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Journal of the Lepidopterists' Society 53(4), 1999, 153-158

THE SPHINGIDAE (HETEROCERA) OF THE "EL OCOTE" RESERVE,

CHIAPAS, MEXICO

Olga Lidia Gomez-Nucamendi, Robert W. Jones1 and Alejandro Moron-Rios 2

El Colegio de la Frontera Sur, Apdo. Postal 63, C.P. 29290 San Cristobal de las Casas, Chiapas, MEXICO

ABSTRACT. A study of the family Sphingidae was conducted in the Reserve, "El Ocote," located in the northeast portion of the state of Chiapas, Mexico. Collections were made principally using a blacklight between 1994 and 1997. A total of 60 species were collected, from 20 genera in five tribes and three subfamilies. Xylophanes, Manduca, Eumorpha and Erinnyis had the greatest number of species. Nyceryx mulleri Clark is a new record for Chiapas. Eighty percent of the species were collected in the first two months of the rainy season. It was estimated that the present collection accounted for 75% of the species of Sphinigidae in the reserve, based on the accumulation of species per collection effort. The fauna of the Sphingidae collected from "El Ocote" was compared with that reported from two other reserves of southern Mexico, "Los Tuxtlas", Veracruz and "Chajul", Chiapas. "El Ocote" and "Chajul" were the most similar with 87% affinity.

Additional key words: tropical forests, biodiversity, neotropical region, invertebrate inventory.

RESUMEN. Se presenta un estudio faunistico de la familia Sphingidae de la selva "El Ocote", ubicada al noreste del estado de Chiapas, Mexico. Se efectuaron colectas con trampa de luz tipo pantalla entre 1994 y 1997, que aportaron 60 especies de 20 generos comprendidos en cinco tribus y tres subfamilias. El mayor numero de especies se distribuye en los generos Xylophanes, Manduca, Eumorpha y Erinnyis. Nyceryx mulleri Clark es un nuevo registro para Chiapas. El 80% de las especies se colect6 en los dos primeros meses de la temporada lluviosa. La estimation de la riqueza de especies por esfuerzo de colecta indica que esta investigation aporta el 75% de las especies de esta region. Se incluye una comparacion con la fauna de Sphingidae de Los Tuxtlas, Veracruz y Chajul, Chiapas, resultando que El Ocote y Chajul presentan una afinidad de 87%.

There has been tremendous international publicity and concern surrounding the unprecedented rate at which tropical forests are being lost. The concern is well warranted because although tropical forests cover only about 7% of the terrestrial surface of the planet, they probably support around 50% of the flora and fauna (Myers 1986). Despite this diversity, these forests are increasingly threatened and destroyed.

Although much of the international attention has been directed at the larger tracts of tropical forest such as the Amazon and the Congo basin of Africa, Mexico has important tracts of tropical forests that merit serious conservation efforts. However, it is estimated that 80% of the tropical forests of Mexico have already been destroyed and those that remain are seriously threatened (Estrada et al. 1995). This is the case with the tropical forest reserve, "El Ocote" in the northeastern Chiapas. Of the 48,140 hectares designated as reserve lands in 1982 (Diario Oficial de la Federacion 1982), only 57%, (27,437 ha), can now be considered as forested and not directly affected by agricultural activities (Garcia et al. 1996).

As for other tropical forests, a high diversity of vertebrates have been reported from "El Ocote" Reserve (Dommguez et al. 1996, Mufioz et al. 1996, Navarrete-Gutierrez et al. 1996), but there is a lack of inventories of the invertebrate fauna. Knowledge of the invertebrate diversity could complement the findings con-

1  Present address: Licenciatura en Biologia, Universidad Auto-noma de Queretaro, Apdo. Postal 184, C.P. 76010, Queretaro, Qro., MEXICO

2 Address correspondence to this author

cerning the vertebrate fauna, and give further information on the species most threatened by habitat destruction, and, in general, give an indication of the biological significance of the reserve based on species richness (Toledo 1988).

The present study is a contribution to the knowledge of the Lepidoptera fauna of southeastern Mexico, and in particular, of the family Sphingidae of the "El Ocote" Reserve of northwestern Chiapas. An inventory was conducted of the Sphingidae of the Reserve, from which a comparison was made of the species richness of this family reported from two other tropical forests of southern Mexico. "Los Tuxtlas" of the Gulf Coast of Veracruz, and "Chajul" of the Lacandon region of eastern Chiapas. These forests, together with "El Ocote", presumably formed a single tract of tropical forest stretching from the Gulf coast of Veracruz to what is now the border with Guatemala (Challenger 1998).

Materials and Methods

Description of the study area. The protected forest and fauna reserve known as the "Selva El Ocote" is located in the northwest portion of the state of Chiapas (16°53'-17°05'N and 93°30/-93°47/W) in the municipality of Ocozocoautla de Espinoza (Fig. 1). The reserve has an area of 48,140 ha (Diario Oficial de la Federacion 1982) and ranges in altitude from 180 to 1500 m above sea level. The mean annual temperature and precipitation is 25.2°C and 2387 mm, respectively (INEGI1984).

The soils of the reserve are thin and fragile, of limestone origin, with large numbers of exposed rocks and boulders (Garcia et al. 1996). The topography is highly

Journal of the Lepidopterists' Society

17° 05'

16° 53'

Fjg. 1. A. Geographic position of the "El Ocote" Reserve in Southeastern Mexico. B. Detail of the reserve and location of sampling areas: 1= El Encajonado, 2 = 9 km SW of Ejido Cuauhtemoc 3 = Finca Nueva Providcncia.

irregular and has contributed to the existence of very diverse floral assemblages, of which the high tropical semi-evergreen tropical forest is the dominant vegetation. This forest type coexists with fragments of medium height evergreen tropical forest, low semi-evergreen tropical forest, low deciduous tropical forest, and associated successional stages: "acahuales," savanna, and pastures (Ochoa-Gaona 1996).

Sampling. Night sampling was conducted at three locations that mostly comprised well preserved medium height evergreen tropical forest: El Encajon-ado, Ejido Cuauhtemoc and Finca Nueva Providen-cia (Fig. 1). These three locations provide a good representation of the spatial complexity of this type of tropical rain forest in the Reserve. Collections were mostly made during 1994. Supplementing field work

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Table 1. Characteristics of the collection localities compared in this study. Vegetation categories are: TRF = Tropical Rain Forest, ESF : Evergreen Seasonal Forest, TDF = Tropical Deciduous Forest, and SS = Short-tree Savanna (Breedlove, 1981).

was conducted in 1995, 1996 and 1997. Each collection coincided with the new moon and had an average duration of five days. A twelve volt ultraviolet light and sheet were placed from 1800 to 0500 h each night to attract moths. At the time of capture, each captured specimen was injected in the thorax with 95% ethyl alcohol and placed in individually labeled glassine envelopes. Species were identified using Hodges (1971), D'Abrera (1986) and by comparison with reference material from the Entomology Collection of the Institute) de Biologia, Universidad Nacional Autonoma de Mexico (IBUNAM), in Mexico City. Collected specimens were deposited in the Entomology Collection of the El Colegio de la Frontera Sur, Unidad San Cristobal, Chiapas (ECOSC-E) and the entomology collection of IBUNAM.

Analysis of data. Estimations of the species richness of the Sphingidae of "El Ocote" Reserve were based on the Clench equation of species accumulation:

S(t) = at/(l+bt)

where S (t) is the expected number of species at time (t), a is the list increase rate, b is the species accumulation parameter, and the asymptote is given as a/b (Soberon & Llorente 1993). The model estimates total number of species present in the study area based on the characteristics of the decrease in new species collected as more time is spent in the field, this process will eventually generate an asymptote as an estimated total number of species (Soberon & Llorente 1993, Leon-Cortes 1995). The model was fitted by the non linear regression module provided by the package SPSS (v.6.1) using Levenberg-Marquardt algorithm.

The fauna of Sphingidae from "El Ocote" Reserve was compared with that reported for the Chajul Biological Station, Chiapas (Leon-Cortes & Pescador 1998) and Los Tuxtlas, Veracruz (Beutelspacher 1989) (Table 1), using the Simpsons similarity Index. This index is appropriate when compared faunas are disproportionate in size and number of shared taxa (Sanchez & Lopez 1988). We applied cluster analyses using the unweighted arithmetic average clustering method

(UPGMA) to show the total relationships among these faunas (Crisci & Lopez 1983).

Results and Discussion

Species richness and seasonal abundance. A total of 60 species of the family Sphingidae were collected from the "El Ocote" Reserve, belonging to three subfamilies, five tribes, and 20 genera (Table 2). The genus with the largest number of species was Xy-lophanes with 15 species, followed by Manduca with 10 species, and both Eumorpha and Erinnyis had five species each. These four genera represented 58% of the species collected in "El Ocote". Leon-Cortes and Pescador (1998) reported the same pattern in the abundance of species per genera from the Chajul Biological Station, in eastern Chiapas, and, in general, this also appears to be common to other tropical forests of America (Leon-Cortes & Pescador 1998).

The species collected from "El Ocote" Reserve represented 64% of the sphingids collected in the state of Chiapas and are all new records for the locality. Of these, Nyceryx mulleri Clark is a new state record. With our records, Chiapas has 49% of the Sphingidae reported from Mexico by White et al. (1991).

We estimated a species accumulation curve using Clench's equation. Our collection represents 75% of the Sphingidae of "El Ocote" Reserve predicted over 100 nights. Additional collection effort may provide only 10 to 15 more sphingid species (Fig. 2).

Three abundance categories were established using the criteria of Rabinowitz et al. (1986) and based on the numbers of specimens of each of the species collected. These categories were: "rare," (1 to 2 specimens); "common," (3 to 19 specimens); and "abundant," (20 to 50 specimens). Using these groupings, 16 of the collected sphingids were rare, 40 were common, and 4 were abundant (Table 2).

The number of species collected varied greatly between seasons. Forty-six species (77%) were only collected during the rainy season (May to October), whereas only one species was found exclusively during the dry season, as compared to 13 species (21%) which were found in both wet and dry seasons. Of the species

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Journal of the Lepidopterists' Society

Table 2. List of species of Sphingidae (Insecta: Lepidoptera) collected in the "Selva El Ocote" Reserve in Chiapas, Mexico during 1994-1997. F = February, A = April, M = May, J = June, Jl = July, Au = August, O = October and N = November, Total = Total number of specimens captured.

Species

Month

Total

Subfamily Sphinginae Tribe Sphingini

Cocytius lucifer (Rothschild and Jordan, 1903)

Neococytius cluentius Cramer, 1775

Manduca dilucida Edwards, 1887

Manduca occulta Rothschild and Jordan, 1903

Manduca lefeburei (Giierin,1844)

Manduca ochus (Klug,1836)

Manduca rustica (Fabricius, 1775)

Manduca albiplaga (Walker, 1856)

Manduca muscosa (Rothschild and Jordan, 1903)

Manduca corallina (Dnice, 1883)

Manduca lichenea (Burmeister,1856)

Manduca florestan Cramer, 1782

Sphinx leucophaeta Clemens, 1870

Sphinx merops (Boisduval,1870)

Subfamily Ambulicinae

Tribe Smerinthini

Protambulyx strigilis (Linnaeus, 1771)

Adhemarius gannascus (Stoll,1790)

Adhemarius ypsilon Rothschild and Jordan, 1903

Subfamily Macroglossinae

Tribe Dilophonotini

Pseudosphinx tetrio (Linnaeus, 1771)

Isognathus rimosus Grote,1865

Erinnys alope (Drury,1770)

Erinnyis lassauxi (Boisduval,1859)

Erinnys ello (Linnaeus,1758)

Erinnys oenotrus (Cramer, 1782)

Erinnyis obscura (Fabricius,1775)

Pachyliaficus (Linnaeus, 1758)

Pachyliodes resumens (Walker, 1856)

Herneroplanes ornatus (Rothschild and Jordan, 1894)

Hemeroplanes triptolemus (Cramer, 1779)

Madoryx oiclus (Cramer,1779)

Madoryx pluto Cramer, 1779

Callionima innus (Rothschild and Jordan,1903)

Callionima parce Fab ricius, 1775

Callionima falcifera (Gehler,1943)

Enyo lugubris (Linnaeus, 1777)

Enyo ocypete (Linnaeus, 1758)

Enyo gorgon (Cramer, 1777)

Perigonia lusca Fabricius,1777

Subfamily Macroglossinae

Tribe Philampelini

Eumorpha anchemola (Cramer, 1780)

Eumorpha triangulum Rothschild and Jordan, 1903

Eumorpha elisa (Smyth, 1901)

Eumorpha satellita Linnaeus,1771

Eumorpha uitis (Linnaeus, 1758)

Subfamily Macroglossinae

Tribe Macroglossini

Cauthetia spuria Boisduval,1875

Nyceryx mulleri Clark, 1917

Nyceryx riscus Schaus,1890

Xylophanes pluto (Fabricius,1777)

Xylophanes tyndarus (Boisduval,1875)

Xylophanes pistacina (Boisduval,1877)

Xylophanes porcus (Hiibner,1829)

Xylophanes ceratomiodes (Grote and Robinson, 1867)

Xylophanes anubus (Cramer, 1877)

Xylophanes amadis amadis Stoll

Xylophanes amadis cyrene (Druce,1777)

Xylophanes beIti (Druce,1878)

Xylophanes eumedon (Edwards,1887)

Xylophanes turbata Edwards, 1887

Xylophanes chiron nechus Drury,1770

Xylophanes libya (Druce,1878)

Xylophanes neoptolemus (Stoll, 1782)

Xylophanes thyelia Linnaeus, 1758

Au

J

M,J

F, M,J, O, N

M,J

J Jl

M,J, O

J J

M M,J,J1, N

M, J, Jl

F F, M,J

J.J1.0

M, J, Jl, Au, O. N M.J,Jl,Au, O, N

],Jl,Au, O, N M,J FJ

J M,J M,J

M M,J1 M,J

M F, M M, Jl

M

Jl

M, J, Jl, N

F, J, Jl, Au, O

M

O

A, M

M,J

M,J

F, J, Jl, Au, O, N

L Jl

FJJl M,J

M,J,J] M M

M,J M,J M,J M,J M, J, Au, O J,Jl,Au J,Au J,Au M,J1

J

M,J

M,J M, J, Jl, A M,J, A, O

O, N

1 1 4 12 5 3

10 4 4 1 7 15 1 4

5 13 20

7 4 3 1 7 10 1 3 4 3 1

14 1 1

18 26 1 1 2 5

3 10

8

25 4

47 1 1 3 6 7 3 8

18 2 9 3 2 4 7 10 4 4

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40

Number of nights

Fig. 2. Species accumulation curve of Sphingidae as a function of collection effort from "El Ocote", Reserve, Mexico. (° = Cumulative number of expected species, (♦ = Cumulative number of observed species.

Los Tuxtlas

Chajul

____Fl nr.nlp

Chajul-Los

Tuxtlas

El Ocote-Los

Tuxtlas

El Ocote-Chajul

El Ocote-Chajul-Los Tuxtlas

a

^M^«^

E

W//////////////A

=F=P

E

5 10 15 20 25 30 35 40 45 50 55 60

D Species

Fig. 4. Number of genera and species from the three localities compared from southern Mexico: A. number of genera and species not shared between sites, and B. number of genera and species shared between sites.

of sphingids collected during the rainy season, 48 of these (80%) were collected during the first two months of the season in May and June. Towards the end of the wet season, the abundance of each species declined and with fluctuations at low densities, in a manner similar to that observed during the dry season. This seasonal pattern in abundance and activity is similar to that reported in other studies (Haber & Frankie 1989, Janzen 1984, 1986, Powell & Brown 1990, Pescador 1994, Gregg et al. 1993). Haber and Frankie (1989) and Janzen (1984) state that precipitation and vegetative productivity are closely linked to the life cycle of the sphingids. During the dry season, the majority of the sphingids are in the pupal stage, while the larval and adult stages are found during the wet season. This pattern is reflected in reduced photosyn-thetic activity and productivity in host plants during the dry season, including the complete loss of leaves in some species, with the resumption of productivity during the wet season.

Similarity among fauna of the Sphingidae in Southern Mexico. The similarity among the sphingid

Fig. 3. Dendrogram of Sphingidae from three sites in southern Mexico. The unweighted arithmetic average (UPGMA) was used to cluster related groups.

fauna at the three sites analyzed are presented in Fig. 3. Within the three localities, 29 genera and 100 species have been collected, of which only 15 genera and 44 species were shared. The dendrogram derived from the cluster analysis indicates that there is greater similarity between "El Ocote" and the "Chajul" region of the Lacandon Forest of eastern Chiapas, than with "Los Tuxtlas" located on the Gulf coast (Fig. 3). These two latter localities shared 18 genera and 58 species, whereas "El Ocote" shared 17 genera and 52 species with "Chajul" and 16 genera and 46 species with "Los Tuxtlas"(Fig. 4).

In 1998 forest fires devastated approximately 7.4% of "El Ocote" reserve during the National severe drought associated with the El Nino phenomenon of 1997 (Aguilera 1998). Since this study represents the only non-vertebrate faunal data available of the species richness of a specific group from the reserve before the fires, the data presented here may serve as a baseline for estimating the speed and nature of the recovery of the reserve following this major disturbance event. In addition, Sphingidae, being highly vagile species, easily monitored, and with relatively high diversity may be an especially appropriate group of organisms in the study of what is hopefully a process of recovery.

Acknowledgments

The authors wish to thank Jean Haxaire of the Museo National d'Histoire Naturelle de Paris and Carlos R. Beutelspacher-Baigs of the Instituto de Biologia, Universidad Nacional Autonoma de Mexico for their comments and help in confirming the identification of some of the species. We are in debt with different colleagues of El Colegio de la Frontera Sur: Pedro F. Quintana-Ascencio read and gentiy commented on a preliminary version of the manuscript and helped with the calculations of the Clench model, Trinidad Aleman-Santillan made valuable comments at the beginning of the project, Miguel A. Vazquez-Sanchez provided important logistic support during field work, and Manuel Giron-Intzin for his help in the field

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and with the preparation of the specimens. El Colegio de la Fron-tera Sur (ECOSUR) provided logistical and financial support to the authors.

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Received for publication 29 December 1998; revised and accepted 2 February, 2000.