The butterflies of Turquino National Park , Sierra Maestra , Cuba ( Lepidoptera , Papilionoidea )

The butterflies of Turquino National Park, Sierra Maestra, Cuba (Lepidoptera, Papilionoidea).— Between February and November 2011, we conducted a species inventory, created a natural history database and a made a first approach to the composition and structure of the butterfly communities present at several vegetation types in the Turquino National Park. The inventory included 83 species, 29 of them endemic. We recorded 57 species (18 endemic) in transects along main vegetation pathways. In disturbed vegetation, species richness was higher (48) and abundance was better distributed, but the proportion of endemism was lower (23%). Species richness decreased and the dominance and proportion of endemism increased with altitude. Numbers of species and the proportions of endemism at natural habitats sampled were: 19 and 58% for evergreen forest, 10 and 60% for rainforest, eight and 100% for cloud forest, and four and 100% for the elfin thicket. Flowers of 27 plants were recorded as nectar sources for 30 butterfly species, and host plants were recorded for nine species.


Introduction
Butterflies are poorly represented in conservation planning in Cuba.Several protected areas of Cuba have species inventories of this group (Fontenla, 1987a(Fontenla, , 1987b(Fontenla, , 2005;;Hernández et al., 1994;Pérez et al., 1999;Núñez, 2004Núñez, , 2010) ) but most lack information relevant for their conservation, such as data concerning preferred habitats, food sources for adults and immature stages, population size, and so on.
Cuba has a rich Lepidoptera fauna that includes 192 species of butterflies and skippers (Núñez & Barro, 2012).Of this total, 33 species and 39 subspecies are endemic to the island.Located in the Sierra Maestra Mountains, the Turquino National Park (TNP) harbors several of the Cuban endemic butterflies (Schwartz & Hedges, 1991).The cited authors recorded 37 species, nine of them endemic species and subspecies, along a transect from the south coast to Pico Real del Turquino, 1,974 m, the highest of all mountains in Cuba.Schwartz & Hedges (1991) incompletely described the habitats occupied by some species.
The goal of the present study was to increase the inventory of TNP butterfly species, to conduct a first approach to the composition and structure of its butterfly communities at different vegetation types, and to record all possible natural history data on this group in order to contribute to its future conservation and management.

Study area and vegetation types
The TNP covers 242.1 km 2 in the highest portion of the Sierra Maestra Mountain range in southeastern Cuba, including the country's highest peak, the Pico Real del Turquino of 1,974 m.Vegetation of the Turquino massif is composed of several natural types of forests and thickets along different altitudes, as well as disturbed vegetation (grasslands and ruderal vegetation) resulting from human activity .Forests include evergreen forest between 300-800 m of elevation, rainforest at 800-1,600 m, and cloud forest between 1,600-1,900 m (Capote & Berazaín, 1989).Pine forests are also present at the TNP but were not sampled in this study.The Pico Real del Turquino and its vicinity are covered by a thicket formation that is known by several names but referred to here as elfin thicket, following Borhidi (1996).

Methods
The TNP was visited in February, June, September, and November 2011.Butterflies were inventoried and counted along two vegetation paths on the TNP northern slope.One is 13 km long.It runs from Alto del Naranjo, 900 m, to Aguada de Joaquín Station, 1,250 m, ending at Pico Real del Turquino, 1,974 m.The other path, 16 km long, starts at 250 m at the Santo Domingo town, goes to El Cojo Station, 1,200 m, and also ends at the top of Pico Real del Turquino.
During each visit, transects were placed at random to take samples, completing three sampling times at each vegetation type by the end of the study.Sampling covered at least 20 to 30% of the length of pathway where there was vegetation.The total number of transects at each vegetation type was 24 for disturbed vegetation, 36 for evergreen forest, 36 for rainforest, 24 for cloud forest, and 14 for elfin thicket.The minimum distance between two transects was 50 m.
Each transect was 100 m long and 4 m to each side of the observer.All butterfly species and individuals flying up to 5 m high were counted.In addition, any data on natural history (oviposition plant, nectar sources, and larval food plant) was recorded.Specimens requiring closer examination were netted and released at the same site after identification.Counts were made between 9:00 and 15:00 h, always on sunny days.Núñez All the species detected are reported in the species list and in the annex 1, where they have been classified as dominant, intermediate, or rare based on their position at the rank abundance curves for each habitat type.Species observed only outside transects were considered as rare.
Records of additional species were obtained by reviewing the entomological collection of the Institute of Ecology and Systematics (CZACC).

Species inventory
To date, 83 butterfly species have been recorded from the TNP, including 29 species and subspecies endemic to Cuba, representing five families (table 1, species list and annex 1).Nymphalidae is the family with the highest number of species (29), followed by Hesperiidae (26), Pieridae (17), Papilionidae (8), and Lycaenidae (3).During the field work, 71 species were recorded, 27 being endemic to Cuba.The other 12 species are only known from literature records or from the CZACC collection.
Butterfly communities 57 species (18 Cuban endemics) were recorded from the transect counts (table 2), while an additional 14 species were recorded outside the transects.Lowland to mid-altitude areas covered by disturbed vegetation were the most diverse, with 48 species, of which 16 were endemic (4 species and 12 subspecies).Butterfly abundance was also higher than at the other vegetation types, and was evenly distributed among species (fig.1).This community is dominated by Calisto herophile herophile, Nymphalidae, and Phoebis sennae sennae, Pieridae.
At higher altitudes with better preserved habitats, species richness was lower but the proportion of endemic butterflies was higher (table 2).All communities at natural habitats Table 1.Turquino National Park vegetation types, altitudinal range (A, in m), and total number of butterfly species (S), endemism (E), and percentage of endemism (%E).Records from the present study, literature, and specimens deposited at the IES collection.Tabla 1. Tipos de vegetación, rango altitudinal (A, en m), y número total de especies de mariposas (S), endemismo (E) y porcentaje de endemismo (%E) en el Parque Nacional Turquino.Registros procedentes de este estudio, referencias bibliográficas y ejemplares depositados en la colección del IES.except the evergreen forest are evidently dominated by Calisto smintheus, the only butterfly endemic to the Sierra Maestra Mountain range, followed by Anetia briarea numidia and Anetia cubana, two other Nymphalids endemic to Cuba (fig.2).The community in the evergreen forest was the richest of all natural habitats and had a more evenly distributed abundance.It was also dominated by C. smintheus, but to a lesser extent than in other habitat types (fig.2).An analysis by family indicated that all communities were dominated by the Nymphalidae (table 3).Only in disturbed habitats, the Hesperiidae had a similar number of species, although their abundance and number of endemisms was almost half that of Nymphalidae.Moreover, this was the only habitat where all the families were present.Habitats at higher elevations were characterized by a stronger dominance by the Nymphalidae, which represented more than two thirds of the communities´ abundance and number of endemisms (table 3).

Natural history data
A total of 27 plant species were recorded as nectar sources for 30 butterfly species (see species list and annex).The plant species most visited were Lantana camara (11 butterfly species), and Stachytarpheta jamaicensis (nine butterfly species).Nectar sources for the endemic species Greta cubana, Anetia cubana, and Anetia briarea numidia are provided here for the first time.

Discussion
The list of butterfly species from the TNP represents 43% of all members of this group in Cuba.The same percentage applies to the species endemic to Cuba, when both species (nine) and subspecies (20) are considered.The only family lacking is Riodinidae, with a single Caribbean species restricted, both ecologically and geographically, to the north keys of central Cuba, the extreme southeastern coast, and northeastern mountains (Alayo & Hernández, 1987;Hernández et al., 1998;Rivero et al., 2003).Compared to the only previous study, performed by Schwartz & Hedges (1991), we observed an additional 34 species during field work, including 20 endemic species.We did not detect four other species that were reported by Schwartz & Hedges.Differences in species richness and composition might be explained by environmental dissimilarities (humidity, habitats, and conservation status) between the northern slope, where present work was conducted, and the southern slope where Schwartz & Hedges (1991) worked.In addition, sampling effort was greater in this study than in the six working days in the previous study.
The species richness per family is comparable to that of the entire Cuban butterfly fauna (Alayo & Hernández, 1987;Smith et al., 1994).Schawrtz & Hedges (1991) found a similar family proportion but observed more Pieridae species, probably because they spent more time collecting in open, disturbed areas.
Habitat disturbance increased species richness by diminishing endemism and adding generalist widespread species.Sixteen of the 17 butterflies listed by Fontenla (1992) as members of the Cuban butterfly generalist group were present at disturbed areas of the TNP.This author also found that Cuban Nymphalidae, including many endemic species, were better represented in the mountain forests, as occurred in the TNP, where this family attained the highest diversity in the rainforest, the cloud forest, and the elfin thicket.
Low to moderate levels of disturbance can even increase butterfly diversity by offering additional resources to some endemic species, such as nectar sources close to evergreen forest (as recorded in La Platica village).Fontenla (1987aFontenla ( , 1987bFontenla ( , 1992) ) and Núñez & Barro (2003) found that edges of several Cuban vegetation types have higher values of butterfly diversity than pure habitats.However, severe disturbances (e.g.forest degradation to grassland by fires), caused impoverishment of the butterfly communities.At TNP, Loma del León suffered this kind of change and during the present study only two generalist non-endemic species, Agraulis vanillae and Leptotes cassius, were found there.
Results from this study show that the TNP has a diverse butterfly fauna that needs to be preserved.More studies should be conducted in this area to increase the inventory of its species and to add more data to the presently known natural history.Information gathered during the present work will help to improve park management and to develop existing ecotourism.