Journal Information
Vol. 60. Issue 1.
Pages 1-7 (January - March 2016)
Share
Share
Download PDF
More article options
Visits
...
Vol. 60. Issue 1.
Pages 1-7 (January - March 2016)
Systematics, Morphology and Biogeography
DOI: 10.1016/j.rbe.2015.11.012
Open Access
Description of immature stages of Hoplia mexicana Harold and H. squamifera Burmeister (Coleoptera, Melolonthidae, Hopliinae)
Visits
...
Miguel Ángel Moróna,
Corresponding author
miguel.moron@inecol.mx

Corresponding author.
, Andrés Ramírez-Ponceb, Concepción Ramírez-Salinasc, Hortensia Carrillo-Ruizd
a Red de Biodiversidad y Sistemática, Instituto de Ecología, Xalapa, Mexico
b Laboratorio de Biología Evolutiva, Centro Tlaxcala de Biología de la Conducta, Universidad Autónoma de Tlaxcala, Tlaxcala, Mexico
c El Colegio de la Frontera Sur (ECOSUR), San Cristóbal de Las Casas, Chiapas, Mexico
d Laboratorio de Entomología, Escuela de Biología, Benemérita Universidad Autónoma de Puebla, Puebla, Mexico
Article information
Abstract
Full Text
Bibliography
Download PDF
Statistics
Figures (6)
Show moreShow less
Tables (1)
Table 1. Main characters in the pupae of some American genera of Scarabaeoidea. D.O., dioneiform organs; H.P., humeral projections; P.P., pronotal projections; T.P., tergal projections; U, urogomphi.
Abstract

Third stage larvae and pupae are described based on specimens collected in Mexico: Oaxaca (Cerro Zempoaltepetl), and Chiapas (Amatenango), respectively. Pupal characters are described for the first time for American Hopliinae. Habitus images and figures of diagnostic characters as well as comments on the differences between these larvae and those of Hoplia callipyge LeConte, 1856 and H. equina LeConte, 1880, the only Hopliinae larvae previously known in New World, are also included.

Keywords:
Biology
Morphology
Pupae
Taxonomy
White grubs
Full Text
Introduction

The immature stages of world Hopliinae are poorly known. Partial descriptions exist for the third instar larvae of the North American species Hoplia callipyge LeConte, 1856 (= H. oregona LeConte, 1856; = H. cazieri Boyer, 1940) and H. equina LeConte (Ritcher, 1949). Additionally, there are larval descriptions for some Russian species of the genus Hoplia Illiger, 1803 and the central European H. philanthus (Fuessly, 1775) (Medvedev, 1952; Hůrka and Bílý, 1966). Micó (2001) redescribed the larva of H. philanthus and described that of H. chlorophana Erichson, 1848, based on specimens from Spain, both with detailed illustrations. Ansari et al. (2006, 2008) detailed the life cycle of H. philanthus and commented about the importance and biological regulation of this turf pest.

Larvae of North American species of Hoplia may be characterized as follows: head without stemmata; labrum symmetrical; maxillary stridulatory area with row of 9–11 short teeth with anteriorly directed points; lacinia with longitudinal row of 3 unci, distal 2 unci fused basally, subapical uncus much smaller; epipharynx with plegmatia, plegmata quite short; proplegmatia present or absent; epizygum absent; haptomerum with single helus; dexiophoba and laeophoba absent; haptolachus without microsensilla; spiracles of abdominal segments 1–3 similar in size, spiracles of segments 4–8 much smaller; anal opening Y-shaped; raster consisting of subtriangular teges of 30–60 fairly long hamate setae with curved tips; claws of fore legs large and falcate, claws of middle legs much reduced in size, claws of hind legs minute or absent (Ritcher, 1966). Larvae of European species of Hoplia share all of the above cited characters, except: maxillary stridulatory area with row of 12–14 short teeth with anteriorly directed points; lacinia with subapical uncus large; epipharynx without proplegmatia; spiracles of abdominal segments 4–6 diminutive (Micó, 2001).

During recent years, we had the opportunity to rear many scarab larvae collected from forest soil, grasslands and corn fields of southeastern Mexico, and some of them belong to species of Hoplia. This material allowed us to describe in the present paper the third instar larvae for H. mexicanaHarold, 1869 and H. squamiferaBurmeister, 1844, for which the immature stages were unknown. We also describe the pupae of these two species, a life stage that was unknown for any New World hopliine scarab.

Material and methods

Throughout the descriptions and discussions, we follow terminology employed by Ritcher (1966) and Morón (1986). Drawings of diagnostic structures were made using a Leica stereomicroscope associated with a camera lucida at 0.8–5.0× magnification. Measurements were taken using an ocular micrometer (Leica microsystems, Germany). Voucher specimens were deposited at Colección proyecto Diversidad en Sistemas de Cultivos, El Colegio de la Frontera Sur (ECOSUR), and Colección Entomológica, Instituto de Ecología, A. C. Xalapa, México (IEXA).

TaxonomyDescriptionsHoplia mexicanaHarold, 1869Third instar (Figs. 1–11 and 13)

Description. Head (Fig. 2). Maximum width of head capsule: 1.8–2.0mm. Surface of cranium smooth, with scattered minute punctures, pale yellowish. Frontal and epicranial sutures hidden. Frons with 2 exterior frontal setae, 3–4 posterior frontal setae, 1 anterior frontal seta, 1 anterior angle seta, 3 dorsoepicranial setae, 8–9 epicranial setae on each side and 7–9 setae behind each antennal support. Clypeus with 2 central setae and 4 lateral seta on each side. Labrum nearly symmetrical with rounded lateral margins irregularly bordered, anterior margin projected forward with irregular border, 6–7 posterior setae, 1 central seta and 3–4 lateral setae on each side. Stemmata absent. Epipharynx (Fig. 5) 1.0–1.1mm wide, 0.8–0.9mm long, epizygum absent and zygum as a irregular, ovate, reddish yellow plate. Haptomerum weakly raised, with 4 heli; each plegmatia formed by 8–10 short plegmata; proplegmatia absent. Each acanthoparia with 10–12 spinose setae. Dexiophoba and laeophoba absent. Dexiotorma narrowed, slightly sinuose. Laeotorma elongate and narrowed, with sclerotized plate unciform, raised. Sense cone short. Crepis weakly defined. Left chaetoparia with 28–30 spinose and medium size setae mixed. Right chaetoparia with 30–36 short, stout setae. Mandibles (Figs. 3 and 4) without ventral stridulatory area; scissorial area in both mandibles without distal blade. Right mandible with 1 small preapical tooth, inner margin simply curved, and distal lobe (M1) of molar area scarcely developed. Calx short. Left mandible with 1 small preapical tooth, inner margin simply curved, brustia moderately setose. Acia absent. Maxillae (Figs. 6–9). Mala with 1 uncus on apex of galea, 3 unci surrounded by 3–4 stout heli on apex of lacinia; stridulatory area with 8 small teeth with anteriorly directed points. Hypopharyngeal sclerome (Fig. 10) asymmetrical with raised and curved projection on the right side. Fourth antennomere elongate, with distal half narrowed, with 1 ovate, dorsal sensory spot on basal half and 2 ventral sensory spots.

Figs. 1–5.

Third instar larva of Hoplia mexicana. 1, lateral habitus. 2, head, frontal view. Mandibles, ventral view: 3, left. 4, right. 5, epipharynx.

(0.39MB).
Figs. 6–12.

Hoplia mexicana. 6–11: third instar larva. Maxillae: dorsal view: 6, left. 7, right. Mesial view: 8, left. 9, right. 10, labium and hypopharynx. 11, raster. 12, MALE pupa, lateral view.

(0.29MB).
Figs. 13–16.

Hoplia mexicana. 13, Third instar, lateral view. 14–16: male pupa. 14, ventral view. 15, lateral view. 16, dorsal view.

(0.26MB).

Thorax. Respiratory plates light yellow, scarcely curved, “C” shaped (Figs. 1 and 13) 0.12–0.14mm long, 0.10–0.11mm wide, bulla small, slightly raised, rounded, distance between lobes longer than dorso-ventral diameter of bulla; microscopic holes of respiratory plate elongate-oval in outline and arranged in irregular transverse rows. Lateral sclerome of pronotum not defined. Pro-prescutum with 20–22 long setae regularly distributed; proscutum with 8–9 slender long setae; mesoprescutum with transverse row of 10–12 long setae; mesoscutum with 12–16 setae; mesoscutellum with row of 8–10 slender long setae; metaprescutum with transverse row of 12–16 mixed short and long setae; metascutum with 10–12 setae; metascutellum with transverse row of 10–12 mixed short and long setae (Figs. 1 and 13). Protarsal claws long, narrowed, sharply pointed, with 1 prebasal inner seta and 1 lateral external seta, moderately longer than mesotarsal claws; meso-tarsal claws long, sharply pointed, with pre-basal and lateral setae; metatarsal claw short, rounded, with 2 setae, much shorter than mesotarsal claws. Apical metatarsomere shortened, rounded, with ventral side clearly convex.

Abdomen. Respiratory plates light yellow, slightly curved, “C” shaped; on segments I–IV with similar diameter, 0.12–0.13mm long, 0.10mm wide; plates on segments V–VII are slightly smaller, 0.10–0.11mm length, 0.09mm width. Plate on segment VIII is smaller than preceding. Dorsa of abdominal segments I–VI each with dense vestiture of yellow stout, short setae; dorsa of segments VII–IX with transverse rows of 20–28 slender, short setae. Venter of abdominal segments I–VIII with transverse rows of 14–18 slender, long setae; venter of segment IX with transverse rows of 12–16 mixed short and long setae.

Raster (Fig. 11) with each palidia formed by 9–11 pali, convergent toward basal and distal extremes, septula wide, oval; tegilla with 20–26 short, stout setae; campus with 14–18 slender, short setae; barbula much more densely setose. Dorsal anal lip with dense vestiture of short setae. Ventral anal lip with scattered short setae toward sides and distal border narrowly notched at middle. Anal slit “Y” shaped. Approximate dorsal body length 16–17mm.

Description: body length 7.5–7.8mm. Widest width 4.4–4.5mm. Head. Surface glabrous, strongly deflexed; frons convex with two rounded prominences; clypeus widely concave on the center; labrum, mandibles, maxillae and palps discernible; antennal theca briefly expanded, stout with apex rounded; eyes small (Fig. 14). Thorax: surface glabrous. Pronotum convex, surface slightly irregular, anterior angles prominent, posterior angles rounded. Meso- and metanotum well-differentiated. Elytral and posterior wing thecae closely appressed, curved ventrally around the body; elytral thecae with irregular depressions and large, prominent, humeral tubercles (Figs. 12 and 15); thecae of the wings slightly longer than elytral thecae. Protibiae with apical and preapical tubercles clearly developed. Meso and metatibiae with apical tubercle. Abdomen: segments I–VI clearly wider and shorter than the distal segments VII–VIII, without dioneiform organs, but segments II–IV with pairs of prominent tubercles and segment V with increased posterior border (Figs. 12 and 15–16). Pleural lobes rounded. Spiracle I elongate, with fine peritreme and covered by wing thecae; spiracles II–IV ovate and high, prominent, with narrow sclerotized peritreme; spiracles V–VIII closed. Sutures between segments VIII and X not complete, partially fused. Last segment slightly pruinose, without urogomphi.

Last abdominal segment of male with large dorsal and lateral rounded prominences, and genital ampulla ventrally exposed (Figs. 12 and 14–16); female without such prominences at the apex of abdomen.

Specimens examined. Four third instar larvae, four pupae and two exuviae of third instar larvae reared to adults, collected at Mexico: Oaxaca: Santa María Tlahuitoltepec municipality, Cerro Zempoaltepetl, 2400m, 8-VI-2010, A. Ramírez col. (IEXA).

Comments. The third instar larvae of H. mexicana resemble those of H. callipyge and H. equina, but H. mexicana has 4 heli on haptomerum, and palidia with well-defined septula. Hoplia callipyge and H. equina each have1 helus on the haptomerum and only subtriangular teges of long hamate setae with curved tips. The dense setiferous vestiture on the dorsum of abdominal segments I–VI and the sides of the last segment, aid in distinguishing H. mexicana from the known larvae of other Mexican melolonthine scarabs. Projections on the humeral area of the elytra and the middle of the abdominal segments of the pupae of both sexes may act as auxiliary supports during pupal development. These projections possibly aid in regulating humidity near the body wall. This type of structure is also observed in pupae of some Scarabaeinae (Edmonds and Halffter, 1978), and Ceratocanthinae (Morón and Arce, 2003).

Biology. Hoplia mexicana is known only from the northern mountains of the Mexican state of Oaxaca, at localities with elevations between 2300 and 2500m. Larvae have been collected in rich organic soil of oak and coniferous forest. Adults are rarely attracted to electric lights during summer, and their host plants are unknown.

Hoplia squamiferaBurmeister, 1844Third instar (Figs. 17–24)

Description. Head (Fig. 18). Maximum width of head capsule: 2.1–2.4mm. Surface of cranium smooth, with scattered minute punctures, pale yellowish. Frontal and epicranial sutures hidden. Frons with 2 exterior frontal setae, 1 posterior frontal seta, anterior frontal seta absent, 1 anterior angle seta, 3–4 dorsoepicranial, 14–18 epicranial setae on each side and 6–8 setae behind each antennal support. Clypeus with 2 central setae and 4 lateral setae on each side. Labrum nearly symmetrical with rounded lateral margins irregularly bordered, anterior margin projected forward with irregular border, basal transverse keel with 5 posterior setae, 1 anterior-central seta and 5–6 lateral setae on each side. Stemmata absent. Epipharynx (Fig. 21) 1.6–1.9mm wide, 1.0–0.2mm long. Epizygum absent and zygum as irregular, ovate, reddish yellow plate. Haptomerum weakly raised, with 4 heli; each plegmatia formed by 9–10 short plegmata; proplegmatia absent. Each acanthoparia with 9–10 spinose setae. Dexiophoba and laeophoba absent. Dexiotorma narrowed, slightly curved. Laeotorma elongate and narrowed with sclerotized plate unciform, raised. Sense cone short. Crepis weakly defined. Left chaetoparia with 30–34 spinose and medium size setae mixed. Right chaetoparia with 28–30 short, stout setae. Mandibles (Figs. 19 and 20) without ventral stridulatory area; scissorial area in both mandibles without distal blade. Right mandible with 1 small rounded, preapical tooth, inner margin simply curved, and distal lobe (M1) of molar area scarcely developed. Calx short. Left mandible without preapical tooth, inner margin simply curved, brustia moderately setose. Acia absent. Maxillae (Fig. 23). Mala with 1 uncus on apex of galea, 3 unci surrounded by 3–4 stout heli on apex of lacinia; stridulatory area with 10–11 small teeth with anteriorly directed points. Hypopharyngeal sclerome (Fig. 22) asymmetrical with curved projection on the right side. Fourth antennomere elongate, with distal half narrowed, with 1 ovate, dorsal sensory spot on basal half and 2 ventral sensory spots.

Figs. 17–21.

Third instar larva of Hoplia squamifera. 17, lateral habitus. 18, head, frontal view. Mandibles, ventral view: 19, left. 20, right. 21, epipharynx.

(0.43MB).
Figs. 22–25.

Hoplia squamifera. 22–24: third instar. 22, labium and hypopharynx. 23, left maxilla, dorsal view. 24, raster. 25, female pupa, lateral view.

(0.29MB).

Thorax. Respiratory plates light yellow, scarcely curved, “C” shaped (Fig. 17) 0.11–0.15mm long, 0.10–0.12mm wide, bulla small, slightly raised, rounded, distance between lobes longer than dorso-ventral diameter of bulla; microscopic holes of respiratory plate elongate-oval in outline and arranged in irregular transverse rows. Lateral sclerome of pronotum not defined. Proprescutum with 14–18 long setae regularly distributed; proscutum with 5–6 slender long setae; mesoprescutum with transverse row of 8–9 long setae; mesoscutum with 10–12 setae; mesoscutellum with row of 6–8 slender long setae; metaprescutum with transverse row of 9–12 mixed short and long setae; metascutum with 8–10 setae; metascutellum with transverse row of 6–8 mixed short and long setae (Fig. 17). Protarsal claws long, narrowed, sharply pointed, with 1 pre-basal inner seta and 1 lateral external seta, moderately longer than mesotarsal claws; mesotarsal claws long, sharply pointed, with pre-basal and lateral setae; metatarsal claw short, rounded, with 2 setae, much shorter than mesotarsal claws. Apical metatarsomere shortened, rounded, with ventral side clearly convex.

Abdomen. Respiratory plates light yellow, slightly curved, “C” shaped; on segments I–IV with similar diameter, 0.13–0.14mm long, 0.11mm wide; plates on segments V–VII slightly smaller, 0.11–0.12mm long, 0.10mm wide. Respiratory plate on segment VIII smaller than plate on preceding segment. Dorsa of abdominal segments I–VI each with dense vestiture of yellow stout, short setae; dorsa of segments VII–IX with transverse rows of 18–20 slender, short setae. Venter of abdominal segments I–VIII with transverse rows of 10–12 slender, long setae; venter of segment IX with transverse rows of 10–12 mixed short and long setae. Raster (Fig. 24) with each palidia formed by 10–12 pali, convergent toward basal and distal extremes, septula wide, oval; tegilla with 24–30 short, stout setae; campus with 14–16 slender, short setae; barbula densely setose. Dorsal anal lip with dense vestiture of short setae.

Ventral anal lip with scattered short setae at sides and distal border narrowly notched at middle. Anal slit “Y” shaped. Approximate dorsal body length 16–18mm.

Pupa (Figs. 25 and 26)

Description: body length 7.8–8.1mm. Widest width 4.5–4.6mm. Head. Surface glabrous, strongly deflexed; frons convex with two rounded prominences; clypeus widely concave on the center; labrum, mandibles, maxillae and palps discernible; antennal theca briefly expanded, stout with apex rounded; eyes small (Fig. 25). Thorax: surface glabrous. Pronotum convex, surface slightly irregular, anterior angles briefly prominent, posterior angles widely rounded. Meso- and metanotum well-differentiated. Elytral and posterior wing thecae closely appressed, curved ventrally around body; elytral thecae with irregular longitudinal striae and large, prominent, semiconical humeral tubercles (Figs. 25 and 26); thecae of the wings slightly longer than elytral thecae. Protibiae with apical and preapical tubercles clearly developed. Meso and metatibiae with apical tubercle. Abdomen: segments I–VI clearly wider and shorter than the distal segments VII–VIII, without dioneiform organs, but segments II–IV with pairs of prominent tubercles and segment V with thicker posterior border (Figs. 25 and 26). Pleural lobes rounded. Spiracle I elongate, with fine peritreme partially covered by wing thecae; spiracles II–IV ovate and high, prominent, with narrow sclerotized peritreme; spiracles V–VIII closed. Sutures between segments VIII and X incomplete, partially fused. Last segment slightly pruinose, without urogomphi. Last abdominal segment of male with dorsal and lateral rounded prominences, and genital ampulla ventrally exposed (Fig. 26); female without such abdominal prominences (Fig. 25).

Fig. 26.

Male pupa of Hoplia squamifera, with third instar exuviae on its abdominal apex.

(0.2MB).

Specimens examined. Seven third instar larvae, collected at Mexico: Chiapas, Amatenango del Valle municipality, El Madronal, 2400m, 20-I-1997, corn field, C. Ramírez col. (IEXA). Two pupae and 2 exuviae of third instar larvae reared to adults, with same data, except 24-X-2005 (IEXA). Two third instar larvae: Chiapas, Amatenango del Valle, 28-X-1996, corn field, C. Ramírez; 4 third instar larvae with same data, except 4-XI-1996; 5 third instar larvae 24-IX-2001 (IEXA, ECOSUR).

Comments. The larvae of Hoplia squamifera resemble those of H. mexicana, but lack antero-frontal setae, have 9–10 wide plegmata on each plegmatia, and have wide septula, whereas Hoplia mexicana have 2 antero-frontal setae, 7–8 narrow plegmata on each plegmatia and narrower septula.

Pupae of both Hoplia mexicana, and H. squamifera lack urogomphi and dioneiform organs. This fact is very interesting because, as far as we know, no other scarab group possesses such a combination of characters. For instance, with the exception of Manopus biguttatus Laporte, 1840 (Neita-Moreno et al., 2012), known pupae of Melolonthinae have urogomphi and lack dioneiform organs (Table 1). On the other hand, known pupae of Rutelinae and Dynastinae, lack urogomphi but, in contrast, have dioneiform organs (Morón, 1993). With the combination of these characters, and the unique paired supporting organs in the humeri and tergites, the pupae of Hopliinae are very distinct from other subfamilies of Melolonthidae. Description of the pupae of many other genera of Melolonthidae surely will offer more characters to be tested in future phylogenetic analyses of Scarabaeoidea.

Table 1.

Main characters in the pupae of some American genera of Scarabaeoidea. D.O., dioneiform organs; H.P., humeral projections; P.P., pronotal projections; T.P., tergal projections; U, urogomphi.

Subfamilies  Genera  P.P.  H.P.  D.O.  T.P.  References 
MelolonthinaePhyllophaga Harris, 1827  No  No  No  No  Yes  Morón (1986) 
Polyphylla Harris, 1841  No  No  No  No  Yes  Morón and Nogueira (2015) 
Clavipalpus Laporte,Castelnau, 1832  No  No  No  No  Yes  Neita-Moreno et al. (2012) 
Demodema Blanchard, 1850  No  No  No  No  Yes  Morón and Salvadori (2006) 
Isonychus Mannerheim, 1829  No  No  No  No  Yes  Neita-Moreno et al. (2012) 
Macrodactylus Dejean, 1821  No  No  No  No  Yes  Ramírez-Salinas et al. (2011) 
Manopus Laporte Castelnau, 1840  No  No  No  No  No  Neita-Moreno et al. (2012) 
HopliinaeRutelinaeHoplia Illiger, 1803  No  Yes  No  Yes  No  Present publication 
Anomala Samouelle, 1819  No  No  Yes  No  No  Micó et al. (2003) 
Macraspis MacLeay, 1819  No  No  Yes  No  No  Morón and Paucar-Cabrera (2003) 
ChlorotaBurmeister, 1844  No  No  Yes  No  No  Jameson and Morón (2001) 
Chrysophora LeP. & Serville, 1828  No  No  Yes  No  No  Pardo-Locarno and Morón (2007) 
Leucothyreus MacLeay, 1819  No  No  Yes  No  No  Pardo-Locarno et al. (2006b) 
DynastinaeLigyrus Burmeister, 1847  No  No  Yes  No  No  Ramírez-Salinas et al. (2004) 
Ancognatha Erichson, 1847  No  No  Yes  No  No  Ramírez-Salinas et al. (2004) 
Coelosis Hope, 1837  No  No  Yes  No  No  Pardo-Locarno et al. (2006a) 
Golofa Hope, 1837  No  No  Yes  No  No  Morón (1995b) 
Dynastes Kirby, 1825  No  No  Yes  No  No  Morón (1987) 
Aegopsis Burmeister, 1847  No  No  Yes  No  No  Neita-Moreno et al. (2014) 
CetoniinaeEuphoria Burmeister, 1842  No  No  No  No  No  Ramírez-Salinas et al. (2001) 
Hoplopyga J. Thomson, 1880  No  No  No  No  No  Morón and Arce (2002) 
Gymnetis MacLeay, 1819  No  No  No  No  No  Morón and Arce (2002) 
Neoscelis Schoch, 1897  No  No  No  No  No  Nogueira et al. (2004) 
TrichiinaeInca LePeletier & Serville, 1828  No  No  Yes  No  No  Morón (1983) 
Archedinus Morón & Krikken, 1990 Krikken, 1990  No  No  Yes  No  No  Morón (1995a) 
Orphninae  Aegidium Westwood, 1846  No  No  No  Yes  Yes  Morón (1991) 
Ceratocanthinae  Ceratocanthus White, 1842  Yes  Yes  No  No  Yes  Morón and Arce (2003) 
Aphodiinae  Termitodius Wasmann, 1894  No  Yes  No  Yes  Yes  Reyes-Castillo and Martínez (1979) 
ScarabaeinaePhanaeus MacLeay, 1819  Yes  No  No  Yes  No  Edmonds (1967) 
Eurysternus Dalman, 1824  No  No  No  Yes  No  López-Guerrero and Morón (1994) 

Biology. Hoplia squamifera has been recorded from the Mexican mountains of northern Chiapas and Oaxaca states, southern Veracruz state, and in western Guatemala at localities with elevations between 1200 and 2100m. Larvae have been collected in soil of corn fields in northern Chiapas, Mexico. Adults are frequently attracted to electric lights during spring and summer, but their host plants are unknown.

Conflicts of interest

The authors declare no conflicts of interest.

Acknowledgements

The aid of Adriana E. Castro (ECOSUR) and Susana Guzmán (Instituto de Biología, UNAM, Mexico, D.F.) during field and technical work is here recognized. The suggestions of two anonymous reviewers were very useful in improving the final version of the manuscript. We thank Bruce Gill (Ottawa, Canada) for his friendly assistance with the English language. This paper was supported in part by the research line “Coleópteros Lamelicornios de América Latina” Instituto de Ecología, A.C. Xalapa, Mexico (# 2004010011).

References
[Ansari et al., 2006]
M.A. Ansari, H. Casteels, L. Tirry, M. Moens.
Biology of Hoplia philanthus (Coleoptera: Scarabaeidae: Melolonthinae). A new and severe pest in Belgian turf.
Environ. Entomol., 35 (2006), pp. 1500-1507
[Ansari et al., 2008]
M.A. Ansari, B.N. Adhikari, F. Ali, M. Moens.
Susceptibility of Hoplia philanthus (Coleoptera: Scarabaeidae) larvae and pupae to entomopathogenic nematodes (Rhabditida: Steirnernematidae, Heterorhabditidae).
Biol. Control, 47 (2008), pp. 315-321
[Burmeister, 1844]
H.C.C. Burmeister.
Handbuch der Entomologie (Coleoptera Lamellicornia Anthobia et Phyllophaga Systellochela) vol. 4, part 1.
T. C. F. Enslin, (1844),
[Harold, 1869]
E. Harold.
Scarabaeidae.
pp. 979-1346
[Edmonds, 1967]
W.D. Edmonds.
The immature stages of Phanaeus (Coprophanaeus) jasius Olivier and Phanaeus (Metallophanaeus) saphirinus Sturm (Coleoptera: Scarabaeidae).
Coleopterist Bull., 21 (1967), pp. 97-105
[Edmonds and Halffter, 1978]
W.D. Edmonds, G. Halffter.
Taxonomic review of immature dung beetles of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae).
Syst. Entomol., 3 (1978), pp. 307-331
[Hůrka and Bílý, 1966]
K. Hůrka, S. Bílý.
Die larve von Hoplia philanthus Fuessly (Coleoptera, Scarabaeidae) nebst Bemerkungen zur Bionomie der Art.
Acta Entomológica Bohemoslovaca, 63 (1966), pp. 459-463
[Jameson and Morón, 2001]
M.L. Jameson, M.A. Morón.
Descriptions of the larvae of Chlorota cincticollis Blanchard and Chasmodia collaris (Blanchard) (Scarabaeidae: Rutelinae: Rutelini) with a key to the larvae of the American genera of Rutelini.
Coleopterists Bull., 55 (2001), pp. 385-396
[López-Guerrero and Morón, 1994]
Y. López-Guerrero, M.A. Morón.
Description of immature stages of Eurysternus mexicanus Burmeister and Sisyphus submonticolus Howden (Coleoptera: Scarabaeidae: Scarabaeinae).
J. Kansas Entomol. Soc., 67 (1994), pp. 347-353
[Medvedev, 1952]
S.I. Medvedev.
Larvae of Scarabaeid beetles of the fauna of the USSR.
Opredeliteri po faune SSSR, Moskva, (1952), pp. 47
[Micó, 2001]
E. Micó.
Los escarabeidos antófilos de la península Ibérica (Col. Scarabaeoidea: Hopliinae, Rutelidae, Cetoniidae): taxonomía, filogenia y biología.
Universidad de Alicante, (2001),
[Micó et al., 2003]
E. Micó, M.A. Morón, E. Galante.
New larval descriptions and biology of some New World Anomalini beetles (Coleoptera: Scarabaeoidea: Rutelinae).
Ann. Entomol. Soc. Am., 96 (2003), pp. 597-614
[Morón, 1983]
M.A. Morón.
Los estados inmaduros de Inca clathrata sommeri Westw (Col. Melolonthidae, Trichiinae).
Folia Entomológica Mexicana, 56 (1983), pp. 31-51
[Morón, 1986]
M.A. Morón.
El género Phyllophaga en México. Morfología, Distribución y Sistemática Supraespecífica (Insecta: Coleoptera)..
Instituto de Ecología, (1986),
[Morón, 1987]
M.A. Morón.
Los estados inmaduros de Dynastes hyllus Chevrolat (Col. Melolonthidae, Dynastinae) con observaciones sobre su biología y el crecimiento alométrico del imago.
Folia Entomológica Mexicana, 72 (1987), pp. 33-74
[Morón, 1991]
M.A. Morón.
The immature stages of Aegidium cribratum Bates (Coleoptera, Scarabaeidae, Orphninae).
Coleopterists Bull., 45 (1991), pp. 360-367
[Morón, 1993]
M.A. Morón.
Observaciones comparativas sobre la morfología pupal de los Coleoptera Melolonthidae neotropicales.
Giornale Italiano di Entomologia, 6 (1993), pp. 249-255
[Morón, 1995a]
M.A. Morón.
Larva and pupa of Archedinus relictus Morón & Krikken (Coleoptera: Melolonthidae, Trichiinae, Incaini).
Pan-Pacific Entomol., 71 (1995), pp. 237-244
[Morón, 1995b]
M.A. Morón.
Review of the Mexican species of Golofa Hope (Coleoptera: Melolonthidae, Dynastinae).
Coleopterist's Bull., 49 (1995), pp. 343-386
[Morón and Arce, 2002]
M.A. Morón, R. Arce.
Descriptions of the immature stages of five Mexican species of Gymnetini (Coleoptera: Scarabaeidae: Cetoniinae).
Proc. Entomol. Soc. Washington, 104 (2002), pp. 1036-1054
[Morón and Arce, 2003]
M.A. Morón, R. Arce.
Description of third instar larva and pupa of Ceratocanthus relucens (Bates) (Coleoptera: Scarabaeidae; Ceratocanthinae).
Coleopterists Bull., 57 (2003), pp. 245-253
[Morón and Nogueira, 2015]
M.A. Morón, G. Nogueira.
Descripción de los estados inmaduros de Polyphylla conspersa Burmeister (Coleoptera: Melolonthidae: Melolonthinae) con observaciones sobre su biología.
Boletín Sociedad Entomológica Aragonesa, 57 (2015),
[Morón and Paucar-Cabrera, 2003]
M.A. Morón, A. Paucar-Cabrera.
Larvae and pupae of species of Macraspis Macleay (Coleoptera: Scarabaeidae: Rutelinae: Rutelini).
Can. Entomol., 135 (2003), pp. 467-491
[Morón and Salvadori, 2006]
M.A. Morón, J.R. Salvadori.
Third stage larva and pupa of Demodema brevitarsis Blanchard (Coleoptera: Scarabaeidae: Melolonthinae).
Proc. Entomol. Soc. Washington, 108 (2006), pp. 511-518
[Neita-Moreno et al., 2012]
J.C. Neita-Moreno, M.A. Morón, C.A. Zuluaga-Correa.
Description of the immature stages of four species of Macrodactylini (Coleoptera: Melolonthidae: Melolonthinae).
Neotrop. Entomol., 41 (2012), pp. 150-162
[Neita-Moreno et al., 2014]
J.C. Neita-Moreno, M.C. Oliveira, M.A. Morón.
Larval and pupal descriptions of two Aegopsis species (Coleoptera: Melolonthidae: Dynastinae).
Florida Entomol., 97 (2014), pp. 114-125
[Nogueira et al., 2004]
G. Nogueira, M.A. Morón, H.E. Fierros-López, J.L. Navarrete-Heredia.
The immature stages of Neoscelis dohrni (Westwood) (Coleoptera: Scarabaeidae: Cetoniinae: Goliathini) with notes on the adult behavior.
Coleopterists Bull., 58 (2004), pp. 171-183
[Pardo-Locarno and Morón, 2007]
L.C. Pardo-Locarno, M.A. Morón.
Larva and pupa of Chrysophora chrysochlora (Coleoptera: Scarabaeidae: Rutelinae: Rutelini).
Can. Entomol., 139 (2007), pp. 80-86
[Pardo-Locarno et al., 2006a]
L.C. Pardo-Locarno, M.A. Morón, A. Gaigl.
Los estados inmaduros de Coelosis biloba (Linné, 1767) (Coleoptera: Melolonthidae: Dynastinae) y notas sobre su biología.
Revista Mexicana de Biodiversidad, 77 (2006), pp. 215-224
[Pardo-Locarno et al., 2006b]
L.C. Pardo-Locarno, M.A. Morón, J. Montoya-Lerma.
Descripción de los estados inmaduros de Leucothyreus femoratus (Coleoptera: Melolonthidae, Rutelinae: Geniatini) con notas sobre su biología e importancia agrícola en Colombia.
Folia Entomológica Mexicana, 45 (2006), pp. 179-193
[Ramírez-Salinas et al., 2001]
C. Ramírez-Salinas, A. Castro-Ramírez, M.A. Morón.
Descripción de la larva y la pupa de Euphoria basalis (G & P.)(Coleoptera: Melolonthidae; Cetoniinae).
Acta Zoológica Mexicana (n.s.), 83 (2001), pp. 73-82
[Ramírez-Salinas et al., 2004]
C. Ramírez-Salinas, M.A. Morón, A. Castro-Ramírez.
Descripción de los estados inmaduros de tres especies de Anomala, Ancognatha y Ligyrus (Coleoptera: Melolonthidae: Rutelinae y Dynastinae) con observaciones de su biología.
Acta Zoológica Mexicana (n.s.), 20 (2004), pp. 67-82
[Ramírez-Salinas et al., 2011]
C. Ramírez-Salinas, M.A. Morón, A. Castro-Ramírez.
Descripción de los inmaduros de cuatro especies de Phyllophaga, Paranomala y Macrodactylus (Coleoptera: Melolonthidae) de los Altos de Chiapas, México.
Acta Zoológica Mexicana (n.s.), 27 (2011), pp. 527-545
[Reyes-Castillo and Martínez, 1979]
P. Reyes-Castillo, A. Martínez.
Nuevos Rhyparini neotropicales, con notas sobre su biología (Coleoptera, Scarabaeidae, Aphodiinae).
Folia Entomológica Mexicana, 41 (1979), pp. 115-133
[Ritcher, 1949]
P.O. Ritcher.
Larvae of Melolonthinae with keys to tribes, genera and species (Coleoptera: Scarabaeidae).
Kentucky Agric. Exp. Station Bull., 537 (1949), pp. 1-36
[Ritcher, 1966]
P.O. Ritcher.
White Grubs and Their Allies.
Oregon State University Press, (1966),
Copyright © 2015. Sociedade Brasileira de Entomologia
Idiomas
Revista Brasileira de Entomologia

Subscribe to our newsletter

Article options
Tools
en pt
Cookies policy Política de cookies
To improve our services and products, we use "cookies" (own or third parties authorized) to show advertising related to client preferences through the analyses of navigation customer behavior. Continuing navigation will be considered as acceptance of this use. You can change the settings or obtain more information by clicking here. Utilizamos cookies próprios e de terceiros para melhorar nossos serviços e mostrar publicidade relacionada às suas preferências, analisando seus hábitos de navegação. Se continuar a navegar, consideramos que aceita o seu uso. Você pode alterar a configuração ou obter mais informações aqui.