Revista Colombiana de Entomología 32(2): 191-199 (2006) Phytophagous from , 191

Phytophagous scarab from the Central Region of Guerrero, Mexico (Coleoptera: Scarabaeidae: Melolonthinae, , , Cetoniinae)

Coleoptera Scarabaeidae fitófagos de la Región Central de Guerrero, México (Melolonthinae, Rutelinae, Dynastinae, Cetoniinae)

CUTBERTO PACHECO F.1, CUAUHTÉMOC DELOYA1 , PEDRO CORTÉS G.2

Abstract. From July 1999 to June 2000, 1.307 specimens of phytophagous Scarabaeidae representing four subfamilies, 11 tribes, and 57 species of the following genera were collected: Calomacraspis, Macraspis, Pelidnota, Chrysina, Anomala, Strigoderma, Cotinis, Hologymnetis, Euphoria, Golofa, Strategus, Cyclocephala, Tomarus, Bothynus, Phyllophaga, Diplotaxis, Polyphylla, Isonychus, and Chnaunanthus. The area surveyed included the Mochitlán, , , and re- gions located in the central part of the state of Guerrero at an altitude of 840-1.600 m. These regions are characterized by six types of vegetation: pine forest, pine-oak forest, oak forest, tropical deciduous forest, palm groves, riparian forest, and pasture land. The 1307 specimens were captured using fer- mented fruit traps and by careful examination of herbaceous, brush, and arboreal vegetation in depos- its of the detritus produced by (Atta mexicana, Hymenoptera: Formicidae) in a dead forest as well as by nocturnal collection. Specific richness for Mochitlán is 22 species, Tixtla 23, Chilpancingo 32, and Atlixtac 34; Phyllophaga and Euphoria make up 66.66% of the species, Euphoria subtomentosa being predominant. Central Guerrero has greater specific similarity with species from the high part of the Balsas Basin (southern Morelos 46%, Cuernavaca, Morelos 38%) than with fauna established on the Mexican Pacific slope (Chamela, Jalisco 30%, and Tepic, Nayarit 29%). Key words: Fauna. Scarab beetles. Balsas Basin. Fruit traps. Resumen. El presente estudio se realizó entre julio de 1999 y junio del 2000 en Mochitlán, Atlixtac y Chilpancingo en la región centro del estado de Guerrero, México, en altitudes entre los 840 y 1.600 m y caracterizadas por seis tipos de vegetación: bosque de Pinus, Pinus-Quercus, bosque tropical caducifolio, palmas, vegetación riparia y pastos inducidos. Se obtuvieron 1.307 especímenes que representan 4 subfamilias, 11 tribus y 57 especies de los géneros: Calomacraspis, Macraspis, Pelidnota, Chrysina, Anomala, Strigoderma, Cotinis, Hologymnetis, Euphoria, Golofa, Strategus, Cyclocephala, Tomarus, Bothynus, Phyllophaga, Diplotaxis, Polyphylla, Isonychus, and Chnaunanthus. Los especímenes capturados fueron obtenidos mediante el uso de trampas con fruta fermentada, en la vegetación arbustiva, herbácea y arbórea, en depósitos de detritos de la hormiga Atta mexicana (Hy- menoptera: Formicidae) y en arbolado muerto y en colectas nocturnas. Chilpancingo presenta la mayor riqueza específica con 32 especies, seguida por Tixtla (23) y Mochitlán (22); Phyllophaga y Euphoria concentran al 66.66% de las especies y E. subtomentosa es la especie predominante. La región central de Guerrero presenta una mayor similitud específica con otras localidades establecidas en la parte alta de la Cuenca del Río Balsas (Sur de Morelos 46%, Cuernavaca, Morelos 38%) que con las establecidas en la vertiente del Pacífico Mexicano (Chamela, Jalisco 30% y Tepic, Nayarit 29%). Palabras clave: Fauna. Escarabajos. Cuenca del Balsas. Trampas de frutas.

Scarabaeidae fauna of the Mexican Pa- Pinus, Quercus-Pinus, Pinus-Quercus, and ment of diverse fauna. Phytophagous cific slope and in the Balsas Basin is Abies forests, mesophilous mountains, Scarabaeidae in Guerrero are represented composed of 29 to 48 genera and 70 to tropical deciduous forest, palm groves, by five subfamilies (Melolonthinae, 120 species (Deloya et al. 1993). The riparian forest, and tropical semideciduous Rutelinae, Dynastinae, Cetoninae, and location, size, orography, and biogeo- forest, among others. These numerous as- Trichiinae) that include 29 genera with graphical history of the state of Guerrero sociations, many of which are endemic to 120 species (Morón et al. 1997). The ob- have created a mosaic of different veg- the region, combine with the other charac- jective of the present study was to perform etation associations: xerophilous brush, teristics of the state to favor the establish- a preliminary analysis of phytophagous

1 Autor correspondencia Cuauhtémoc Deloya. Departamento de Entomología, Instituto de Ecología, A.C. (CONACYT), Km 2.5 carretera antigua a Coatepec 351, Congregación El Haya, CP 91070, Xalapa, Veracruz, MEXICO. E-mail: [email protected] 2 Laboratorio de Fitopatología, Facultad de Ciencias Químico Biológicas, Universidad Autónoma de Guerrero. Av. Lázaro Cárdenas s/n, Ciudad Universitaria, Chilpancingo, Guerrero. 192 Revista Colombiana de Entomología Cutberto Pacheco F. & Cols.

Scarabaeidae fauna inhabiting the cen- cept for February and April (Table 3, Fig. Key for the identification of tral region of the state of Guerrero in 1). Specific richness (S) and abundance phytophagous Scarabaeidae species order to elaborate a key for the iden- (a) throughout the year were as follows: found in the central part of Guerrero tification of the species and its compari- July S=8, a=1.68%; August S=4, a= 1 Base of antennal scape covered by son with other fauna obtained from other 0.91%; September S=7, a=1.29%; Octo- the anterior angle of front and ocu- localities. ber S=19, a=37.69%; November S=12, lar canthus, not visible from above a=8.33%; December S=7, a=1.83%; Janu- Study areas. The central region of the ...... 2 ary S=3, a=0.99%; March S=1, a=0.15%; state of Guerrero, Mexico, is located be- May S=36, a=29.96%, and June S=25, a 1’ Base of antennal scape visible from tween 17°30’ and 17°39’N and 99°23’ and =17.12%. As for seasonal richness, 14 spe- above through anteocular indenta- 99°42’W. Climatic characteristics, alti- cies coexisted in summer, 23 in autumn, tion. Mesepimeres not covered by tude, annual rainfall, and mean annual 4 in winter, and 39 in spring. base of elytra. Lateral borders of temperature for the study areas are shown elytra with wide indentation and in Table 1. Vegetation type per region is The following species constituted short, abundant setae. Metatarsus as follows: a) Chilpancingo: pine forest, 89.15% of the total sampling (N=1307) shorter than metatibia ...... pine-oak, oak, tropical deciduous forest, and were represented by 15 or more ...... CETONIINAE 4 palm grove, riparian forest, and pasture specimens: C. mutabilis (4.96%), E. ba- land; b) Mochitlán: tropical semide- salis (8.56%), E. leucographa (1.29%), 2 All tarsal claws equal in length and ciduous forest, oak, induced pasture, E. iridescens (3.28%), E. biguttata thickness, dentate, bifid, or entire. thorny brush, riparian forest; c) Tixtla: (1.75%), E. subtomentosa (27.9%), A...... 3 oak, palm grove, tropical deciduous for- inconstans (3.66%), Anomala sp. 2’ All tarsal claws differing in length est, riparian forest, and pasture land; d) (2.37%), S. aloeus (1.22%), C. lunulata and thickness, the majority grooved : tropical deciduous forest (12.69%), D. atramentaria (2.29%), P. and the minority entire ...... featuring trees less than 15 m high with ardara (1.98%), P. crinalis (1.6%), P...... RUTELINAE 17 robust, twisted trunks. integriceps (1.68%), P. brevidens 3 Claws entire or bifid. Mandible apex Material and Methods (1.29%), P. fulviventris (1.14), P. cre - nonycha (1.75%), P. obsoleta (6.11%) hidden under clypeus, not dorsally From July 1999 to June 2000, monthly and P. scabripyga (3.36%), while 38 other visible ..... MELOLONTHINAE 34 sampling was done in Chilpancingo, species made up only 10.85% of the to- 3’ Claws entire or single (at least inter- Mochitlán, Tixtla, and Atlixtac (Chi- tal with 12 specimens or fewer. mediate and posterior claws). Apex chihualco). Samples were collected both day and night from the arboreal, brush,

and herbaceous strata as well as from flow- -( -(

ers and fruits. Samples were taken from ,) ,)

public lighting installations at night and

by using fermented fruit traps (banana and ,( ,(

pineapple with beer) during the day

+) +)

(Morón 1997). Voucher specimens were

B DA ? deposited in the Entomological Collec- +( +(

tion (IEXA) of the Instituto de Ecología,

*) *) 0 ??@?  <? @A A.C. and M. A. Morón (MXAL) in Xalapa,

Veracruz, Mexico. *( *(

For the data analysis, the number of spe-

) )

cies obtained was recorded (species rich- ness; S = alpha diversity), as was the total ( ( number of specimens (N) for each site. ./0123.45/5. Sørensen’s (1948) Similarity Index was 5  used to determine beta diversity QS = 2(c) Figura 1. Monthly relation between the percent abundance (Line; N=1307) and specific / a + b, where a is the number of species in richness (bars; S= 57 species) of the phytophagous Scarabaeidae (Melolonthinae, Rutelinae, community A, b is the number of species Dynastinae Cetoniinae) from the Central Region of Guerrero, Mexico, July 1999-June 2000. in community B and c is the number of species shared by communities A and B. The key was prepared following the taxo- nomic criteria used by Morón (1984) and Deloya et al. (1995). Table 1. Abiotic factors for study areas in the central region of the State of Guerrero, Mexico (García 1988).

Results Region Rainfall Temperature Altitude Climate A total of 1.307 specimens of phyto- mm/year °C (annual mean) maslmosl phagous Scarabaeidae were collected, representing 4 subfamilies, 11 tribes, 19 Mochitlán 1239.0 24.2 840 Aw(w)igw” genera, and 57 species (Table 2). Atlixtac 717.5 23.2 1210 BS1(h´)w(w)igw” Chilpancingo 827.4 21.7 1360 A(C)wo(w)(i´)w” Melolonthinae from the central part of Tixtla 1014.9 21.5 1600 A(C)w1(w)(i’)g Guerrero were recorded year round ex- Phytophagous Scarabaeidae from Guerrero, Mexico 193

Table 2. Species of Scarabaeidae pleurosticti from the central region of the state of Guerrero, of mandible visible from dorsum .. Mexico, captured between July 1999 and June 2000...... DYNASTINAE 27

1 Melolonthinae 4 Scutellum covered by the basal lobe A) Melolonthini of pronotum. Gymnetini...... 5 Phyllophaga (Phyllophaga) ardara Saylor, 1943 P. (P.) brevidens (Bates, 1888) 4’ Scutellum exposed from above P. (P.) crinalis (Bates, 1888) ...... Euphoria P. (P.) dasypoda (Bates, 1888) P. (P.) disca Saylor, 1943 7 P. (P.) eniba Saylor, 1943 P. (P.) fulviventris (Moser, 1918) 5 Head with frontal projection fused P. (P.) integriceps (Moser, 1918) or partially free Cotinis Burmeister P. (P.) ravida (Blanchard, 1851) P. (P.) martinezpalaciosi Morón, 1988 ...... 6 Phyllophaga (P.) sp. P. (P.) setifera (Burmeister, 1855) 5’ Head lacking frontal projection. P. (P.) crenonycha Saylor, 1943 Anterior margin of clypeus highly P. (Phytalus) epulara Sanderson, 1958 marginate. Mesometasternal projec- P. (Ph.) obsoleta (Blanchard, 1851) P. (Listrochelus) sp. tion with sharp apex internally pro- P. (Chlaenobia) scabripyga (Bates, 1850) jected ...... Hologymnetis cinerea Diplotaxis atramentaria Bates, 1888 D. cribriceps Bates, 1889 6 Frontal projection free, less than D. megapleura Vaurie, 1960 D. trapezifera Bates, 1887 50% of length. Apically enlarged in Polyphylla petiti (Guérin-Méneville, 1844) dorsal view. Projection of clypeus variable. Coloration opaque, black, B) Macrodactylini Isonychus ocellatus Burmeister, 1855 greenish, and velvety. Total length 22-30 mm ...... Cotinis mutabilis C) Incerta sedis Chnaunanthus discolor Burmeister, 1844 6’ Frontal projection fused for 65% of 2 Rutelinae its length from vertex to clypeus. A) Dorsal coloration dark green. Meso- Calomacraspis splendens (Burmeister, 1844) metasternal projection rounded. To- Macraspis aterrima (Waterhouse, 1881) Burmeister, 1844 tal length 19-22.5 mm ...... Chrysina macropus (Francillon, 1795) ...... Cotinis pauperula B) Anomalini 7 Species longer than 15 mm ...... 8 Anomala cincta Say, 1835 7’ Species shorter than 15 mm ...... 11 A. foraminosa Bates, 1888 A. forreri Bates, 1888 8 Clypeus square with rounded lateral A. inconstans Burmeister, 1847 Anomala sp. margins ...... 9 Strigoderma sulcipennis Burmeister, 1844 Strigoderma tomentosa Bates, 1888 8’ Clypeus trapezoidal ...... 10 3 Dynastinae 9 Dorsal surface with long setae. A) Dynastini Antennal club shorter than rest of Golofa imperialis Thomson, 1858 antennomeres. Pronotum with two B) Oryctini pairs of longitudinal bands and dark Strategus aloeus (Linné, 1758) parallels; each pair with anterior C) Cyclocephalini convergence. Total length 18 mm. Cyclocephala lunulata Burmeister, 1847 ...... Euphoria iridescens C. stictica Burmeister, 1847 C. sexpuctata Laporte, 1840 9’ Dorsal surface with short setae. D) Antennal club much longer than the Tomarus nasutus (Burmeister, 1847) rest of the antennomeres. Pronotum T. sallei (Bates, 1888) with one pair of dark longitudinal Bothynus complanus (Burmeister, 1847) bands in a “db” shape; each pair with 4 Cetoniinae anterior convergence. Total length A) Gymnetini 18 mm ...... Euphoria vestita Cotinis mutabilis (Gory & Percheron, 1833) C. pauperula Burmeister,1847 10 Anterior tibia externally tridentate, Hologymnetis cinerea (Gory & Percheron, 1833) all teeth equidistant and basal tooth B) Cetoniini smaller. Dorsal coloration green, (Gory & Percheron 1833) E. biguttata (Gory & Percheron, 1833) elytra with whitish sculpture. Total E. canescens (Gory & Percheron, 1833) length 21.0-21.5 mm ...... E. dimidiata (Gory & Percheron, 1833) ...... Euphoria westermani E. iridescens Schaum, 1841 E. lineoligera Blanchard, 1850 10’ Anterior tibia externally tridentate, E. pulchella (Gory & Percheron, 1833) E. sepulcralis leucographa (Gory & Percheron, 1833) with two anterior teeth close to- E. subtomentosa Mannerheim, 1837 gether and basal tooth slightly E. vestita (Gory & Percheron, 1833) smaller. Total length 15-18 mm .... E. westermanni (Gory & Percheron, 1833) ...... Euphoria biguttata 194 Revista Colombiana de Entomología Cutberto Pacheco F. & Cols.

11 Basal half of elytra reddish, the rest 21’ Third protarsal joint situated at same 29 Elytra glabrous. Anterior margin of black with or without variable sculp- height as apical tooth of protibia . clypeus straight. Pronotum and ture ...... 12 ...... Anomala forreri elytra with irregular spots ...... Cyclocephala lunulata 11’ Basal half of elytra always lacking 22 Pronotum and scutellum green; reddish spots...... 14 protibia tridentate ...... 29’ Elytra setiferous. Anterior margin of clypeus sinuate. Pronotum and 12 Clypeus almost square; posterior ...... Anomala cincta elytra with a more or less defined half of elytra black and lacking 22’ Pronotum yellowish-brown with a pattern ...... 30 sculpture ..... Euphoria dimidiata blackish-red anterocentral spot vari- 30 Each elytra with three irregular black 12’ Clypeus triangular; posterior half of able in shape; protibia bidentate ...... Anomala inconstans points. Disc of pronotum with two black elytra with sculpture ...... 13 black spots...... 13 Anterior margin of clypeus rounded; 23 Pronotum with a longitudinal fur- ...... Cyclocephala sexpunctata sides of pronotum rounded and with row and two diagonal furrows on 30’ Each elytra with three longitudinal whitish sculpture ...... each side; pronotal setae widely spots: two short lateral spots and one ...... Euphoria canescens spaced ....Strigoderma sulcipennis large spot that widens at the poste- 13’ Anterior margin of clypeus truncate; 23’ Pronotum lacking furrows and rior half ...... Cyclocephala stictica sides of pronotum angled and lack- setiferous, setae abundant and erect 31 Apex of meso- and metatibiae trun- ing whitish sculpture...... Strigoderma tomentosa cate or with dorsolateral projection; ...... Euphoria pulchella 24 Basal margin line of pronotum com- sexual dimorphism rare. Head with 14 Pronotum black ...... 15 plete. Exterior border of mandibles carinae or tubercles . Pentodontini ...... 32 14’ Pronotum reddish ...... 16 clearly indented. Dorsal coloration yellow with green highlights. Total 31’ Apex of metatibiae denticulate or 15 Elytra bicolored, black with yellow length 22-26 mm ...... scalloped. Sexual dimorphism Euphoria basalis ...... Pelidnota virescens accentuated. Males and females 15’ Elytra black with irregular whitish without horns on head, only two 24’ Basal margin line of pronotum in- transversal tubercles. Male prono- sculpture on posterior half and sides. complete or absent ...... 25 Total length 10-11 mm ...... tum with three horns or bumps that ...... Euphoria lineoligera 25 Basal margin of pronotum incom- surround a wide central fovea. Total plete ...... Chrysina macropus length 30-50 mm ...... 16 Anterior margin of clypeus straight, ...... Strategus aloeus projected upward and curved in a 25’ Basal margin line of pronotum ab- 32 Apex of meso- and metatibiae ex- “u” shape; sides of pronotum with sent...... 26 tended dorsolaterally; propygidium whitish sculpture ...... 26 Scutellum longer than pronotum. extended towards the back with a ...... Euphoria leucographa Color shiny black. Metaepisternum stridulatory area. Pronotum with one 16’ Anterior margin of clypeus slightly rugose and punctate. Total length tubercle and a postapical concavity. rounded, never projected upward; 23-29 mm ..... Macraspis aterrima Total length 27-36 mm ...... sides of pronotum lacking whitish 26’ Scutellum shorter than pronotum...... Bothynus complanus sculpture ...... Clypeus semitrapezoidal. Labrum 32’ Apex of meso- and metatibiae trun- ...... Euphoria subtomentosa visible above. Protibia bidentate in cate; propygidium not extended 17 External border of elytra with mem- males, tridentate in females, with backwards and lacking stridulatory branous margin ...... Anomalini very small basal tooth. Dorsal col- area. Pronotum lacking tubercles ...... 18 oration bright metallic green. Total and depressions. Clypeus lacking length 14-16 mm ...... preapical carina; front with transver- 17’ External border of elytra lacking ...... Calomacraspis splendens sal carina...... Tomarus membranous margin ...... 24 ...... 33 27 Protarsus equal to or larger than 18 Elytrae wider than posterior region. protibia. Male pronotum with tuber- 33 Protibia tetradentate with a well- Dorsum convex ...... Anomala 19 cle; head with thin horn ...... formed denticle between the second 18’ Elytrae longer in humeral than in ...... Golofa imperialis and third tooth. Frontal carina posterior region. Dorsum flat and bituberculate. Pygidium smooth, 27’ Protarsus shorter than protibia . 28 glabrous ...... Strigoderma 23 polished, and punctate. Total length 20-21 mm...... Tomarus sallei 19 Pronotum blackish-red ...... 20 28 Head and pronotum without carinae, tubercles, or depressions. Tarsomeres 33’ Protibia tridentate, lacking denticles 19’ Pronotum green or yellowish-brown semicylindrical. Meso- and meta- between teeth. Frontal carina con- ...... 22 tibiae with wide apex ...... tinuous. Pygidium smooth, pol- 20 Elytra red-brown...... Anomala sp. Cyclocephala ...... 29 ished, with three scattered punctures. Total length 17-19 mm ...... 20’ Elytra yellowish-brown ...... 21 28’ Head and pronotum with carinae, tubercles, and depressions. Tarso- ...... Tomarus nasutus 21 Second protarsal joint situated at meres triangular or semicylindrical. 34 Dorsal and ventral region covered same level as apical tooth of protibia Meso- and metatibiae apex, scal- with yellow scale-like setae ...... Anomala foraminosa loped or dentate ...... 31 ...... Isonychus ocellatus Phytophagous Scarabaeidae from Guerrero, Mexico 195

34’ Dorsal region smooth or covered 40’ Pygidium setiferous, setae short and and sharpness of intermediate tooth. with setae of varying lengths ... 35 sparse. Total length 8.0 mm ...... Body length 14 to 19 mm (P. ravida ...... Diplotaxis trapezifera group, P. dentex complex). Color 35 Anterior coxae more or less conical, reddish-chestnut. Total length 15-17 prominent. Length less than 12 mm. 41 Vertex generally with a well-marked mm ...... Phyllophaga ravida Sexual dimorphism rare ...... 37 transversal carina. Claws serrate or pectinate ...... 47’ External claws of male mesotarsus 35’ Anterior coxae transversal. Total .... Phyllophaga (Listrochelus) sp. slightly deformed and lacking distal length generally greater than 12 mm bifurcate appearance, as intermedi- (except in Phyllophaga oblongula, 41’ Vertex lacking transversal carina. All ate tooth is very short and rounded. which measures 8.5 mm). Sexual di- three pairs of claws bifid, grooved, Body length 16 to 23 mm (ravida morphism apparent or quite notice- or dentate, especially in males ...... group, dasypoda complex) ...... 48 able. Reproductive organ complex ...... 42 ...... 36 48 Clypeus semicircular, slightly si- 42 Tarsal claws unidentate ...... nuate. Tegument shiny. Antennal la- 36 Antennal club formed by three Phyllophaga (sensu stricto)..... 46 mella in males as long as the first antennomeres, both in males and fe- 42’ Tarsal claws grooved or bifid .. 43 seven antennomeres combined. males. Sexual dimorphism apparent Pygidium with long setae ...... or quite noticeable . Phyllophaga 43 Ventral region of tarsomeres with ...... P. dasypoda ...... 41 abundant setae (more conspicuous in males). Dorsum glabrous, shiny. 48’ Clypeus with anterior margin si- 36’ Antennal club formed by seven Body elongate and yellow ...... nuate. Tegument shiny. Antennal la- antennomeres in males and five in Phyllophaga (Chlaenobia) scabripyga mella in males shorter than first females ...... Polyphylla petiti seven antennomeres combined. 43’ Ventral region of tarsomeres with Pygidium with short setae. Color 37 Antennal club small and oval. Ab- sparse or no setae. Dorsal appear- reddish-chestnut. Total length 21- dominal sternites totally or partially ance variable ...... Phyllophaga 22.5 mm ...... fused to each other at medial thirds. (Phytalus) ...... 44 ...... Phyllophaga fulviventris Head, pronotum, and ventral regions dark brown, elytra shiny straw-yel- 44 Pronotum opaque. Front with abun- 49 Claws of intermediate denticle as low. Total length 3 mm ...... dant, erect, medium-length setae; long as or longer than apical denti- ...... Chnaunanthus discolor pronotum and elytra setiferous, se- cle; widely separated from both ends, tae short; anterior region of prono- with a slight, rounded basal dilation. 37' Antennal club elongate. Abdominal tum with sparse, long setae in Tropical species ...... 50 sternites never totally or partially anterior region and area in front of fused to each other at medial thirds apical callus of elytra ...... 49’ Claws with intermediate denticle of Diplotaxis ...... 38 ...... P. (Ph.) epulara variable length and position. Both male metatibial spurs articulate with 38 Species longer than 8 mm. Clypeus: 44’ Pronotum shiny ...... 45 apical border...... 51 trapezoidal, anterior margin truncate 45 Elytra and scutellum shiny. Front with rounded lateral angles ..... 39 50 Exterior spur of male metatibiae and anterocentral region of pro- fused to apical border and at least 38’ Small species, length less than 8 mm. notum with short setae, conspicu- 60% shorter than interior spur. Clypeus trapezoidal, anterior mar- ous and/or inconspicuous ...... Parameres short, compact, fused to gin sinuate and sharply elevated ...... P. (Ph.) obsoleta base and apex. Dorsal surface vari- with lateral angles projected. Lateral 45’ Elytra and scutellum opaque. Long able although generally velvety. margins of elytra with long setae .. setae on front and anterocentral re- Aedeagus slightly sclerotized and ...... 40 gion of pronotum ...... lacking complex ornamentation...... 39 Elytra twice as long as pronotum...... P. (Ph.) crenonycha P. (Phyllophaga), P. rorulenta group Second elytral interval unipunctate. 46 Claws with dilated and dentate base, P. martinezpalaciosi Lateral margins of pronotum near intermediate tooth flanked by nar- apex not strongly marginate or el- row, deep indentations. External 50’ Exterior spur of male metatibiae ar- evated. Color black with red high- claws of male mesotarsus with api- ticulate with apical border, length lights. Total length 9 mm ...... cal portion curved or angled down- greater than 50% that of interior spur...... Diplotaxis cribriceps ward so that intermediate tooth juts Dorsal and pygidial dressing velvety or setiferous. Dorsal and pygidial 39’ Elytra three times longer than out laterally causing deformationP. (Phyllophaga) ravida group .....47 surface formed by short, very abun- pronotum. Second elytral interval dant setae. Male and female claws multipunctate. Lateral margins of 46’ Claws with intermediate tooth far similar. Parameres short, wide, fused pronotum near apex strongly margi- from apex or base or from both ends. at base and apex. Aedeagus long, nate and slightly elevated. Color Claws with intermediate denticle sclerotized, with thick, very con- reddish black. Total length 9 mm . variable in structure and position, spicuous setiferous ornaments...... Diplotaxis atramentaria inferior border seldom serrate ...49 ...... P. (Phyllophaga) setidorsis 40 Pygidium setiferous, setae long and 47 External claws of male mesotarsus group...... 53 abundant. Total length 7.5-8.2 mm deformed with a distal bifurcate ap- 51 Male anal plate with anterior flange ...... Diplotaxis megapleura pearance due to their great length that reaches lateral ends, middle sec- 196 Revista Colombiana de Entomología Cutberto Pacheco F. & Cols.

tion indented, one sinus or bilobed eight states at an altitude of 300 to 1.650 May (43), June (33), July (2), and Octo- or bidentate projection. Frequently m (Deloya et al. 1995). The only P. petiti ber (3) in Mochitlán (3), Atlixtac (27), fifth sternite visible with dark, sample was captured at lights in Tixtla and Chilpancingo (57). grainy medial area. Dorsum pruinose in December in an area near croplands. Strigoderma Burmeister. The 8 speci- or setiferous dorsum. Parameres Isonychus Mannerheim. Isonychus oce- mens captured correspond to two species, short, fused, ring-like, with a small llatus is a common species of wide distri- S. sulcipennis (6) and S. tomentosa (2) ventral bidentate projection and bution; its food preferences are unknown and were collected in Mochitlán (3), symmetrical latero-distal denticles. (Morón et al. 1997). In May, the 3 speci- Chilpancingo (1), and Tixtla (4) during Aedeagus highly ornamented, with mens were captured at lights in Atlixtac September (2), October (3), November thorns, setae, and curved filaments (2) and Tixtla (1). (1), and December (1). Strigoderma P. (Phyllophaga) anodentata sulcipennis was captured on Tagetes group...... 52 Chnaunanthus Burmeister. Chnaunan- erecta (Cempanzuchitl) flowers and S. thus discolor is a very common species 51’ Male anal plate narrow and slightly tomentosa on Compositae flowers. in central Mexico (Morón et al. 1997). excavated, but lacking notable The four samples reviewed were captured Golofa Hope. Golofa imperialis lives at flange in anterior margin. Parameres in Compositae blossoms during October an altitude of 500-2.100 m on the inter- short, fused, ring-shaped. Dorsal re- in Chilpancingo (3) and Atlixtac (1). nal slopes of principal mountains and gion with velvety or setiferous cover. mesetas or woody high plateaus (Morón Aedeagus sclerotized, highly orna- Calomacraspis Bates. Calomacraspis 1993). The only specimen was captured mented with plates and groups of splendens has been found on verbena and at lights in Tixtla during October. macroscopic setae ...... asclepiadaceous flowers; larvae develop (Phyllophaga), P. porodera group on detritus deposits of the Atta Strategus Kirby. Strategus aloeus has P. eniba mexicana (Fr. Smith) (Deloya 1988). The wide neotropical distribution (Morón et species is exclusive to Mexico (Jameson al. 1988). The 16 specimens were col- 52 Antennae formed by 10 antenno- et al. 1994) and lives in warm parts of lected at light (15) and on rotting wood meres. Dorsum opaque .. P. ardara Jalisco, Puebla, Veracruz, Chiapas, (1) during May (7), June (1), July (4), and 52’ Antennae formed by 9 antennome- Morelos, Hidalgo, and Guerrero (Deloya August (4) in Mochitlán (7), Tixtla (2), res. Dorsum shiny. Total length 15- et al. 1994, 1995; Morón, 1994). The only Atlixtac (2), and Chilpancingo (5). 16 mm ...... P. brevidens specimen in this study was captured on Cyclocephala Dejean. This has a Compositae plants during October in 53 Superior metatibial spur curved in wide neotropical distribution. The 168 Chilpancingo. an open “s” shape. 3-6 antennal specimens represent three species: C. antennomeres of equal length, 7 Macraspis MacLeay. Macraspis ate- lunulata (166), C. stictica (1), and C. shorter ...... P. crinalis rrima is a species exclusive to Mexico; sexpunctata (1), all of which were cap- adults consume annona fruits, and larvae tured at lights during the months of May 53’ Superior metatibial spur curved. 3- develop on rotten Persea americana (113), June (46), September (1), and Oc- 4 antennomeres of equal length, 5-7 trunks (Deloya et al. 1995). The only tober (8) in Mochitlán (3), Atlixtac (15), shorter ...... P. setifera specimen reviewed was captured in flight Chilpancingo (148), and Tixtla (2). during September in Mochitlán. General comments about Tomarus Erichson. The 9 specimens rep- phytophagous Scarabaeidae found in Pelidnota MacLeay. Pelidnota virescens resent 2 species: T. sallei (8) and T. nasutus the Central Region of the State of adults exhibit nocturnal habits and feed (1) and were collected at lights lighting Guerrero, Mexico on the foliage of various trees. Larvae are during the months of May (6), July (2), Phyllophaga Harris. Adults exhibit crep- found inside rotten stumps, and the life and August (1) in Chilpancingo (6) and uscular or nocturnal habits and feed on cycle is complete in one year; they are Tixtla (3). widely distributed between Mexico and the foliage of various plants; larvae, in Bothynus Hope. Bothynus complanus, 5 Costa Rica (Deloya et al. 1995; Morón contrast, eat roots (Deloya et al. 1995; specimens were captured at lights during et al. 1997). The only specimen collected Morón et al. 1988). 293 specimens from May (3) and November (2) in Tixtla. 17 species were captured at lights in was captured at lights in Mochitlán dur- Mochitlán (9), Atlixtac (172), Chilpan- ing May. Cotinis Burmeister. It has a wide distri- bution from northern South America to cingo (84), and Tixtla (28) during May Chrysina Kirby. Chrysina macropus the southern United States. Most species (130), June (75), July (1), August (1), and adults are frequently attracted to various are diurnal and favor flowers and ripe September (2). types of light. Their alimentary prefer- fruits (Deloya et al. 1995). A total of 75 ences are unknown; larvae have been Diplotaxis Kirby. Adults exhibit crepus- samples from 2 species were collected: collected from rotten stumps. This spe- cular or nocturnal habits; larvae consume C. mutabilis (65) and C. pauperula (10) cies life cycle requires two years (Morón roots (Deloya et al. 1995). 39 samples during January (5), May (14), June (23), et al. 1997). The two specimens studied from 4 species were captured at lights, D. July (2), September (6), October (9), No- were captured at lights in Atlixtac dur- megapleura (4), D. cribriceps (2), D. vember (7), and December (9) in Mo- ing May. atramentaria (30), and D. trapezifera (3) chitlán (24), Atlixtac (2), Chilpancingo in Mochitlán (1), Atlixtac (37), and Tixtla Anomala Samouelle. 87 specimens from (34), and Tixtla (15). Cotinis mutabilis (1) during May (20) and June (19). 5 species were captured at lights: A was collected from Tagetes erecta flow- Polyphylla Harris. The genus is widely forreri (5), A. cincta (2), A. foraminosa ers and Pithecelobium dulce foliage as distributed from Canada to Guatemala. (1), A. inconstans (48), and one undeter- well in flight and from fermented fruit In Mexico, captures have been made in mined species (31) during March (2), traps. Phytophagous Scarabaeidae from Guerrero, Mexico 197

Hologymnetis Martínez. Hologymnetis Table 3. Phytophagous Scarabaeidae (Melolonthinae, Rutelinae, Dynastinae, Cetoniinae) species cinerea is found in Guatemala and widely captured by region in Central Guerrero, Mexico. (Numbers correspond to abundance). distributed throughout the Mexican ter- ritory, except on the Baja California pe- Species/ Mochitlan Atlixtac Chilpancingo Tixtla ninsula and in Yucatán (Deloya et al. Altitude 840 m 1210 m 1360 m 1600m 1995). The 9 samples were captured in fermented fruit traps and on Tagetes Cotinis mutabilis 22 2 27 14 erecta flowers during the months of Sep- C. pauperula 2 0 71 tember (1), October (5), and November Hologymnetis cinerea 1 2 4 2 (3) in Mochitlán (1), Atlixtac (2), Chipan- Euphoria basalis 0 0 1 12 cingo (4), and Tixtla (2). E. s. leucographa 8 4 3 2 E. dimidiata 3 1 7 1 Euphoria Burmeister. It is widely distrib- E. iridescens 0 43 0 0 uted across most of the American conti- E. canescens 0 3 0 0 nent. Adults favor flowers, while larvae are E. vestita 0 0 0 1 saprophagous (Morón et al. 1988; Deloya E. biguttata 15 8 0 0 et al. 1995). The 585 samples collected represent 11 species: E. basalis (112), E. E. subtomentosa 67 247 16 35 leucographa (17), E. dimidiata (12), E. E. lineoligera 2 1 0 0 iridescens (43), E. canescens (3), E. vestita E. pulchella 0 2 0 0 (1), E. biguttata (23), E. subtomenstosa E. westermanni 1 0 2 1 (365), E. lineoligera (3), E. pulchela (2), Macraspis aterrima 1 0 0 0 and E. westermanni (4), all captured in Calomacraspis splendens 0 0 1 0 fermented fruit traps and on pumpkin flow- Strigoderma sulcipennis 2 0 0 4 ers (Cucurbita pepo), campanzuchitl S. tomentosa 1 0 1 0 (Tagetes erecta), huizache (Acacia Anomala forreri 0 2 3 0 schaffneri) and other Compositae in A. cincta 0 0 2 0 Mochitlán (96), Atlixtac (309), Chilpan- A. foraminosa 0 0 1 0 cingo (140), and Tixtla (40) during Janu- A. inconstans 3 25 20 0 ary (8), May (5), June (3), September (3), Anomala sp. 0 0 31 0 October (460), November (95), and De- Pelidnota virescens 1 0 0 0 cember (11). Chrysina macropus 0 2 0 0 Strategus aloeus 7 2 5 2 Discussion Cyclocephala lunulata 3 15 148 0 C. stictica 0 0 0 1 With regard to specific richness in each C. sexpunctata 0 0 0 1 region, 22 species were captured in Golofa imperialis 0 0 0 1 Mochitlán, 34 in Atlixtac, 32 in Chilpan- Tomarus nasutus 0 0 1 0 cingo, and 23 in Tixtla (Table 3; Fig. 2); T. sallei 0 0 5 3 43.65% of the specimens were obtained Bothynus complanus 0 0 0 5 in Atlixtac, 36.92% in Chilpancingo, Chnaunanthus discolor 0 1 3 0 11.39% in Mochitlán, and only 8.02% in Polyphylla petiti 0 0 0 1 Tixtla. Of the 57 species studied, C. Isonychus ocellatus 0 2 0 1 mutabilis, H. cinerea, E. leucographa, E. Diplotaxis megapleura 1 3 0 0 dimidiata, E. subtomentosa, S. aloeus, P. D. cribriceps 0 2 0 0 ardara and P. brevidens were found in D. atramentaria 0 30 0 0 the four regions at an altitude of 840- D. trapezifera 0 2 0 1 1.600 meters. The similarity index (Søren- Phyllophaga ardara 1 5 614 sen 1948) between the localities is as P. dasypoda 0 0 04 follows: Mochitlán-Atlixtac 53%, Mo- P. crinalis 0 1 146 chitlán-Chilpancingo 55%, Mochitlán- P. setifera 0 2 5 0 Tixtla 48%, Atlixtac-Chilpancingo 60%, P. integriceps 0 4 180 Atlixtac-Tixtla 38% and Chilpancingo- P. eniba 0 2 20 Tixtla 47%; this indicates a greater simi- P. disca 0 0 10 larity between Atlixtac and Chilpancingo, P. brevidens 1 4 102 intermediate regions found at altitudes of P. ravida 0 7 40 1.210 and 1.360 m, respectively. P. fulviventris 0 0 132 The vegetation of Balsas River Basin and P. martinezpalaciosi 0 3 7 0 the Mexican Pacific slope is mainly de- Phyllophaga sp. 0 2 0 0 ciduous tropical forest, xerophyllous P. epulara 1 0 0 0 scrub, and lowland oak forest mixed with P. crenonycha 0 23 0 0 pine forest at intermediate altitudes. As P. obsoleta 0 80 0 0 such, studies of Scarabaeidae Pleurosticti Phyllophaga sp. 1 1 3 0 were carried out in the south of Morelos P. scabripyga 5 38 1 0 198 Revista Colombiana de Entomología Cutberto Pacheco F. & Cols.

phagous (28%), grouping Calomacraspis, *R(( -( Cotinis, Hologymnetis and Euphoria; c)

*Q(( ,) phylo-xilophagous (3.5%), represented by Macraspis aterrima and Pelidnota vires- *-(( ,( cens, and d) sapro-rizo-xilophagous *+(( (3.5%), which included Golofa imperialis +) and Chrysina macropus. Feeding habits *((( +( are unknown for adults and larvae of the R(( species Isonychus ocellatus and Bothynus /D@Y A [

*) 0 ??@?  Altitude (masl) complanus. Q(( *( The structure of the trophic guilds in cen- -(( tral Guerrero suggests that there is a pre- +(( ) dominance of phylo-rhizophages during spring owing to the reproductive habits of ( ( 5 ? /A? U A ?  WA the Melolonthinae: larvae must feed and X ?A make it to the third larval instar as quickly as possible because the rainy season – Figura 2. Relation between altitude (bars) and specific richness (line) by locality of the when vegetation renews its foliage and phytophagous Scarabaeidae (Melolonthinae, Rutelinae, Dynastinae Cetoniinae) from the Cen- tral Region of Guerrero, Mexico (July 1999-June 2000). roots are produced – is very short. Sapro- meliphages predominate during autumn when the Asteraceae are flowering and and Tepic, Nayarit between 800 and 1.200 sites located between 850 and 1.850 masl, when the cellulose based material that the masl, where there is deciduous tropical and relatively high compared to Tentzo, larvae consume has accumulated (decom- forest. Species richness was 72 and 78 spe- Puebla. Results in central Guerrero suggest posing litter and wood) on the soil. The cies in these two locales, respectively. In that as altitude increases (840 to 1.360 limited representation of the phylo- Cuernavaca, Morelos (1.250-1.850 masl) masl) species richness increases (Mochi- rhizophages and the sapro-meliphages for tropical deciduous forest and Pinus- tlán840m S = 22, Atlixtac1210 m S = 34, could be an indication that small scale Quercus forest there were 93 species. In Chilpancingo1360 m S = 32), up to 1.600 removal of wood from the forest could be Tentzo, Puebla (2.000-2.350 masl) in masl in Tixtla where it decreases (S = 23). contributing to the extinction of these xerophyll scrub and oak forest there were populations on a local scale. only 32 species (Deloya et al. 1993,1995; Based on the feeding habits of Scarab lar- Morón et al. 1998, 2000). Considering the vae and adults in the regions studied, the New Records. Of the total species cap- altitudinal ranges (350, 400 and 600 masl) following guilds were represented: a) tured (N=57), 30% (17) represent new and the species richness recorded, in the phylo-rizophagous (61.4%), covering records for Guerrero; they correspond to central region of Guerrero over an alti- Phyllophaga, Diplotaxis, Polyphylla, the genera Anomala, Strigoderma, Eu- tudinal range of 760 masl, 57 species were Anomala, Strigoderma, Strategus, Cyclo- phoria, Phyllophaga, Polyphylla and recorded; a relatively low number for the cephala and Tomarus; b) sapro-meli- Chnaunanthus (Table 4), making the to-

Table 4. New records of phytophagous Scarabaeidae species for the State of Guerrero, Mexico (Melolonthinae, Rutelinae, Dynastinae, Cetoniinae).

Species Distribution New records for Guerrero

Phyllophaga ardara Hgo Mochitlan, Atlixtac, Chilpancingo, Tixtla P. dasypoda Chis,Oax,Pue,Ver Tixtla P. crinalis Hgo,Pue Atlixtac, Chilpancingo, Tixtla P. setifera Hgo, Jal,Mex,Nay, Oax,Pue,Sin Atlixtac, Chilpancingo P. eniba Mex,Nay,Sin,Son Chilpancingo P. disca Mex Chilpancingo P. brevidens Jal, Mex,Mor,Nay, Pue,Sin Mochitlan, Atlixtac, Chilpancingo, Tixtla P. fulviventris Chis,Col,Jal,Mich, Mor,Nay Chilpancingo, Tixtla P. crenonycha Mex Atlixtac P scabripyga Oax Mochitlan, Atlixtac, Chilpancingo Isonychus ocellatus Sin,Hgo,Oax,Pue,Ver Atlixtac, Tixtla Polyphylla petiti Mor,Pue Tixtla Chnaunanthus discolor Gto,Mich,Oax,Pue Atlixtac, Chilpancingo Anomala forreri Sin Atlixtac, Chilpancingo A. foraminosa BCS,Chis,Hgo,Mor, Oax,Pue,QRo,Sin,Tab, Tamps,Ver Chilpancingo Strigoderma tomentosa Oax Mochitlan, Chilpancingo Euphoria canescens Ags,Gto,Pue,Ver Atlixtac

Ags (Aguscalientes), BCS (Baja California Sur), Chis (Chiapas), Col (Colima), Gto (Guanajuato), Hgo (Hidalgo), Jal (Jalisco), Mex (México), Mich (Michoacán), Mor (Morelos), Nay (Nayarit), Oax (Oaxaca), Pue (Puebla), QRo (Quintana Roo), Sin (Sinaloa), Son (Sonora), Tab (Tabasco), Tamps (Tamaulipas), Ver (Veracruz). Phytophagous Scarabaeidae from Guerrero, Mexico 199 tal present number of phytophagous cal history in the Balsas River Basin. This coleópteros edafícolas americanos. Pu- scarab species 137. is in contrast to Chamela and Tepic which blicación especial de la Benemérita are farther away from the central region Universidad Autónoma de Puebla y la Agricultural aspects. In the central part of Guerrero at 577 km and 692 km, re- Sociedad Mexicana de Entomología. of Guerrero, the so-called “wire worms” spectively. Although Tentzo is located at GARCÍA, E. 1988. Modificaciones al sistema (Coleoptera: Elateridae) and “white de clasificación climática de Koppen, an intermediate distance (307 km), its al- grubs” (Coleoptera: Scarabaeidae) are México. p. 113-117. titude and cold climate could be a cause serious agricultural pests, having caused JAMESON, M. L.; RACTCLIFFE, B. C.; of the low similarity and species richness. great economic losses to basic crops in MORÓN, M. A. 1994. A synopsis of the The other sites are located at an altitude most temporal zones (Morón 1999, 1988; neotropical genus Calomacraspis Bates of 1.200 masl, with the exception of with a key to larvae of the American gen- Cortés et al. 1994). Species of Phyllo- Chamela at 100 masl, which has a humid era of Rutelini (Coleoptera: Scarabaeidae: phaga, Diplotaxis, Anomala, Cycloce- tropical climate. Rutelinae) Annals of the Entomological phala, Strategus and Tomarus could also Society of America 87 (1): 43-58. be considered potential pests, given that Acknowledgments MORÓN, M. A. 1984. Escarabajos, 200 in the phylo-rhizophagous guild, there millones de años de evolución, Publ. 14, are species with rhizophagous, sapropha- We are grateful to Miguel Angel Morón Instituto de Ecología, México. 132 p. gous or facultative larvae, as observed and Leonardo Delgado for determinations MORÓN, M. A. 1988. Las especies de for Cyclocephala lunulata. This species of Phyllophaga and Diplotaxis species Phyllophaga (Coleoptera: Melolonthidae) behaves like a saprophage in soils with respectively. This work is a contribution con mayor importancia agrícola en México. a high content of decomposing organic to the project “Systematics and ecology p. 81-102. En: III Mesa Redonda sobre of phytophagous and saprophagous in- plagas del suelo. Sociedad Mexicana de material, and eats the remains of har- Entomología, Morelia, Michoacán, México. vested rice and sugar cane crops, but has sects” of the Department of Entomology (902-08/044), Instituto de Ecología, A.C. MORÓN, M. A.; DELOYA, C.; DELGADO, been observed eating roots in sandy soils L. 1988. Fauna de coleópteros Melolon- with little organic material (Deloya References cited thidae, Scarabaeidae y Trogidae de la región 1998). de Chamela, Jalisco, México. Folia CORTÉS GENCHI, P.; MARURIS, M. R.; Entomológica Mexicana. 77: 313-378. Zoogeographical aspects. The fauna ACEVEDO ROSAS, J. L. 1994. Pruebas MORÓN, M. A. 1997. Inventarios faunísticos obtained from the central region of en campo con Metarhizium anisopliae para de los Coleoptera Melolonthidae neotro- Guerrero makes up 5.6% of all phytopha- el control de la «gallina ciega» en el cultivo picales con potencial como bioindicadores. gous scarabs recorded in the country and del maíz. p. 137. En: 1er. Encuentro de Giornale Italiano di Entomologia 8: 265- 13.47% of all species recorded in the Investigación y Educación Agropecuaria y 274. state. Melolonthinae (24) and Ceto- Forestal en Guerrero. MORÓN, M. A.; RATCLIFFE, B. C.; niinae (14) are the subfamilies with the DELOYA, C. 1988. Coleópteros Lamelicornios DELOYA, C. 1997. Atlas de los escara- greatest specific richness; Phyllophaga asociados a depósitos de detritos de Atta bajos de México, Coleoptera: Lame- and Euphoria are the most diverse gen- mexicana (Smith) (Hymenoptera: Formi- llicornia, Vol. I, Familia Melolonthidae. era and include 66.66% of species. In cidae) en el sur del estado de Morelos, Sociedad Mexicana de Entomología, México. Folia Entomológica Mexicana 75: México, México. 274 p. terms of genera, six are in the subfamily 77-91. MORÓN, M. A.; DELOYA, C.; RAMÍREZ, Rutelinae, five each in Dynastinae and DELOYA, C.; A. BURGOS, A.; BLACKA- A. C.; HERNÁNDEZ, S. R. 1998. Fauna Melolonthinae, and only three in Ceto- LLER, J.; LOBO, J. M. 1993. Los coleóp- de Coleoptera Lamellicornia de la región niinae. The composition of the fauna teros Lamelicornios de Cuernavaca, de Tepic Nayarit. Acta Zoológica Mexicana observed was compared in that of five Morelos, México (Passalidae, Trogidae, (n.s.) 75: 73-116. different regions previously studied: Scarabaeidae y Melolonthidae). Boletín MORÓN, M. A. 1999. Coleoptera Melo- Cuernavaca, southern Morelos, Sierra Sociedad Veracruzana de Zoología 3 (1): lonthidae. p. 43-59. En: Deloya López, A. del Tentzo, Puebla in the Balsas Basin 15-55. C.; Valenzuela González, J. E. (eds.). and Chamela, Jalisco, and Tepic, Nayarit DELOYA, C.; MORÓN, M. A. 1994. Listados Catálogo de insectos y ácaros plagas de on the Mexican Pacific slopes. Similar- faunísticos de México, V. Coleópteros los cultivos agrícolas de México. Sociedad Lamelicornios del Distrito de Jojutla, Mexicana de Entomología, Publicaciones ity (Sørensen 1948) was greatest between Morelos, México (Melolonthidae, Scara- Especiales No. 1, México. southern Morelos (QS = 0.46) and baeidae, Trogidae y Passalidae). Instituto MORÓN, M. A.; ARAGÓN, A.; TAPIA- Cuernavaca, Morelos (QS = 0.38; Morón de Biología, UNAM, México. 49 p. ROJAS, A. M.; ROJAS-GARCÍA, R. et al. 1988, 1998, 2000; Deloya et al. DELOYA, C.; MORÓN, M. A.; LOBO, J. M. 2000. Coleoptera Lamellicornia de la Si- 1993), compared with Chamela, Jalisco 1995. Coleoptera Lamellicornia (Macleay, erra del Tentzo, Puebla, México. Acta (QS = 0.30), Tepic, Nayarit (QS = 0.29) 1819) del Sur del Estado de Morelos, Zoológica Mexicana (n.s.) 79: 77-102. and Tentzo, Puebla (QS = 0.23). Geo- México. Acta Zoológica Mexicana (n.s.) SØRENSEN, T. 1948. A method of establish- graphic distance could influence the 65: 1-42. ing groups of equal amplitude in plant so- similarity between the fauna of central DELOYA, C. 1998. Cyclocephala lunulata ciology based on similarity of species Guerrero and that of southern Morelos Burmeister, 1847 (Coleoptera: Melolon- content and its application to analysis of thidae, Dynastinae) asociada al cultivo de the vegetation on Danish commons. and Cuernavaca. The latter are the two maíz en Pueblo Nuevo, Morelos, México. Biologiske Skrifter 5: 1-34. closest localities, at 123 km and 169 km p. 121-130. En: Morón M.A.; Aragón A. from the central region of Guerrero and (eds.). Avances en el estudio de la diver- they also share a common biogeographi- sidad, importancia y manejo de los Recibido: 18-oct-05 • Aceptado: 12-abr-06