the Jaccard index analysis (see the footnote 7 of the Appendix 1 for its rationale).

Figure 4.

Proportion of species richness at the family level at Los Cerritos and Heloderma Reserve.

Family-level species composition also differed between the sites, and the magnitude of this difference varied among the six families (Table 2). The dissimilarity indices for Riodinidae, Lycaenidae, and Papilionidae were considerably larger, indicating that species composition differed more greatly between the sites in these families. Pieridae and Nymphalidae, by contrast, had smaller indices with many shared species, demonstrating that their species composition was relatively similar between the sites.

Ninety-three species (42.7%) occurred in both dry and rainy seasons, whereas 103 (47.2%) appeared only in the rainy season and 22 species (10.1%) only in the dry season. The most frequently recorded species was Euremadairaeugenia (Wallengren, 1860) (Pieridae: Coliadinae), which was collected or observed throughout the year (Appendix 1). The second most frequently recorded species (in 11 months) were Kricogonialyside (Godart, 1819) (Coliadinae) and Hamadryasglauconomeglauconome (H. Bates, 1864) (Nymphalidae: Biblidinae), followed by Pyrisitiaproterpia (Fabricius, 1775) (Coliadinae: in ten months), Phoebissennaemarcellina (Cramer, 1777) (Coliadinae), Mestraamymone (Ménétriés, 1857) (Biblidinae), and Urbanusdorantesdorantes (Stoll, 1790) (Hesperiidae: Eudaminae: all in nine months).

Discussion

A total of 218 species were recorded at the two dry forest sites during our 10-year field surveys, which confirms the relatively high lepidopteran diversity of Guatemalan seasonally dry forests for the small areas that comprise the study sites (<70 ha each). The estimated species richness suggests that nearly a quarter of the species inhabiting each site have yet to been recorded. The number of the additional species yielded in the subsequent surveys was more than twice greater at Heloderma Reserve than at Los Cerritos. The proportion of newly recorded species was much higher in Lycaenidae and Riodinidae; seven lycaenid species were added to the list for Los Cerritos, and seven lycaenid and four riodinid species were added to that of Heloderma Reserve, which nearly doubled the species richness of each family at this site (six lycaenid and five riodinid species in Yoshimoto et al. 2019). Among these species, the record of Chalybshassan (Stoll, 1790) at Heloderma Reserve is highly important (Fig. 3a), as this species had not been reported for more than 100 years in Guatemala before we collected four individuals at Los Cerritos in 2011 and 2012 (Yoshimoto and Salinas-Gutiérrez 2015). These results highlight the importance of continuing butterfly surveys at both sites to create more exhaustive inventories, especially on small and taxonomically difficult taxa such as Lycaenidae and Riodinidae. Moreover, it is important to conduct research in other dry regions (e.g., the Nentón Valley in northwestern Guatemala) and to make quantitative among-site comparisons of species richness and composition as well. All these studies will contribute to a comprehensive understanding of Neotropical butterfly fauna and distribution, and would serve as a scientific baseline for biodiversity conservation in Guatemalan dry regions.

More than half of the species sampled at each site were shared between the sites, suggesting that species composition is partially and moderately similar between Los Cerritos and Heloderma Reserve. Importantly, between-site similarity greatly differed among the six families. Higher similarity in Pieridae (especially in Coliadinae) would likely be associated with the distribution and abundance of their host plants, considering that coliadine larvae mostly feed on fabaceous plants such as Senna (e.g., DeVries 1987) and that these plants appear to be abundant at both sites.

In Lycaenidae and Riodinidae, species composition largely differed between the sites; in particular, Theclinae had considerable differences in species richness and composition (Appendix 1). In addition, most of these thecline species tended to be highly seasonal, as 25 out of 30 species were sampled only in the rainy season. In contrast to their marked seasonal pattern, Strymonmegarus (Godart, [1824]) and S.rufofusca (Hewitson, 1877) occurred frequently also in the dry period at Heloderma Reserve; three and five individuals of each species were collected in both December and January at this site (Appendix 1). It should also be mentioned that Hechtiaguatemalensis Mez (Bromeliaceae), one of the dominant bromeliad species at Heloderma Reserve (Fig. 1b), may be a possible foodplant for S.megarus at this site, as the larvae of this species are known to feed on bromeliads (Robbins 2010). Examination of abundance and distribution of host- and nectar-plants, as well as of larval and adult feeding behavior in relation to their phenology, would be an initial step to elucidate the bionomics of these species. Such surveys may also identify factors underlying the regional similarity and dissimilarity in the butterfly fauna.

We recorded Amblyscirteselissaelissa Godman, 1900, Repensflorus (Godman, 1900), and Niconiadesnikko Hayward, 1948 (Hesperiidae: Hesperiinae) for the first time in Guatemala (Fig. 3c, d, e). Austin et al. (1998) listed A.e.elissa and N.nikko as species with a potential distribution in Guatemala. Repensflorus could have been included in this category as well, as it is known to be distributed in the adjacent countries (Mexico, Belize, and Nicaragua; Warren et al. 2017). These results indicate that there still exists a gap in our knowledge of geographic distribution of Neotropical skipper species, again emphasizing the importance of more intensive research in Guatemala to bridge this gap.

Four individuals of Pirunaaea (Dyar, 1912) (two in the previous survey and two in the subsequent one: Figs 2g, 3b) were collected at Heloderma Reserve. This is an interesting result, since most species in this genus are distributed in humid areas at higher elevation (1000–2700 m; Warren and González-Cota 1998). As Yoshimoto et al. (2019) pointed out, the wing pattern of these individuals is somewhat different from Mexican P.a.aea (Dyar, 1912), implying that Pirunacingosombra Evans, 1955, described from Guatemala and currently considered a synonym of P.a.aea, may be a valid subspecies-level taxon. At present, this is difficult to determine, as very few specimens of this species have been sampled in Guatemala (Barrios et al. 2006).

Appendix 1

Table A1.

Butterfly species observed in 2011–2021 at two dry forests in Guatemala: Los Cerritos Municipal Park and Heloderma Natural Reserve, based on our previous studies (Yoshimoto et al. 2018, 2019) and on subsequent field surveys (July 2014 to August 2021 at Los Cerritos and October 2017 to November 2021 at Heloderma Reserve). Species and months in bold indicate the data newly obtained in the subsequent surveys. Year information is also shown with sampling months, when necessary. Nomenclature follows Warren et al. (2017).

Family		Months when observed
Subfamily		Los Cerritos	Heloderma Reserve
Species and subspecies
Papilionidae
Papilioninae
1	Neographiumepidausepidaus (E. Doubleday, 1846) PH, A	Apr, May, Jun, Jul, Aug, Nov	–
2	Neographiumphilolausphilolaus (Boisduval, 1836) PH, A	Jun, Sep	Mar, Apr, May, Jun
3	Battuspolydamaspolydamas (Linnaeus, 1758) PH, A	Mar, Jul, Aug, Sep, Dec	–
4	Paridesphotinus (E. Doubleday, 1844)	–	Sep
5	Heraclideserostratuserostratus (Westwood, 1847) A, Y	May, Oct	–
6	Heraclidesthoasautocles (Rothschild & Jordan, 1906) PH, A	Feb, Mar, Apr, Jun, Aug, Nov	–
7	Heraclidesornythionornythion (Boisduval, 1836) PH	–	May, Jun
8	Heraclidesrumiko Shiraiwa & Grishin, 2014 MI, PH	Jul	Oct’16MI, Dec
9	Papiliopolyxenesasterius Stoll, 1782 PH	Mar, Apr, May, Nov	–
Pieridae
Coliadinae
10	Kricogonialyside (Godart, 1819) PH, A	Mar	Jan, Feb, Mar, Apr, May, Jun, Jul, Aug, Sep, Nov, Dec
11	Euremadairaeugenia (Wallengren, 1860) PH, A, Y	Jan, Feb, Aug, Nov	Jan, Mar, Apr, May, Jun, Jul, Aug, Sep, Oct, Dec
12	Euremaboisduvaliana (C. Felder & R. Felder, 1865) PH, A	Feb, Oct, Nov	Jun, Jul, Aug, Sep, Oct, Nov
13	Abaeisnicippe (Cramer, 1779) MI, PH	Nov	Jul’16MI
14	Pyrisitiaproterpia (Fabricius, 1775) PH, A	May, Jun, Jul, Oct, Dec	Feb, Apr May, Jun, Jul, Aug, Sep, Oct, Nov
15	Pyrisitiadinawestwoodi (Boisduval, 1836) PH, A	–	Jan, Feb, Jun, Oct, Nov, Dec
16	Pyrisitianisenelphe (R. Felder, 1869) PH, A, Y	Jun, Jul, Aug, Nov	Jun, Jul, Aug, Sep, Oct, Nov, Dec
17	Zerenecesoniacesonia (Stoll, 1790) PH, Y	Jun, Aug	Jun, Jul
18	Anteosmaerula (Fabricius, 1775) PH, A	Jun, Sep, Nov	May, Jun, Jul, Aug, Sep, Oct
19	Anteosclorinde (Godart, [1824]) PH, A	Apr, Jun	Jun, Aug
20	Phoebissennaemarcellina (Cramer, 1777) PH, A	Feb, Mar, Apr, Jun, Jul, Nov	Mar, May, Jun, Jul, Aug, Sep
21	Phoebisphileaphilea (Linnaeus, 1763) A, Y	May	Jul
22	Phoebisargante ssp. A	May	Jul
23	Aphrissastatirastatira (Cramer, 1777) A	Oct	–
Pierinae
24	Hesperochariscroceacrocea H. Bates, 1866	Mar, Aug	–
25	Asciamonustemonuste (Linnaeus, 1764) PH, A, Y	Feb, Jun	Jun
26	Ganyrajosephinajosepha (Salvin & Godman, 1868) PH, A	–	Jan, Oct
27	Leptophobiaaripaelodia (Boisduval, 1836) Y	Jan	–
28	Itaballiademophilecentralis Joicey & Talbot, 1928	–	Jan
29	Glutophrissadrusillatenuis (Lamas, 1981) A	–	Jun, Aug
Lycaenidae
Theclinae
30	Evenusregalis (Cramer, 1775) A	Sep	–
31	Atlidesgaumeri (Godman 1901)	Aug	–
32	Atlidescarpasia (Hewitson, 1868) A	Aug	–
33	Rekoazebina (Hewitson, 1869)	Jun, Sep	–
34	Rekoastagira (Hewitson, 1867) A	Aug	–
35	Arawacussito (Boisduval, 1836) A, Y	Aug	–
36	Arawacusjada (Hewitson, 1867) A	Jul	–
37	Kolanalyde (Godman & Salvin, 1887) A	Sep	–
38	Chlorostrymonsimaethissarita (Skinner, 1895) A	Nov	–
39	Cyanophrysherodotus (Fabricius, 1793) A	Aug	–
40	Cyanophrysmiserabilis (Clench, 1946)	–	Oct
41	Electrostrymonhugon (Godart, [1824])	–	Jul
42	Kisutamsyllis (Godman & Salvin, 1887) A	–	Oct
43	Calycopisclarina (Hewitson, 1874)	Jun	–
44	Calycopisisobeon (A. Butler & H. Druce, 1872)	Aug, Sep	–
45	Strymonmelinusfranki W. D. Field, 1938	Aug, Sep	–
46	Strymonrufofusca (Hewitson, 1877) PH	Jul, Nov	Jan, Aug, Oct, Nov, Dec
47	Strymonbebrycia (Hewitson, 1868) PH	Jun, Aug	–
48	Strymonyojoa (Reakirt, [1867]) A	Jul	–
49	Strymoncestri (Reakirt, [1867]) A	Aug	–
50	Strymonbazochiibazochii (Godart, [1824]) A	Jul	–
51	Strymonistapaistapa (Reakirt, [1867])	Aug, Nov	–
52	Strymonmegarus (Godart, [1824]) MI, PH	–	Jan, Jul, Oct’16MI, Dec
53	Strymonziba (Hewitson, 1868)	Jul	–
54	Ministrymonazia (Hewitson, 1873) A	Jun	Jul
55	Ostrinoteskeila (Hewitson, 1869) A, Y	Aug	–
56	Panthiadesbitias (Cramer, 1777) A	Jun	–
57	Michaelushecate (Godman & Salvin, 1887)	Sep	–
58	Eroragabina (Godman & Salvin, 1887)	May, Jun, Aug, Oct	–
59	Chalybshassan (Stoll, 1790)	Aug, Sep, Nov	Jul
Polyommatinae
60	Celastrinaechogozora (Boisduval, 1870) Y	Nov	–
61	Leptotescassiuscassidula (Boisduval, 1870) PH, A, Y	Jun, Dec	Sep, Oct
62	Cupidocomyntastexana (F. Chermock, 1945) PH, A, Y	Sep, Nov	Oct, Nov, Dec
63	Hemiargusceraunusastenidas (Lucas, 1857) A	Mar, Jul, Nov	Feb, Jun, Dec
64	Hemiargushannohanno (Stoll, 1790) PH, A	–	Jul, Aug, Sep, Oct
65	Echinargusisola (Reakirt, [1867])	Feb, Dec	Jun, Dec
Riodinidae
Riodininae
66	Rhetusarciuscastigatus Stichel, 1909 A	Sep	–
67	Calephelis spp. PH, 1	Jan, May, Jul, Aug, Oct, Nov, Dec	Jan, Jul, Aug, Sep, Oct, Nov, Dec
68	Lasaiasulasula Staudinger, 1888 PH	–	Jun, Oct
69	Lasaiamariamaria Clench, 1972	–	Jun, Jul, Oct
70	Melanispixepixe (Boisduval, 1836) A	Feb, Sep, Nov, Dec	–
71	Anteroscarausiuscarausius Westwood, 1851 A	Aug, Nov	Sep, Nov
72	Calydnasturnula (Geyer, 1837) PH	Aug, Sep, Oct	–
73	Emesismandana furor A. Butler & H. Druce, 1872 A	Aug	–
74	Emesistenedia C. Felder & R. Felder, 1861 A, Y, 2	Jul	–
75	Emesislupinalupina Godman & Salvin, 1886 2	Oct	–
76	Curvieemesia (Hewitson, 1867) PH, A, 3	–	Jun, Oct
77	Thisbelycorias (Hewitson, [1853]) A	Jun, Jul, Oct, Nov	–
78	Judithacaucana (Stichel, 1911)	–	Oct
79	Synargismycone (Hewitson, 1865) A	Mar, Jun, Jul	Jul
80	Hypophyllazeurippa Boisduval, 1836	–	Jan
81	Theopevirgilius (Fabricius, 1793) A	–	Feb, Nov
Nymphalidae
Libytheinae
82	Libytheanacarinentamexicana Michener, 1943 PH, A, Y	Jul	Jun, Aug, Sep
Danainae
83	Lycoreahaliaatergatis E. Doubleday [1847] * PH, A, Y	–	May, Sep
84	Danauseresimusmontezuma Talbot, 1943 PH, A	Aug, Nov, Dec	Jun, Aug, Sep
85	Danausgilippusthersippus (H. Bates, 1863) A	–	Mar
86	Mechanitislysimniautemaia Reakirt, 1866 PH, A, Y	–	May, Aug, Oct
87	Mechanitispolymnialycidice H. Bates, 1864 PH, A, Y	Sep	Sep, Oct
88	Dircennaklugiiklugii (Geyer, 1837) Y	Sep, Oct, Nov	–
Heliconiinae
89	Agraulisvanillaeincarnata (N. Riley, 1926) PH, A	Jun, Jul, Nov	Aug
90	Dionemonetapoeyii A. Butler, 1873 Y	Jun, Nov	–
91	Dionejunohuascuma (Reakirt, 1866) PH, A	Feb, Mar, Dec	Jul
92	Dryasiuliamoderata (N. Riley, 1926) PH, A, Y	Aug	Aug, Sep, Oct
93	Eueidesisabellaeva (Fabricius, 1793) A	Nov	–
94	Heliconiuscharithoniavazquezae W. Comstock & F. Brown, 1950 A, Y	Jul	Jul, Oct
95	Euptoietahegesiameridiania Stichel, 1938 PH, A, Y	Jun, Jul, Sep	Jun, Jul
Limenitidinae
96	Adelphaparoecaparoeca (H. Bates, 1864) Y	Oct, Nov	–
97	Adelphaiphicleolaiphicleola (H. Bates, 1864) PH	Aug	Jun, Jul, Sep, Oct
98	Adelphamelanthe (H. Bates, 1864) A	Aug, Sep	–
Biblidinae
99	Biblishyperiaaganisa Boisduval, 1836 PH, A	Jul, Dec	Sep
100	Mestraamymone (Ménétriés, 1857) A	May, Jun, Jul	Mar, Jul, Aug, Sep, Oct, Nov, Dec
101	Catonephelemexicana Jenkins & R.G. Maza, 1985 A	Sep, Oct, Nov	–
102	Eunicamonima (Stoll, 1782) PH	Jun, Aug	Mar, Jun, Jul, Aug, Oct, Dec
103	Eunicatatilatatila (Herrich-Schäffer, [1855]) A	Jun	–
104	Hamadryasatlantisatlantis (H. Bates, 1864) PH	Sep	Jun, Jul, Nov
105	Hamadryasfebruaferentina (Godart, [1824]) PH, A	Apr, May, Jun, Jul, Nov, Dec	Feb, Jul, Oct
106	Hamadryasglauconomeglauconome (H. Bates, 1864) PH	Jan, Jul, Oct, Nov, Dec	Jan, Feb, Mar, May, Jun, Jul, Aug, Sep, Oct
107	Hamadryasguatemalenaguatemalena (H. Bates, 1864) PH, A	May, Jun, Jul	Jul
108	Bolboneurasylphissylphis (H. Bates, 1864) PH	Jul, Aug, Sep	Mar, Jun, Jul, Sep, Oct
109	Epiphileadrastaadrasta Hewitson, 1861 Y	Aug, Oct	–
110	Temenislaothoehondurensis Fruhstorfer, 1907 A	–	Oct
111	Dynaminedyonis Geyer, 1837 A	Jul, Aug, Oct, Nov, Dec	–
112	Dynaminepostvertamexicana R.F. d’Almeida, 1952 PH, A	Jun, Jul, Oct, Nov	Sep, Oct
113	Dynaminetheseus (C. Felder & R. Felder, 1861) A	Aug, Sep, Oct	–
114	Diaethriaastalaastala (Guérin-Méneville, [1844]) A, Y	May, Jul, Oct, Nov	Oct
Cyrestinae
115	Marpesiapetreus ssp. A, Y	Jul, Sep	Jun
Nymphalinae
116	Historisodiusdious Lamas, 1995 PH, A	Jun, Jul, Aug, Sep	Jul
117	Smyrnablomfildiadatis Fruhstorfer, 1908 PH, A, Y	Jul	May, Nov
118	Anartiafatimafatima (Fabricius, 1793) PH, A, Y	May	Jun, Sep, Oct
119	Siproetaepaphusepaphus (Latreille, [1813]) A, Y	Sep	Sep
120	Siproetastelenesbiplagiata (Fruhstorfer, 1907) PH, A, Y	Jul, Sep	Jun, Aug, Sep, Oct
121	Junoniaevarete (Cramer, 1779) PH, A	Jun, Jul, Aug	Jan, Jul, Oct
122	Chlosynejanaisjanais (Drury, 1782) A	Jun, Aug	–
123	Chlosyneerodyleerodyle (H. Bates, 1864) MI	Jul’12MI, Jul’19, Oct’11MI, Oct’17	–
124	Chlosynerositarosita A. Hall, 1924 MI	Sep’11MI	Jun, Jul, Sep
125	Chlosynetheonatheona (Ménétriés, 1855) PH	Apr, Jun, Sep	Jun, Jul, Aug, Sep
126	Chlosynelacinialacinia (Geyer, 1837) PH, A, Y	Mar, Jun, Jul, Aug, Nov	Jun, Jul, Aug
127	Chlosynemelanarge (H. Bates, 1864) PH	–	Aug, Sep, Oct
128	Microtiaelvahorni Rebel, 1906 PH	Jun, Jul, Aug, Nov	Jun, Jul, Aug, Sep, Oct, Nov
129	Anthanassatulcis (H. Bates, 1864) MI, A	May’11MI	Jun, Sep, Dec
130	Anthanassaptolycaptolyca (H. Bates, 1864) Y	Aug	Dec
131	Tegosaguatemalena (H. Bates, 1864) A	Feb, Nov	–
Charaxinae
132	Zaretisellops (Ménétriés, 1855) A	Jul, Sep, Nov	–
133	Anaeaaidea (Guérin-Méneville, [1844]) PH, A	May, Nov	Jun, Aug, Oct, Nov
134	Fountaineaglyceriumglycerium (E. Doubleday, [1849]) PH	Aug, Sep, Oct, Nov	–
135	Archaeopreponademophoncentralis (Fruhstorfer, 1905) * PH, A	–	Jul
Satyrinae
136	Morphohelenor ssp.* A	–	Sep
137	Caligotelamoniusmemnon (C. Felder & R. Felder, 1867) * PH, A	–	Oct
138	Manatariahercynamaculata (Hopffer, 1874) A, Y	–	Jun
139	Cissiasimilis (A. Butler, 1867) MI, PH, A	May’12MI, Jun’12MI, Oct, Nov	Jan, Feb, Apr, May, Jun, Oct, Nov, Dec
140	Cissiathemis (A. Butler, 1867) MI, PH	Jul, Aug’11MI	Feb, Jun, Jul, Aug’16MI, Oct’16MI, Dec
141	Cyllopsisgemmafreemani (D. Stallings & J. Turner, 1947)	Sep, Nov	–
142	Cyllopsishedemannihedemanni R. Felder, 1869 Y	Feb	–
143	Cyllopsishilaria (Godman, 1901)	Sep, Nov	–
144	Cyllopsispephredo (Godman, 1901) Y	Jun, Nov	–
145	Euptychiafetna A. Butler, 1870	Aug, Sep	–
146	Hermeuptychiahermes (Fabricius, 1775) A, Y, 4	Jul	Jan, Feb, Sep, Oct
147	Taygetisthamyra (Cramer, 1779) PH	Nov	Jun, Oct
Hesperiidae
Eudaminae
148	Phocidespolybiuslilea (Reakirt, [1867]) A	Nov	–
149	Phocidesuraniaurania (Westwood, 1852)	Aug	–
150	Proteidesmercuriusmercurius (Fabricius, 1787) PH, A	Jun	Jun, Sep
151	Epargyreusexadeuscruza Evans, 1952 A, Y	Feb, Mar, Apr, Jun, Jul	Aug
152	Polygonusleoarizonensis (Skinner, 1911)	Jul	Jun, Jul, Aug, Sep, Oct
153	Chioidesalbofasciatus (Hewitson, 1867) A	Jun	–
154	Chioideszilpa (A. Butler, 1872) A	Jan, Mar	–
155	Typhedanusundulatus (Hewitson, 1867) A	Feb, Mar, May	–
156	Typhedanusampyx (Godman & Salvin, 1893) A	–	Oct
157	Polythrixasine (Hewitson, 1867) PH, A, 5	–	Jan, Dec
158	Polythrixoctomaculata (Sepp, [1844]) A	–	May
159	Cephiseaelius (Plötz, 1880)	–	Oct
160	Codatractusalcaeusalcaeus (Hewitson, 1867)	Mar, May	–
161	Codatractusmelon (Godman & Salvin, 1893)	–	Jun, Jul
162	Urbanusviterboana (Ehrmann, 1907) MI, A, Y	Sep, Nov’11MI	Sep, Oct
163	Urbanusesmeraldus (A. Butler, 1877) A, Y	Aug, Sep	Jun
164	Urbanusdorantesdorantes (Stoll, 1790) PH, A	May, Jul, Aug, Dec	Apr, Jun, Jul, Aug, Sep, Oct, Nov
165	Urbanusprocne (Plötz, 1881) PH, A, Y	May, Jul, Nov	–
166	Urbanusdoryssusdoryssus (Swainson, 1831) A	Jul	–
167	Astraptesfulgeratorazul (Reakirt, [1867]) A, Y, 6	Oct, Nov	–
168	Astraptesalectorhopfferi (Plötz, 1881) A	Sep	Jan
169	Astraptesanaphusannetta Evans, 1952 PH, A, Y	Jun, Oct	Jul
170	Achalarustoxeus (Plötz, 1882) A	–	Mar, Apr, Oct
171	Achalarusalbociliatusalbociliatus (Mabille, 1877) A	Feb, Mar	Oct, Nov
172	Cabarespotrillopotrillo (Lucas, 1857) A, Y	–	Jun, Jul, Aug, Oct, Nov
173	Cogiacajetaeluina Godman & Salvin, 1894	May, Jun	–
Pyrginae
174	Mysoriaaffinis (Herrich-Schäffer, 1869)	–	Oct
175	Celaenorrhinusfritzgaertneri (Bailey, 1880) PH	Feb, Aug	Mar, Jun
176	Noctuanastator (Godman, 1899) PH, A, Y	Feb, Mar, May, Jul, Sep	–
177	Bollaevippe (Godman & Salvin, 1896)	Mar	–
(177)	Bolla sp. 7	–	Oct 7
178	Staphylusascalaphus (Staudinger, 1876) Y	May	Jan, Sep
179	Staphylusazteca (Scudder, 1872)	–	Feb, Aug, Nov
180	Gorgythionvox Evans, 1953 A, Y	–	Jun, Jul, Aug, Sep, Oct
181	Mylonsalvia Evans, 1953	Nov	–
182	Mylonpelopidas (Fabricius, 1793) A	–	May, Jun
183	Graisstigmaticusstigmaticus (Mabille, 1883) PH	–	Jan, Jun, Jul
184	Timocharestrifasciatatrifasciata (Hewitson, 1868) A	–	Jan
185	Chiomarageorginageorgina (Reakirt, 1868)	Jan, Jul, Sep, Nov	Aug
186	Erynnisfuneralis (Scudder & Burgess, 1870)	Aug	Jun
187	Eantistamenund (W. H. Edwards, 1871) PH	Feb, Jul, Aug, Nov, Dec	–
188	Atarnessallei (C. Felder & R. Felder, 1867) A	Jul	–
189	Carrhenesfuscescensfuscescens (Mabille, 1891)	–	Jun
190	Antigonuserosus (Hübner, [1812]) PH, A	Mar, Oct	Feb, Mar, Jun, Aug, Oct
191	Antigonuscorrosus Mabille, 1878 A, Y	–	Sep
192	Zopyrionsandace Godman & Salvin, 1896	Mar, Aug	Jan, Apr, May, Jun, Sep, Dec
193	Pyrgusoileus (Linnaeus, 1767) PH, A, Y	Apr	Jul, Aug, Sep, Oct, Dec
194	Pyrgusorcus (Stoll, 1780)	–	Jun
195	Heliopyrgusdomicelladomicella (Erichson, [1849])	Sep	Sep, Oct, Nov
196	Heliopeteslavianalaviana (Hewitson, 1868) A	–	Feb
197	Heliopetesmacairamacaira (Reakirt, [1867]) MI, A	Jul’12MI, Nov	Oct
198	Heliopetesalana (Reakirt, 1868) A, Y	Jul	–
Heteropterinae
199	Pirunaaea (Dyar, 1912) PH	–	Jun 8, Jul, Sep 8, Oct
Hesperiinae
200	Pericharesadela (Hewitson, 1867) A	Aug, Sep	Oct
201	Copaeodesaurantiaca (Hewitson, 1868) MI	Mar’11MI	–
202	Panoquinalucas (Fabricius, 1793) A		Jan
203	Zenisjebushemizona (Dyar, 1918)	Jan	–
204	Synapteshiva Evans, 1955	Aug	Jun, Dec
205	Synaptesyraces (Godman, 1901)	–	Sep
206	Callimormussaturnus (Herrich-Schäffer, 1869) A	–	Oct
207	Amblyscirteselissaelissa Godman, 1900 NR , MI	–	Jun, Aug’16MI, Sep’16MI
208	Amblyscirtestoltecatolteca Scudder, 1872 A	Jun, Jul	May
209	Methionopsisina (Plötz, 1882) A	–	Jan, Oct, Nov
210	Repensflorus (Godman, 1900) NR	–	Oct
211	Cymaenestrebius (Mabille, 1891) MI, A	Aug	Sep’16MI
212	Leremaliris Evans, 1955	–	Jul, Sep
213	Niconiadesnikko Hayward, 1948 NR	Nov	–
214	Vettiusfantasos (Cramer, 1780) A, Y	Aug, Sep, Oct	Jan, Oct
215	Hylephilaphyleusphyleus (Drury, 1773) A	Aug	–
216	Politesvibexpraeceps (Scudder, 1872) A	Mar, Apr	–
217	Pompeiuspompeius (Latreille, [1824]) A, Y	–	Jun
218	Atrytonopsisovinia (Hewitson, 1866) PH	Jan, Feb, Mar, Apr, Oct	Oct, Dec

^PH Species photographed.

^A Species reported at Tikal in northern Guatemala by Austin et al. (1996).

^Y Species reported at Parque Cayalá in Guatemala City by Yoshimoto et al. (2021).

^* Specimens not collected (recorded only by direct observation or photographs).

^MI Species misidentified in Yoshimoto et al. (2018 or 2019); their previous identification results are shown in Table 1.

¹ Listed at the genus level because of the difficulty of species-level identification derived from the confused taxonomic state of this genus.

² These two species, together with Emesistegula and E.toltec, can be treated as a species complex. These taxa require further study, according to Trujano-Ortega et al. (2021).

³ The genus name, treated as Emesis in Yoshimoto et al. (2019), was modified according to Zhang et al. (2019).

⁴ The name “hermes” is correctly applied to a South American species, according to Cong and Grishin (2014). Thus, the individuals collected might include multiple species, none of which are true hermes.

⁵ One sample which was photographed at Heloderma Reserve in October 2016 and was identified as this species in Yoshimoto et al. (2019) was excluded, as we found that it might have been of another species of this genus, which is unable to de determined because of the lack of a specimen.

⁶ This is a species complex, which includes several species in Costa Rica (Hebert et al. 2004; Brower, 2006, 2010). Thus, the individuals collected might also be of multiple species.

⁷ These data were included in the species count and Chao II analyses for Heloderma Reserve, since at this site none of the identified species of this genus had been recorded. In contrast, these data have been excluded from the Chao II analysis for all data (pooled across the sites) and from the Jaccard index analyses, as this individual might be of Bollaevippe (Godman & Salvin, 1896), which is unable to be examined because of the heavily damaged specimen of Bolla sp.

⁸ Identified to genus (Piruna sp.1) and incorrectly listed as Hesperiinae in Yoshimoto et al. (2019).

^NR New record for Guatemala.

Citation

Yoshimoto J, Salinas-Gutiérrez JL, Barrios M, Warren AD (2022) An updated list of butterflies (Lepidoptera, Papilionoidea) of two Guatemalan seasonally dry forests. ZooKeys 1118: 21–38. https://doi.org/10.3897/zookeys.1118.85810