A Preliminary Assessment of the Butterfly Fauna of El Edén Ecological Reserve: Species Richness and Habitat Preferences

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Chapter 13

A Preliminary Assessment of the Butterfly

Fauna of El Edén Ecological Reserve:

Species Richness and Habitat Preferences

Jorge L. León-Cortés Robert W. Jones Olga L. Gómez-Nucamendi

INTRODUCTION

The Yucatán Peninsula is a district subregion—not only biologically, but also geologically and climatically. It includes parts of southern Mexico, Belize, and Guatemala. The peninsula was once covered with forest, but at present, this forest does not have the same characteristics before human settlements (Welling 1966). Data from the Yucatán Peninsula indicate that the late Pleistocene and early Holocene epochs were dry (Metcalfe et al. 2000). In many cases, lake levels were strongly influenced by lower glacial sea levels and groundwater levels. Lake levels started to rise after 9000 yr. B.P. Moist conditions, possibly wetter than present, were established by about 6800 yr. B.P., but this was followed by a dry period between about 6000 and 5000 yr. B.P. (Metcalfe et al. 2000).

The late Holocene was marked by strong variations in water balance, and the driest period of the Holocene was recorded at many sites sometime between 1500 and 900 yr. B.P. This very dry period is of considerable interest as it corresponds with the time of the collapse of the lowland Maya. Wetter conditions were then reestablished. Many sites in this region also provide records of human disturbance throughout the late Holocene (Metcalfe et al. 2000), and sometimes deliberate modification of the natural environment. The authors wish to thank Marco Lazcano-Barrero for his help in field collections and Manuel Girón-Intzin for his help in mounting material. This is the paper number 4 of the “Dinámica de poblaciones y comunidades de insectos” research group at ECOSUR.

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Leyden et al. (1998) have suggested that pollen records show a very strong human disturbance signal. Hence, both historical human and natural destructive forces have been influential in greatly decimating the natural forest cover along with the associated fauna in the Yucatán Peninsula. Because the area of primary forest and other semi-natural habitats have suffered dramatic changes in much of the peninsula over the course of its history, it is likely that this has also precipitated changes in animal species composition and diversity. Compared to other insect groups in the Neotropics, butterflies have become such well-studied species that they have been used as a representative insect group for many regions. Unlike most insects in tropical forests, butterflies are taxonomically tractable, occur at moderate levels of species richness, and are easy to sample (Gilbert 1984; Brown 1991; New 1991; Sparrow et al. 1994; Lewis, Wilson, and Harper 1998). These characteristics have made tropical butterflies a popular group for investigations into the effects of small-scale disturbances (Bowman et al. 1990; Spitzer et al. 1993), selective logging (Hill et al. 1995), and forest fragmentation (Daily and Ehrlich 1995). Several studies have revealed consistent differences in the responses of butterflies with large and small geographical ranges to human disturbance of tropical forest habitats (Thomas 1991). Whereas widespread butterfly species typically become more abundant in human-modified tropical forests, endemic or geographically restricted species decline (Thomas 1991; Spitzer et al. 1993; Hill et al. 1995).

The aim of this paper was to investigate the patterns of species composition and richness, species habitat preferences, and geographic ranges of butterflies from a specific region in the Yucatán Peninsula—namely, the El Edén Ecological Reserve in the northern part of the peninsula. Whether current species composition patterns could reflect historical (and recent) recurrent habitat modifications in the peninsula is the focus of investigation. Can current butterfly species composition tell us something about past and present habitat disturbance in the Yucatán Peninsula? Are geographically widespread butterfly species more common in Yucatán landscapes than elsewhere in Mexico and Central America?

METHODS

Study site

The El Edén Ecological Reserve (hereafter, El Edén) is located in the Mexican State of Quintana Roo, in the northern portion of the Yucatán Peninsula (Figure 13.1). El Edén occupies an area of approximately 1,560 hectares (ha). Medium-height tropical subperennial forest is the main

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El Edén

100 0 100 200 Miles

N

FIGURE 13.1. The Mexican State of Quintana Roo is shown in black in southern Mexico (inset), and the circle shows the location of El Edén Ecological Reserve. The bar scale applies to Quintana Roo.

vegetation type in the region. A variety of habitat types are also present at El Edén, namely secondary forest, inundated mixed forest (“tintal”), savanna, forest gaps, hedges, and forest borders. All major habitat types recognized in the vicinity of El Edén (see as follows).

Sampling protocol and fauna size

During late summer and early autumn 1995, mostly Papilionidae, Pieridae, and Nymphalidae butterflies were caught using direct netting and traps of vertical cylindrical netting (Van Someren-Rydon traps). For other families (e.g., Hesperiidae and Lycaenidae), limited sampling efforts were carried out. Van Someren-Rydon traps were baited with decaying fruits and hung above trees near lightgaps, forest margins, and moist trails (DeVries 1987; Raguso and Llorente-Bousquets 1991). Traps were inspected daily for captured specimens. Sampling efforts began at 9:00 a.m. and ended at 6:00 p.m. Sampling was carried out only when conditions were ideal to observe active butterflies (Pollard and Yates 1993). All captured individual butterflies were mounted, labeled as to locality, and deposited at the Entomological Collection of El Colegio de la Frontera Sur (ECOSUR). Specimens were identified using a variety of sources, including De la Maza-Ramírez (1987);

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De la Maza-Elvira and De la Maza-Elvira (1993); and DeVries (1987). The reference entomological collection at ECOSUR was also used.

As a way of checking the accuracy of our sampling effort at El Edén, the cumulative number of species versus the number of individuals sampled (see Figure 13.2) was plotted. The species accumulation curve is expected to reach a maximum value when all species at a site have been collected. The faunistic data was fitted to a Michaelis-Menten function type (known in the Entomological literature as Clench’s function), which has provided accurate estimates for butterfly faunas in a variety of tropical sites (Clench 1979; Soberón and Llorente 1993; Colwell and Coddington 1994; Raguso and Bousquets 1991; León-Cortés, Soberón-Mainero, and Llorente-Bousquets 1998). Clench’s equation is expressed as follows:

S(t) = __a*t

(1 + b*t) (1) In Equation (1), S(t) is the expected size of the list, a is the slope at the beginning of the collection, b is a parameter related to the shape of the accumulation of new species during the collection, and t is the collecting effort (e.g., some unit of time) (Soberón and Llorente 1993; León-Cortés, Soberón-Mainero, and Llorente-Bousquets 1998). Using this method, the size and the amount of effort needed to achieve the total sample size of the El Edén butterfly fauna was assessed.

Habitat distribution and geographic range

The habitat distribution and geographic range data was analyzed for El Edén Papilionidae, Pieridae, Nymphalidae, Hesperiidae, and Lycaenidae butterflies from field observations as well as from the information given in the species account in DeVries (1987); Tyler, Brown, and Wilson (1994); and Opler and Wright (1999). Although Hesperiidae and Lycaenidae were also considered for the present analysis, some species were excluded because their habitat description was too scant or ambiguous.

The present analysis deals with 76 species of the 86 species reported in this paper. Following Thomas (1991), butterflies species were classified as occupying unmodified vegetation (when no mention was made that they occupy secondary or agricultural habitats), or as occupying modified vegetation (when mention was made that they occupy secondary, agricultural, or other human-modified habitats, although most of these species also occupy unmodified habitats). Some species that were classified as occupying only unmodified vegetation do make some use of modified habitats, but in most cases the latter are used very rarely by these species, and usually only when modified and unmodified habitats are adjacent (DeVries 1987). A habitat

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Number of individuals sampled 600 500 400 300 200 100 0 Cu m u la ti ve s p e c ie s 140 120 100 80 60 40 20 0

FIGURE 13.2. Plot of cumulative species versus the number of individuals sampled at El Edén. The data for the observed cumulative number of butterfly species are shown as circles on the plot, while the estimates (continuous line) of richness are based on Clench’s equation (R² = 0.99). Asymptotic value for Clench´s function is 163 species.

designation to each species before examining its geographic range category. Geographic range categories were consulted in DeVries (1987); Tyler, Brown, and Wilson (1994); and Opler and Wright (1999). The three geographic range categories used were defined as follows:

1. Restricted to southeast Mexico and Central America, but not farther south than Panama

2. North America to Central America, and/or southern Mexico to northern South America

3. Widespread in the Neotropics (at least reaching Brazil or Bolivia) Although the habitat distribution and geographic range categories were somewhat arbitrary, the general pattern, rather than the habitat-use patterns of a particular species. We also wanted to examine a possible overall association between habitat use and geographic range within each subfamily was also examined. Because the sample size for each subfamily was rather small, samples were collated to examine the overall trend. The proportion of species in each geographic range category that was restricted to unmodified habitats only was calculated. Thus, the whole sample had up to three values. Values were examined according to a declining or increasing proportion of species restricted to unmodified (or primary) habitats with increasing geographic range.

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RESULTS

Butterfly richness

Eighty-six species of butterflies were collected during a 35-day sampling period at El Edén (see Appendix 1). Nymphalidae was the richest family with 53 species (62 percent), followed by Pieridae with 13 species (15 percent), Hesperiidae with 12 species (14 percent), Papilionidae with 5 species (6 percent), and Lycaenidae with 3 species (3 percent).

The plot of cumulative species versus the number of individuals sampled for the El Edén data is shown in fig. 2. Using Equation (1), the estimated richness at El Edén is 163 species. If this estimate is accurate, then we sampled 53 percent of the El Edén butterfly fauna in 35 days. Although the regression line is a reasonably close match to the observed data points (R² = 0.99, Norusis [1994], Figure 13.2), the collections were conducted over a consecutive 35-day period during portions of two seasons (late summer and early autumn). Therefore, model predictions may be biased due to natural changes in the seasonal patterns and abundance of butterfly species. An important number of butterfly species remains to be recorded, mostly for Lycaenidae, Riodiniidae, and Hesperiidae families, which are extremely rich in Central America.

Habitat use and geographic range categories

The habitat use and geographic range categories assigned to each of the species included in the analysis are shown in Appendix 1. Fifteen species were placed in the unmodified habitats category, and 61 species were placed in the modified habitats category. A significant association between habitat use and geographic range for 76 butterfly species at El Edén (Table 13.1, χ² = 17.23, d.f. = 2, p < .001). Species with narrow geographic ranges were usually grouped in unmodified habitats, whereas widespread species were much more likely to make some use of human-modified habitats.

A declining proportion of species restricted to unmodified habitats with increasing geographic range (Figure 13.3, Table 13.2). Thus, the general trend at El Edén suggested that widespread butterfly species were more likely to make some use of modified habitats.

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TABLE 13.1. Association between habitat use and geographic range for Nymphalidae, Pieridae, Hesperiidae, Papilionidae, and Lycaenidae butterfly species at El Edéna.

Geographic range categories

Habitat use at El Edén 1 2 3

Unmodified habitats 6 2 7

Modified habitats 2 13 46

a

Association between habitat use and geographic range: chi square = 17.23, d.f. = 2, p < .001.

1 = Restricted to southeast Mexico and Central America, but not farther south than Panama; 2 = North America to Central America, and/or southern Mexico to northern South America; 3 = Widespread in the Neotropics (at least reaching Brazil or Bolivia).

DISCUSSION

Species richness

The butterfly inventory reported here represents a preliminary list of the butterfly fauna of El Edén. The faunistic data closely fit the model (see Figure 13.2) but, as mentioned previously, the data were collected over a consecutive 35-day period during portions of two seasons (late summer and early autumn) of 1995. Thus, the estimate of 163 species at El Edén is tentative. A species that is rare or absent one day is likely to be rare or absent the next, but may be common in another season or during the same season in a different year. For this reason, we expect a greater increase in the number of new species, particularly for largely unknown butterfly families such as Lycaenidae, Riodiinidae, and Hesperiidae. More conspicuous butterfly species (e.g., Papilionidae, Pieridae, and Nymphalidae) were sampled vigorously during the course of this study. Therefore, we would not expect to add many more new species for these families.

Habitat preferences and geographic ranges

Habitat availability (and the disturbance of it) may influence a species’ abundance and its distribution (e.g., Hughes 2000). Although disturbance may be brought about by human activities, natural events such as hurricanes, fires, and falling trees may also cause ecological disturbance of comparable magnitude. Depending on its historical scale and frequency, habitat modification may elicit a variety of responses from the members of a given Neotropical butterfly community (Raguso and Llorente-Bousquets 1991).

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Geographic range categories 3 2 1 P rop ort io n o f sp e c ie s 1.0 .8 .6 .4 .2 0.0 0.75 0.13 0.13

FIGURE 13.3. The proportion of butterfly species at El Edén, using unmodified habitats with increasing geographic range (see Table 13.2). For a description of geographic range categories, see text.

For instance, recent evidence supports the hypothesis that historical climate variations have had a great effect on British butterfly species assemblages (Dennis 1977; Thomas 1994). Janzen (1988) has emphasized that the resulting mosaic of habitats and successional stages of vegetation can accommodate more species of Lepidoptera than a pristine tropical dry forest (either past or future) could realistically support.

The prevalence of weedy, cosmopolitan species at El Edén conform well to the pattern observed by Welling (1966) for Papilio, Euptoieta, Zerene, and Phoebis species in patches cut into dense thorn forests of the Yucatán Peninsula. Nearly 20 percent of the species reported here have been considered as a fauna characteristic of disturbed habitats (see Raguso and Llorente-Bousquets 1991). Some of these butterflies are migratory habitat generalists with nonspecialized host plant requirements and wide distributions throughout the Neotropics, and could rapidly invade disturbed rainforest patches in the Yucatán Peninsula (e.g., Papilio toas, Ascia monuste, Phoebis sennae, P. Argante, Dannaus gilippus, and Marpesia chiron).

From a habitat point of view, there are important differences between cosmopolitan species that can exploit disturbed forest habitat and organisms that are adapted to undisturbed primary forests. When examining a species’ abundance and distribution across many habitat types, one must consider the availability of the habitats (MacNally 1989). It is possible to have a species that can only use one habitat type, but is relatively abundant in that habitat. If the habitat type is widespread, then the species will be distributed across a large range; conversely, if the habitat is rare, then the species will be narrowly

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TABLE 13.2. Association between habitat use and geographic range for butterflies in each subfamily of Nymphalidae, Pieridae, Hesperiidae, Papilionidae, and Lycaenidae at El Edén.

Geographic range categories

1 2 3 Habitat U M U M U M Nymphalidae Apaturinae 0 0 0 0 0 1 Nymphalidae Brassolinae 0 0 0 0 0 2 Nymphalidae Charaxinae 2 0 0 0 1 6 Nymphalidae Danainae 0 0 0 1 0 1 Nymphalidae Heliconiinae 0 0 0 0 0 4 Nymphalidae Melitaeinae 0 0 0 1 0 1 Nymphalidae Morphinae 0 0 0 1 0 0 Nymphalidae Nymphalinae 3 1 1 6 5 12 Nymphalidae Satyrinae 0 0 0 0 1 0 Pieridae Coliadinae 0 0 0 3 0 8 Pieridae Pierinae 0 0 0 0 0 2 Hesperiidae Pyrginae 0 0 1 1 0 5 Papilionidae Papilioninae 1 1 0 0 0 3 Lycaenidae Polyommatinae 0 0 0 0 0 1 Number of species 6 2 2 13 7 46

U = unmodified habitats; M = modified habitats

1 = Restricted to southeast Mexico and Central America, but not farther south than Panama; 2 = North America to Central America, and/or southern Mexico to northern South America; 3 = Widespread in the Neotropics (at least reaching Brazil or Bolivia).

distributed. Thus, abundance-distribution relationships can change depending on the geographic extent of the habitats in which species’ densities are measured (e.g., Gaston and Lawton 1990).

Few forest species that possess relatively narrow geographic ranges were classified as making use of modified habitats. These included: Achalarus toxeus, Dynamine talassina, Hamadryas februa ferentina, H. guatemalena guatemalena, H. iphtime, Memphis morvus boisduvali, Pyrrhogyra neaerea hypsenor, Taygetis virgilia rufomarginata, and Temenis laothoe agatha. Geographically restricted or endemic butterfly species are often regarded as biotope specialists, although they can be among the most abundant species at

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sites where they occur. Some biotope specialists might occupy disturbed habitats, but in most cases the latter are used very rarely by these species, and usually only when modified and unmodified habitats are adjacent (Thomas 1991). For instance, many savanna and tropical dry forest butterflies appear to congregate in mesic habitats, such as gallery forests, or migrate to higher elevations in the dry season (e.g., DeVries 1987). The microdistribution of butterflies in a habitat is influenced by a variety of abiotic and biotic factors such as the distribution of food, shelter, microclimatically favorable spots, or intraspecific and interspecific interactions (Loertscher, Erhardt, and Zettel 1995). Furthermore, many species may be specialists on naturally occurring successional habitats and so be able to occupy human-modified areas that provide the same successional conditions.

If the relationship between habitat use and geographic range reported here is repeated elsewhere in Mexico and Central America as past surveys suggest it is (e.g., Thomas 1991; Raguso and Llorente-Bousquets 1991), an increasing proportion of secondary vegetation throughout the Neotropics will produce a decreasing difference between the faunas of different regions (i.e., endemic species will be exterminated with the loss of primary forest, and the same widespread species will be left inhabiting secondary vegetation in each region). If unmodified habitats continue to be lost, local (alpha) diversity may not noticeably decrease—and may sometimes increase (e.g., Raguso and Llorente-Bousquets 1991)—but regional (beta) diversity will decline.

Although our conclusions are preliminary, the relatively equal time and effort we spent at different habitats within El Edén produced results that support this conclusion. It would be desirable to establish uniform biodiversity-monitoring protocols at a wide range of spatial scales and for different regions in the Yucatán Peninsula. These protocols would build on the results presented here and attempt to generalize the present status of butterfly diversity in the peninsula with the factors that are affecting changes in species composition.

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APPENDIX:

CHECKLIST OF THE BUTTERFLIES AT EL EDÉN ECOLOGICAL RESERVE,

QUINTANA ROO, MEXICO. Taxon Habitat use Geographic range HESPERIOIDEA HESPERIIDAE Pyrginae

Proteides mercurius (Fabricius) 2 3

Poligonus leo Gmelin 2 3

Chioides zilpa (Butler) 2 2

Urbanus proteus (Linnaeus 1758) 2 3

Urbanus dorantes (Stoll 1790) 2 3

Urbanus tanna Evans NA NA

Astraptes aulensis NA NA

Achalarus toxeus (Plötz 1782) 1 2

Xenophanes tryxus (Stoll 1780) 2 3

Heliopetes arsalte (Linnaeus 1767) NA NA

Oligoria maculata NA NA

Spathilepia clonius NA NA

PAPILIONOIDEA PAPILIONIDAE Papilioninae

Battus polydamas polydamas (Linnaeus 1758) 2 3 Papilio thoas Rothschild & Jordan 1906 2 3 Eurytides protesilaus (Rohschild & Jordan 1906) 2 3 Mimoides ilus branchus (Doubleday 1846) 2 1

Mimoides phaon (Boisduval 1836) 1 1

PIERIDAE Pierinae

Appias drusilla (Cramer 1777) 2 3

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Taxon Habitat use

Geographic range Coliadinae

Anteos clorinde Godart 1823 2 3

Anteos maerula Fabricius 1775 2 3

Phoebis argante (Fabricius 1775) 2 2

Phoebis sennae Linnaeus 1758 2 3

Aphrissa statira (Cramer 1777) 2 3

Eurema proterpia (Fabricius 1775) 2 3

Eurema boisduvaliana (Felder 1865) 2 2

Eurema dina westwoodi (Boisduval 1836) 2 2

Eurema albula (Cramer 1775) 2 3

Coliadinae

Eurema nise (Cramer 1775) 2 3

Eurema daira (Godart 1819) 2 3

LYCAENIDAE Theclinae

Pseudolycaena damo (Druce 1785) NA NA

Thereus zebina NA NA

Polyommatinae

Hemiargus ceraunus (Butler & Druce 1872) 2 3

NYMPHALIDAE Charaxinae

Prepona omphale octavia Fruhstorfer 1904 2 3 Archaeoprepona demophon centralis Fruhstorfer 1905 2 3 Archaeoprepona demophoon gulina Fruhstorfer 1904 2 3

Siderone marthesia (Cramer 1777) 2 3

Consul electra (Westwood 1850) 1 1

Anaea aidea (Guérin-Ménéville 1844) 1 1 Memphis morvus boisduvali (Comstock 1961) 1 3

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Taxon Habitat use

Geographic range

Memphis pithyusa (Felder 1869) 2 3

Memphis sp. NA NA

Apaturinae

Doxocopa laure (Drury 1773) 2 3

Nymphalinae

Colobura dirce (Linnaeus 1764) 2 3

Historis odius (Fabricius 1775) 2 3

Smyrna blomfildia datis Fruhstorfer 1908 2 2

Biblis hyperia (Cramer 1782) 2 3

Mestra amymone Menetries 1857 2 2

Hamadryas februa ferentina (Godart 1824) 1 3 Hamadryas glauconome glauconome (Bates 1864) 1 1 Hamadryas feronia farinulenta (Fruhstorfer 1916) 2 3 Hamadryas guatemalena guatemalena (Bates 1864) 1 3

Hamadryas iphtime (Bates 1864) 1 3

Hamadryas sp. NA NA

Dynamine thalassina Boisduval 1870 1 2

Dynamine mylitta (Cramer 1782) 2 3

Marpesia petreus (Cramer 1778) 2 3

Marpesia chiron (Fabricius 1775) 2 3

Marpesia alcibiades (Staudinger 1876) 1 1 Eunica tatila caerula Godman & Salvin 1887 2 2

Eunica monima modesta Bates 1864 2 3

Temenis laothoe agatha (Fabricius 1787) 1 3 Pyrrhogyra neaerea hypsenor Godman & Salvin 1884 1 3

Nymphalinae

Diaethria astala (Guérin-Ménéville 1844) 2 2

Adelpha basiloides (Bates 1865) 1 1

Adelpha iphiclus (Linnaeus 1758) 2 3

Adelpha fessonia Hewitson 1847 2 1

Siproeta stelenes biplagiata (Fruhstorfer 1907) 2 3

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Taxon Habitat use

Geographic range

Anartia jatrophae (Linnaeus 1763) 2 3

Junonia evarete Cramer 1782 2 3

Junonia coenia Hübner 2 2

Heliconiinae

Agraulis vanillae (Linnaeus 1758) 2 3

Dryas iulia (Fabricius 1775) 2 3

Heliconius charitonius (Linnaeus 1767) 2 3 Heliconius erato petiverana Doubleday 1847 2 3

Melitaeninae

Chlosine lacinia (Geyer 1837) 2 3

Thessalia theona (Menétries 1855) 2 2

Danainae

Danaus gilippus thersippus Bates 1863 2 2 Danaus eresimus montezuma Talbot 1943 2 3

Morphinae

Morpho peleides limpida Butler 1872 2 2

Brassolinae

Opsiphanes cassina fabricii (Boisduval 1870)

Opsiphanes cassina chiriquensis Stichel 1904 2 3 Caligo memnon memnon (Felder & Felder 1866) 2 3

Satyrinae

Taygetis virgilia rufomarginata Staudinger 1888 1 3

Cissia sp. NA NA

Note: Habitat use: (1) unmodified habitats only; (2) some use of modified

habitats. Geographic range: (1) Restricted to southeast Mexico and Central America but not farther south than Panama; (2) North America to Central America and/or southern Mexico to northern South America; (3) Widespread in the Neotropics (at least reaching Brazil or Bolivia). NA: Taxon for which habitat use and/or geographic information was not available; these species were excluded from habitat analyses. For taxonomic authorities see DeVries (1987); Raguso and Llorente-Bousquets (1991); Tyler, Brown, and Wilson (1994); and Opler and Wright (1999).

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