SYSTEMATICS Biology and New Larval Descriptions for Three Cetoniine (Coleoptera: : Cetoniinae: : Cetoniina, Leucocelina)

E. MICO´ AND E. GALANTE

Centro Iberoamericano de la Biodiversidad (CIBIO), Universidad de Alicante, 03080-Alicante, Downloaded from https://academic.oup.com/aesa/article/96/2/95/27977 by guest on 24 September 2021

Ann. Entomol. Soc. Am. 96(2): 95Ð106 (2003) ABSTRACT The larval morphology of squalida (Scopoli) and Aethiessa floralis (Fabricius) is described. The latter is the Þrst description of a in this . The third-instar larva of funesta (Poda) is redescribed. These species are included in a revised key to the larvae of Palaearctic Cetoniinae. The life cycle and larval biology of these ßower chafers are described.

RESUMEN Se describe la morfologõ´a larvaria de (Scopoli) y Aethiessa floralis (Fabricius). Esta u´ ltima constituye la primera descripcio´n larvaria de este ge´nero. Se redescribe la larva de O. funesta (Poda). Las mencionadas especies Se han incluido en una clave de identiÞcacio´n para las de las especies Palea´rticas de Cetoniinae. se describe el ciclo biolo´gico y la biologõ´a larvaria de dichas especies.

KEY WORDS Scarabaeidae, Cetoniinae, Tropinota squalida, , Aethiessa floralis, larval biology

Oxythyrea MULSANT (Cetoniini: Leucocelina), Tropi- tae) species are abundant (Mico´ and Galante 1998). nota Mulsant, and Aethiessa (Cetoniini: Cetoniina) The morphology and biology of the immature stages of Burmeister are small genera (10, 10, and 7 species, the genus Aethiessa were unknown until now. respectively) that occur in the Palaearctic region. The aims of our study were: 1) to describe the Tropinota squalida (Scopoli, 1763) is distributed in the third-instar larva of T. squalida and A. floralis and to occidental Mediterranean basin, and Oxythyrea fun- redescribe the larva of O. funesta; 2) to provide a esta (Poda, 1761) occurs in the occidental Palaearctic revised key with the speciÞc diagnostic characters of region. Adults of both species have been considered the larvae of Palaearctic Cetoniinae; and (3) to con- occasional pests of crops and ornamental plants (Jans- tribute to the knowledge of the larval biology and life sens 1960, Balachowsky 1962, Molina 2001). Although cycles of these species. several experiments have tested the larval survival rates at different temperatures (Hurpin 1958, Bala- chowsky 1962), or by altering the substrate (Abdalla Materials and Methods 1991), larval morphology is little known. Golovjanko A total of 91 larvae of T. squalida, 36 larvae of O. (1936) and Korschefsky (1940) illustrated the venter funesta, and 7 larvae of A. floralis were reared from egg of the last abdominal segment of O. funesta. Medvedev to adults to study the life cycle of the species under (1952) illustrated the frontal view of the head and the laboratory conditions. These larvae were fed milled venter of the last abdominal segment with details of rabbit dung heaps, decaying vegetative matter, and the raster, including a short description of these struc- manure. The breeding cages were maintained in an tures. Janssens (1960) described some morphological environmental chamber at 25ЊC:20ЊC (L : D), 80 Ϯ characters of the larvae of T. squalida, but no illustra- 5% RH, with a photoperiod of 15:9(L:D).The tion was provided. None of these contributions rep- breeding cages were examined weekly, and the results resented a complete description, and the character- were recorded. Larvae used for morphological de- istics provided are insufÞcient to distinguish them scriptions were reared in the above laboratory con- from their congeners. Aethiessa floralis (Fabricius, ditions. The different larval instars of each species 1787) occurs in the occidental Mediterranean basin were Þxed in KAAD solution (Carne 1951) for 24 h mainly in formerly cultivated areas characterized by and preserved in 70% ethanol. Specimens are depos- nitrophilous vegetation where Carduinae (Composi- ited in the Entomological Collection of the University

0013-8746/03/0095Ð0106$04.00/0 ᭧ 2003 Entomological Society of America 96 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 2 of Alicante, Spain (CEUA). In the description of lar- six sensilla. Lateral lobe with seven to eight setae on vae, the terminology of Ritcher (1966) and Mico´ et al. each side. Antenna (Fig. 5). Four segmented. Apical (2001) was used. For the suprageneric nomenclature, segment with two dorsal and three ventral sensory we have followed the scarab classiÞcation of Krikken spots. (1984). Thorax. Spiracles (Fig. 4). Respiratory plate of tho- Phenology of adults was based on the labeled spec- racic spiracles with 10Ð16 holes across diameter; lobes imens provided by CEUA, the National Natural Sci- unequal. Legs (Fig. 3). Tarsunguli short (0.5 times the ence Museum (MNCN, Madrid, Spain), the Zoology preceding segment), cylindrical, bearing 12Ð13 setae. Museum (Barcelona, Spain), and the National Natural Abdomen. Spiracles of abdominal segments II-VII History Museum (MNHN, Paris, France), as well as similar in size; those of I and VIII conspicuously our direct and indirect (baited trap) captures. smaller. Dorsa of abdominal segments I-VIII with three to four rows of short setae, each posterior row

with long to short setae. Abdominal segments IX-X Downloaded from https://academic.oup.com/aesa/article/96/2/95/27977 by guest on 24 September 2021 Results and Discussion fused, densely setose with short setae and single row of long to short setae at the middle and apex (Fig. 1). T. squalida (Scopoli, 1763) Third-Instar Larva Tegilla and lower anal lip composed of short and long (Figs. 1Ð13) setae (Fig. 6). Raster with pair of palidia diverging The larval description is based on two third-instar posteriorly, each palidium consisting of 17Ð21 short, larvae reared from eggs laid by adults collected at “La acute pali (Fig. 7). Granadella,” Alicante (Spain), IV-1997, Mico´ and Diagnosis. One of the most remarkable diagnostic Verdu´ leg.; one third-instar larva reared from eggs laid characters of T. squalida is the wide stridulatory area by adults collected at “Sierra de Aixorta´,” Alicante of the mandibles (Fig. 11). This area is conspicuously (Spain), V-1996, Mico´ and Verdu´ leg.; one third-instar wider than in other known larvae of Palaearctic Ce- larva collected at “La Mata,” Torrevieja, Alicante toniini (Mico´ and Galante 2003). The short tarsungu- (Spain), IV-1995, Mico´ and Verdu´ leg.; one third-instar lus of this species is also noticeable, being 0.5 times the larva collected at “Palacio de Don˜ ana,” Huelva length of the previous segment (Fig. 3). (Spain), VIII-1999, Mico´ and Verdu´ leg. Head. Maximum width of head capsule, 3.2 mm. O. funesta (Poda, 1761) Third-Instar Larva Cranium (Fig. 2). Color, light yellow. Frons sparsely (Figs. 14Ð26) punctate, with one anterior seta, one posterior seta, one external seta, and one anterior angle seta on each The larval description is based on six third-instar side. Frontal suture widely sinuated. Labrum (Fig. 2). larvae reared from eggs laid by adults collected at Trilobed, narrower than clypeus; clithra present. “Arenales del Sol,” Alicante (Spain), Mico´ leg.; three Epipharynx (Fig. 8). Plegmatium absent. Corypha third-instar larvae collected at Boren, Le´rida, (Spain), with four long setae ßanked by one to two sensilla on 5-VIII-1998, Mico´ and Verdu´ leg. each side. Acanthoparia with 10Ð12 setae decreasing Head. Maximum width of head capsule 2.7 mm. in size posteriorly. Right chaetoparia more developed Cranium (Fig. 15). Color, light yellow. Frons with one than left chaetoparia, covered with longitudinal rows posterior seta and one anterior angle seta on each side; of setae. Pedium reduced. Laeotorma short with pter- anterior and external setae reduced to single micro- notorma present. Dexiotorma long, one-thirds length seta on each side. Frontal suture slightly sinuated. of the base of epipharynx, pternotorma present. Hap- Labrum (Fig. 15). Trilobed, narrower and shorter than tomeral region with slightly curved, transverse row of clypeus; clithra present. Epipharynx (Fig. 21). Pleg- 12Ð13 heli and three to four sensilla over the row. matium absent. Corypha with four long setae ßanked Haptolachus with four sensilla (two at base and two at by one to two sensilla on each side. Acanthoparia with left margin). Sensorial cone acute at apex. Sclerotized Þve to six short setae. Chaetoparia covered with four plate and crepis absent. Mandibles (Figs. 10Ð12). to Þve longitudinal rows of setae. Laeotorma short Asymmetrical, with two scissorial teeth anterior to with pternotorma present. Dexiotorma long, one- notch; two posterior to notch at left mandible; and one thirds length of the base of epipharynx, pternotorma posterior to notch at right mandible. Scissorial teeth of present. Haptomeral region with slightly curved, left mandible S3 and S4 well developed, parallel to S1 transverse row of 12Ð13 heli and three to four sensilla and S2. Stridulatory area oval, well developed, with over the row. Haptolachus with four sensilla (two at Ϸ20 stridulatory ridges (Fig. 12). Dorsal surface with base and two at left margin). Middle of haptolachus two setae near proximal end of the scissorial area. with slightly sclerotized longitudinal area. Sensorial Basomedial angle with brustia of short setae. Maxilla cone present. Sclerotized plate and crepis absent. (Fig. 13). Galea and lacinia fused forming mala. Mala Mandibles (Figs. 23, 24, 25). Asymmetrical, with two with large uncus at apex and two subterminal unci scissorial teeth anterior to notch; two posterior to fused at base. Stridulatory area consisting of a row of notch at left mandible, and one posterior to notch at four to Þve acute teeth and one small anterior conical right mandible. Scissorial teeth of left mandible S3 and process. Labium (Fig. 9). Hypopharyngeal sclerome S4 well developed, parallel to S1 and S2. Stridulatory with well-developed truncate process on right side. area elongated-oval, with 9Ð11 stridulatory ridges Glossa with three to Þve setae set in two rows on each (Fig. 25). Dorsal surface with two setae near the prox- side; base with transverse row of short setae; apex with imal end of the scissorial area. Basomedial angle with March 2003 MICO´ AND GALANTE:CETONIINAE LARVAL DESCRIPTIONS 97 Downloaded from https://academic.oup.com/aesa/article/96/2/95/27977 by guest on 24 September 2021

Figs. 1–7. T. squalida third-instar larva. 1. Habitus. 2. Frontal view of head. 3. Tarsungulus of anterior leg. 4. Thoracic spiracle. 5. Ventral view of antenna (a) and dorsal view of apical antennal segment showing sensory spots (b). 6. Venter of last abdominal segment. 7. Detail of palidium. 98 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 2 Downloaded from https://academic.oup.com/aesa/article/96/2/95/27977 by guest on 24 September 2021

Figs. 8–13. T. squalida third-instar larva. 8. Epipharynx. 9. Dorsal view of hypopharynx. 10. Dorsal view (a), lateral view (b), and ventral view (c) of left mandible. 11. Ventral view (a), lateral view (b), and dorsal view (c) of right mandible. 12. Stridulatory area of mandible. 13. Ventral (a) and dorsal (b) view of left maxilla. Apex of mala showing unci (c). March 2003 MICO´ AND GALANTE:CETONIINAE LARVAL DESCRIPTIONS 99 Downloaded from https://academic.oup.com/aesa/article/96/2/95/27977 by guest on 24 September 2021

Figs. 14–20. O. funesta third-instar lava. 14. Habitus. 15. Frontal view of head. 16. Tarsungulus of anterior leg. 17. Thoracic spiracle. 18. Ventral view of antenna (a), dorsal view (b), and lateral view (c) of apical antennal segment showing sensory spots. 19. Venter of last abdominal segment. 20. Detail of palidium. 100 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 2 Downloaded from https://academic.oup.com/aesa/article/96/2/95/27977 by guest on 24 September 2021

Figs. 21–26. O. funesta third-instar larva. 21. Epipharynx. 22. Dorsal view of hypopharynx. 23. Dorsal view (a), lateral view (b), and ventral view (c) of left mandible. 24. Ventral view (a), lateral view (b), and dorsal view (c) of right mandible. 25. Stridulatory area of mandible. 26. Ventral (a) and dorsal (b) view of left maxilla. Apex of mala showing unci (c). March 2003 MICO´ AND GALANTE:CETONIINAE LARVAL DESCRIPTIONS 101 brustia of short setae. Maxilla (Fig. 26). Galea and with 9Ð12 setae decreasing in size posteriorly. lacinia fused forming mala. Mala with large uncus at Chaetoparia well developed, with longitudinal rows of apex and two subterminal unci fused at base. Stridu- setae. Laeotorma short with pternotorma present. latory area consisting of a row of Þve to seven acute Dexiotorma short, one-fourths length of the base of teeth and one small anterior conical process. Labium epipharynx, pternotorma present. Haptomeral region (Fig. 22). Hypopharyngeal sclerome with well-devel- with slightly curved, transverse row of 12Ð13 heli and oped truncate process on right side. Glossa with three three to four sensilla over the row. Haptolachus with to Þve setae set in two rows on each side; base with four sensilla (two at base and two at left margin). transverse row of short setae; apex with four to six Sensorial cone present. Sclerotized plate and crepis sensilla. Lateral lobe with 6Ð10 setae on each side. absent. Mandibles (Figs. 36, 37, 38). Asymmetrical, Antenna (Fig. 18). Four segmented. Apical segment with two scissorial teeth anterior to notch; two pos- with two dorsal and three ventral sensory spots; apical terior to notch at left mandible, and one posterior to

segment two times the length of third segment. notch at right mandible. Scissorial teeth of left man- Downloaded from https://academic.oup.com/aesa/article/96/2/95/27977 by guest on 24 September 2021

Thorax. Spiracles (Fig. 17). Respiratory plate of dible S4 well developed. Stridulatory area elongate- thoracic spiracles with 10Ð14 holes across diameter; oval, with Ϸ14 stridulatory ridges (Fig. 38). Dorsal lobes unequal. Legs (Fig. 16). Tarsunguli cylindrical, surface with two setae near the proximal end of the bearing 9Ð10 setae. scissorial area. Basomedial angle with brustia of short Abdomen. Spiracles of abdominal segments II-VII setae. Maxilla (Fig. 39). Galea and lacinia fused form- similar in size; those of I and VIII slightly smaller. ing mala. Mala with large uncus at apex and two sub- Dorsa of abdominal segments I-VIII with three to four terminal unci fused at base. Stridulatory area consist- rows of short setae, each posterior row with long to ing of a row of seven to nine acute teeth and one small short setae. Abdominal segments IX-X fused. Dorsa of anterior conical process. Labium (Fig. 35). Hypopha- last abdominal segment characterized by the presence ryngeal sclerome with well-developed truncate pro- of conspicuously shorter setae in the apical half of the cess on right side. Glossa with three to Þve setae set in segment (Fig. 14). Tegilla and lower anal lip composed two rows on each side; base with transverse row of of short and long setae (Fig. 19). Raster with a single short setae; apex with four to Þve sensilla. Lateral lobe palidium horseshoe shaped consisting of Ϸ29 short with 10Ð18 setae on each side. Antenna (Fig. 31). Four and acute pali (Fig. 20). segmented. Apical segment with two dorsal and three Diagnosis. Known larvae of Leucocelina (Cetoni- ventral sensory spots. inae: Cetoniini) differ from those of Cetoniina by the Thorax. Spiracles (Fig. 30). Respiratory plate of horseshoe-shaped palidium (Donaldson 1987). Larvae thoracic spiracles with 25 holes across diameter. Legs of O. funesta share this character with other Leuco- (Fig. 29). Tarsunguli cylindrical, bearing 13 setae. celina such as Oxythyrea cintella (Schaum) (Med- Abdomen. Spiracles of abdominal segments similar vedev 1952), Oxythyrea hemorrhoidalis (Fabricius), in size. Dorsa of abdominal segments I-VIII with four Oxythyrea testaceoguttata Blanchard, Oxythyrea rubra to six rows of short setae, each posterior row with long (Gory and Percheron), Mausolopsis amabilis (Scham), to short setae. Abdominal segments IX-X fused, and Leptothyrea perroudi (Schaum) (Donaldson 1987). densely setose; setae of apical half longer. Tegilla com- Another diagnostic characteristic of O. funesta is the posed of short, acute setae. Lower anal lip with short presence of conspicuously shorter setae in the apical setae curved at apex and a posterior row of long setae half of the dorsum of the last abdominal segment (Fig. (Fig. 32). Raster with a pair of palidia convergent 14). The dorsum of the last abdominal segment has not posteriorly, each palidium consisting of 17Ð21 short been illustrated for the other known Leucocelina. and acute pali (Fig. 33). Diagnosis. The description of the larvae of A. floralis constitutes the Þrst description of a larva for this ge- A. floralis (Fabricius, 1787) Third-Instar Larva nus. One of the diagnostic characteristics used to sep- (Figs. 27Ð39) arate this genus from the other known Palaearctic The larval description is based on Þve third-instar genera is the presence of several posterior frontal larvae reared from eggs laid by adults collected at “El setae (Fig. 28) and the scissorial area of the left man- Garbinet,” Alicante (Spain), 15-VII-1997, Mico´ leg.; dible showing a prominent fourth tooth disposed per- two third-instar larvae collected at “La Mata,” Torre- pendicular to the mandibular axis (Fig. 37, a and c). vieja, Alicante (Spain), IV-1998, Mico´ and Verdu´ leg.; The epipharynx was similar to those of Tropinota, four third-instar larvae collected at “Arenales del Sol,” characterized by the Þner and decreasingly small setae Alicante (Spain), V-1996, Mico´ and Verdu´ leg. of the acanthoparia, while in the other Palaearctic Head. Maximum width of head capsule 4.6 mm. species these setae are stout and shorter (Mico´ and Cranium (Fig. 28). Color, yellowish-brown. Frons Galante 2003). sparsely punctate with one anterior seta, three to Þve posterior setae, one external seta, and one anterior angle seta on each side. Frontal suture widely sinu- Key to Species of the Known Third-Instar Larvae of ated. Labrum (Fig. 28). Trilobed, narrower than the Palaearctic Cetoniinae clypeus; clithra present. Epipharynx (Fig. 34). Pleg- matium absent. Corypha with four long setae ßanked The most important contribution to the knowledge by two to three sensilla on each side. Acanthoparia of Palaearctic cetoniine larvae has been provided 102 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 2 Downloaded from https://academic.oup.com/aesa/article/96/2/95/27977 by guest on 24 September 2021

Figs. 27–33. A. floralis third-instar larva. 27. Habitus. 28. Frontal view of head. 29. Tarsungulus of anterior leg. 30. Thoracic spiracle with details of respiratory holes. 31. Ventral view of antenna showing sensory spots. 32. Venter of last abdominal segment. 33. Detail of palidium. March 2003 MICO´ AND GALANTE:CETONIINAE LARVAL DESCRIPTIONS 103 Downloaded from https://academic.oup.com/aesa/article/96/2/95/27977 by guest on 24 September 2021

Figs. 34–39. A. floralis third-instar larva. 34. Epipharynx. 35. Dorsal view of hypopharynx. 36. Ventral view (a), lateral view (b), and dorsal view (c) of right mandible. 37. Dorsal view (a), lateral view (b), and ventral view (c) of left mandible. 38. Stridulatory area of mandible. 39. Dorsal view (a) and ventral view (b) of left maxilla. Apex of mala showing unci (c).

by Golovjanko (1936), Korschefsky (1940), Van Emden (1941). We supplement the knowledge of Emden (1941), and Medvedev (1952). Later, Palaearctic Cetoniinae larvae with the redescription Klausnitzer and Krell (1997) provided a key based of one species and the Þrst descriptions of two on the illustrations of Korschefsky (1940) and Van species and one genus. 104 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 2

Table 1. Described Larvae of Palaearctic Cetoniidae 5. Frons and clypeus coarsely pitted ...... Cetonischema aeruginosa (Drury) Species Publication 5Ј. Frons and clypeus smooth ...... 6 Aethiessa floralis (F.) This publication 6. Head with anterior frontal setae reduced to mi- aeratula Reitter Medvedev 1952 crosetae or absent (Fig. 15) ...... 7 Golovjanko 1936, Van Edmen 1941, Ј (Linnaeus) Medvedev 1952 6 . Head with anterior frontal setae and external Cetonischema aeruginosa Medvedev 1952 frontal setae medium to long (Figs. 2 and 28). (Drury) (Netocia karelini [Zoubkov]) ...... 9 Eupotosia affinis Ting 1934 (Andersch) 7. Raster with horseshoe-shaped palidium (Fig. Gnorimus Le Pelletier Van Edmen 1941, Medvedev 1952 20). [Oxythyrea cintella Schaum] ...... and Serville ...... Oxythyrea funesta (Poda) Liocola lugubris Medvedev 1952 7Ј. Palidium consisting of two rows of pali (Figs. 7 (Herbst) Netocia hungarica Medvedev 1952 and 33). (Cetonia aurata [Linnaeus], Cetonia Downloaded from https://academic.oup.com/aesa/article/96/2/95/27977 by guest on 24 September 2021 (Herbst) aeratula Reitter, Liocola lugubris [Herbst], Netocia karelini Medvedev 1952 Potosia fieberi [Medvedev], Eupotosia affinis (Zoubkov) [Andersch]) ...... 8 Netocia morio Mico´ and Galante 2002 (Gory and Percheron) 8. Stridulatory area of mandibles consisting of Netocia oblonga Mico´ and Galante 2002 24Ð25 transverse ridges; inter-ridge area 5Ð8 (Gory and Percheron) times ridge width ...... Netocia trojana Medvedev 1952 (Gory and Percheron) ...... Potosia cuprea brancoi Baraud Osmoderma eremita Medvedev 1952 8Ј. Stridulatory area of mandibles consisting of (Scopoli) 15Ð19 transverse ridges; inter-ridge area 1.7Ð Oxythyrea cintella Medvedev 1952 2.0 times ridge width . Potosia opaca Fabricius Schaum Oxythyrea funesta Golovjanko 1936, Medvedev 1952, 9. Palidia with 17Ð22 set of pali. Pali acute at apex. (Poda) This publication ...... 10 Potosia cuprea brancoi Mico´ and Galante 2002 9Ј. Palidia with 11Ð14 set of pali. Pali round at apex. Baraud Potosia cuprea metallica Korchefsky 1940, Van Edmen 1941, (Netocia hungarica [Herbst]) ...... 11 (Herbst) Medvedev 1952 10. Frons with one posterior seta on each side (Fig. Potosia fieberi Medvedev Medvedev 1952 2). Stridulatory area of mandibles broadly Potosia hieroglyphica Medvedev 1952 oval (Fig. 12). (Tropinota hirta [Poda]) . . . (Me´ne´trie`s) Potosia opaca (Fabricius) Mico´ and Galante 2002 ...... Tropinota squalida (Scopoli) Trichius Fabricius Van Edmen 1941, Medvedev 1952 10Ј. Frons with 3Ð5 posterior setae on each side Tropinota hirta (Poda) Golovjanko 1936, Medvedev 1952 (Fig. 28). Stridulatory area of mandibles Tropinota squalida This publication elongate-oval (Fig. 38) ...... (Scopoli) (L.) Medvedev 1969 ...... A. floralis (Fabricius) 11. Frons with one anterior seta on each side. Te- gilla with short setae and isolated long setae...... Netocia morio (Fabricius) Redescriptions of many Palaearctic Cetoniinae lar- 11Ј. Frons with 1Ð3 anterior setae on each side. vae (see Table 1 for a complete listing of Palaearctic Tegilla with many long setae and sparse larval descriptions) are needed before they can be short setae ....Netocia oblonga (Gory and included in our key. Taxa in brackets key out here, but Percheron) because of a lack of adequate information in the avail- able descriptions they cannot be keyed farther. 1. Labrum entire ...... 2 1Ј. Labrum trilobed ...... 4 2. Dorsum of last abdominal segment divided into two cushion-like areas by a median groove ...... Valgus hemipterus (Linnaeus) 2Ј. Dorsum of last abdominal segment without these areas ...... 3 3. Raster with palidium. Dorsum of mandibles with a longitudinal deep furrow ...... Gnorimus Le Pelletier and Serville 3Ј. Raster without palidium. Dorsum of mandibles without this furrow .....Trichius Fabricius 4. Legs bearing apical, short, subconical claws with two distal setae ...... Osmoderma eremita (Scopoli) 4Ј. Legs bearing cylindrical tarsungulus, rounded apically and with several setae ...... 5 Fig. 40. Life cycle of the studied species. March 2003 MICO´ AND GALANTE:CETONIINAE LARVAL DESCRIPTIONS 105

Table 2. Field Observations of Tropinota squalida and Oxythyrea funesta Third-instar Larvae and Immatures

Species Stage Resource Date Localities Leg. Tropinota squalida Immature Decaying shrub leaves 7 Dec 1997 Embalse de Celemõ´n (Cadiz, Spain) Mico´ (Fabricius) Immature Decaying shrub leaves 24 July 1997 Altura (Castello´n, Spain) Mico´ Immature Decaying shrub leaves 3 Jan 1998 Villar de Olalla (Cuenca, Spain) Larvae Oryctolagus cuniculus (L.) 2 April 1998 Pen˜ a Rubia (Villena, Spain) Mico´ latrine Larvae O. cuniculus latrine 26 April 1997 Benitachel (Alicante, Spain) Mico´ and Verdu´ Larvae O. cuniculus latrine 3 May 1997 Benitachel (Alicante, Spain) Mico´ and Verdu´ Larvae Old dung pat 6 Aug 1999 Palacio de Don˜ ana (Huelva, Spain) Verdu´ Oxythyrea funesta Larvae Manure 4 May 1996 Bo´ren (Le´rida, Spain) Mico´ and Verdu´ (Poda) Larvae Old dung pat 15 Aug 1999 (Cuenca, Spain) Mico´ and Verdu´ Downloaded from https://academic.oup.com/aesa/article/96/2/95/27977 by guest on 24 September 2021

Larval Biology cuniculus (n ϭ 3), decaying shrub leaves (n ϭ 3), and manure (n ϭ 1). All larvae reached the third-instar; Hurpin (1958) described the life cycle duration of however, only larvae that were given manure Þnished T. squalida and O. funesta as it related to temperature the life cycle. Adults of A. floralis (65.3% females and and estimated that the optimal temperature for larval 34.7% males) are attracted to ripe fruit and wine- development ranged between 20 and 25ЊC. Our results baited traps (n ϭ 700). Conversely, only females were showed that the life cycle of T. squalida was shorter captured in dung-baited traps (n ϭ 20). This fact than that of O. funesta under laboratory conditions, indicates that manure is an appropriate substrate for and both species overwinter as adults (Fig. 40). The female egg laying. larval development period lasted 6Ð7 wk for T. A. floralis is abundant in brushwood areas and in squalida and 8Ð9 wk for O. funesta. Both species have waste ground areas, and adults are attracted to the been reported from decaying vegetative matter and large ßowers of Carduiinae (Compositae) species manure (Molina 2001, Balachowsky 1962). However, (Puche Roca 1988, Mico´ and Galante 1998). Unfortu- it is the Þrst time that immature stages of Cetoniinae nately, these areas are shrinking because of the in- are reported from rabbit dung heaps. crease of urban development. Moreover, the decrease The European rabbit Oryctolagus cuniculus (Lin- of farming activity could threaten the Iberian popu- naeus) is abundant in the southeastern Iberian Pen- lations of A. floralis. insula. This species produces piles of dung heaps (la- trines) that constitute a stable, moist, and organic substrate for many Scarabaeoidea larvae (Galante and Acknowledgments Cartagena 1999, Mico´ and Galante 2003). T. squalida larvae frequently exploit this resource (Table 2), and We thank Jose´ R. Verdu´ and M. C. Cartagena for allowing us to study the cetoniine beetles collected in their dung- our Þeld observations showed that O. funesta larvae baited and wine-baited traps. Kate Burke checked an early were more commonly found feeding on manure (Ta- English version. The research was supported in part by the ble 2). However, laboratory data showed that both Spanish Ministry of Science and Technology, BOS 2000-0148 species were able to exploit the milled rabbit dung project. heaps and decaying leaves of shrubs and manure with a high survival rate. Larvae of T. squalida (n ϭ 91) and References Cited O. funesta (n ϭ 33) were reared from eggs under laboratory conditions. They fed on milled dung heaps Abdalla, E. F. 1991. Activity of the slaked lime mixed with of O. cuniculus (n ϭ 32, n ϭ 17, respectively), decaying cattle manure against the grubs of the scarabeid, shrub leaves (n ϭ 37, n ϭ 8), and on manure (n ϭ 22, Tropinota squalida (Scop.) (Scarabaeidae: Coleoptera). n ϭ 8). Mortality of grubs was always below 40% for Pakistan J. Zool. 23(1): 89Ð91. T. squalida, being especially low when using milled Balachowsky, A. S. 1962. Entomologie applique´ a` lÕagriculture. Cole´opte`res I. Masson et C ie (ed.). Paris, dung heaps of rabbits (10%). No mortality was ob- France. 1: 24Ð204. served in O. funesta grubs. Carne, P. B. 1951. Preservation techniques for scarabaeid Both species have been called rose chafers because and other insectÕs larvae. Proc. Linn. Soc. N.S.W. 76: of the feeding behavior of adults on rose buds and 26Ð30. other ornamental ßowers (Balachowsky 1962, Molina Donaldson, J. M. 1987. Descriptions of, and a key to, larvae 2001). The presence of organic manure near orna- of some South African Cetoniinae (Coleoptera: Scar- mental or cultivated areas during adult activity from abaeidae). Entomol. Mon. Mag. 123: 1Ð14. March to August (Fig. 40) could constitute a source of Galante, E., and M. C. Cartagena. 1999. Comparison of infestation by these species. Mediterranean dung beetles (Coleoptera, Scarabae- oidea) in cattle and rabbit dung. Environ. Entomol. 28(3): Under laboratory conditions, the larval develop- 420Ð424. ment period of A. floralis lasted 22 wk (Fig. 40) and, Golovjanko, Z. S. 1936. Les larves les plus communes des as occurs in many Cetoniini species (Mico´ et al. 2000), Cole´opte`res Lamellicornes de la partie ´enne de they overwinter as adults. A total of seven larvae lÕU.R.S.S. (tableaux analytiques de la faune de lÕU.R.S.S.). reared from eggs were given milled dung heaps of O. 20: 1Ð65. 106 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 2

Hurpin, B. 1958. Inßuence de certains facteurs e´cologiques Mico´, E., A.B.T. Smith, and M. A. Moro´n. 2000. New larval sur la ponte et le developement des ce´toines grises. En- descriptions for two species of Burmeister (Co- tomol. Exp. Appl. 1: 246Ð259. leoptera: Scarabaeidae: Cetoniinae: Cetoniini: Eupho- Janssens, A. 1960. Faune de Begique: insectes Cole´opte`res rina) with a key to the known larvae and a review of the Lamellicornes. Institut Royal des Sciences Naturelles de larval biology. Ann. Entomol. Soc. Am. 93(4): 795Ð801. Belgique, Belgique, Bruxelles. Mico´, E., J. R. Verdu´ , and E. Galante. 2001. Larval morphol- Klausnitzer, B. D., and F.-T. Krell. 1997. 6. U¨ berfamilie: ogy of some Anisopliini grain beetles with a key to their Scarabaeoidea. In B. Klausnitzer (ed.), Die Larven der larvae (Coleoptera: Scarabaeoidea: Rutelidae: Anomali- Ka¨fer Mitteleuropas. 3. Band. Teil 2. Gustav nae). Eur. J. Entomol. 98: 311Ð320. Fischer Verlag, Jena. Molina, J. M. 2001. Incidencia de Tropinota squalida (Sco- Korschefsky, R. 1940. Bestimmungstabelle der hau¨ Þgsten poli, 1783) (Coleoptera: Scarabaeidae) en el cultivo del deutschen Scarabaeiden larven. Arb. Physiol. Angw. En- ara´ndano en Huelva (Espan˜ a): problema´tica asociada a su tomol. Berlin-Dahlem. 7: 41Ð52. control. Zapateri Rvta. Aragon. Ent. 9: 93Ð98. Krikken, J. 1984. A new key to the suprageneric taxa in the Puche Roca, J. A. 1988. Datos ecolo´gicos y de distribucio´n beetles family Cetoniidae, with annotated lists of the de Cetoniinae (Coleoptera, Scarabaeidae) del litoral le- Downloaded from https://academic.oup.com/aesa/article/96/2/95/27977 by guest on 24 September 2021 known genera. Zoologische Verhandelingen 210: 4Ð71. vantino. Actas III Congreso Ibe´rico de Entomologõ´a 693Ð Medvedev, S. I. 1952. Larvae of Scarabaeid beetles of the 706. fauna of the USSR. Opredeliteli po faune SSSR 47, Ritcher, P. O. 1966. White grubs and their allies: a study of Moskva, Leningrad. North American Scarabaeoid larvae. Studies in Entomol- Medvedev, S. I. 1969. On the larva of Valgus hemipterus L. ogy no. 4. Oregon State University Press, Corvallis, OR. (Coleoptera, Scarabaeidae). Entomol. Rev. 48(1): 100Ð Van Emden, F. I. 1941. Larvae of British beetles. II. A key 101. to the Br. Lamellicornia larvae. Entomol. Mon. Mag. 77: Mico´, E., and E. Galante. 1998. The behaviour of Aethiessa 117Ð192. floralis (Fabricius, 1787) (Coleoptera: Scarabaeoidea: Ting, P. C. 1934. Back-crawling scarabaeid grubs (Potosia Cetoniidae) visiting Onopordum macracanthum Schous- afÞnis Andersch) intercepted in quarantine at San Fran- boe (Compositae). Elytron 12: 69Ð76. cisco. Mon. Bull. Calif. Dep. Agric. 23: 185Ð191. Mico´, E., and E. Galante. 2003. Larval morphology and bi- ology of four fruit chafers (Col., Scarabaeoidea: Cetoni- Received for publication 4 March 2002; accepted 22 July idae: Cetoniinae). Eur. J. Entomol. (in press). 2002.