Dtsch. Entomol. Z. 71 (1) 2024, 185-192 | DOI 10.3897/dez.71.113727 > PENSUFT. Gag MuseuM TOR BERLIN Larval morphology of a Palearctic Rutelini, Parastasia ferrieri (Coleoptera, Scarabaeidae), with discussions on their feeding habits Xiao-Yu Sun', Xu-Ming Dong?, Lu Jiang! 1 Key Laboratory of Economic and Applied Entomology of Liaoning Province, College of Plant Protection, Shenyang Agricultural University, Shenyang, Liaoning 110866, China https://zoobank. org/694F ED$& F-4B56-4CBD-95E3-DB50230DAED6 Corresponding author: Lu Jiang (jianglu@syau.edu.cn) Academic editor: Emmanuel Arriaga Varela # Received 5 October 2023 Accepted 4 April 2024 Published 2 July 2024 Abstract Rutelini is one of the largest tribes of Rutelinae, widely distributed but primarily in the New World. Recently, both larvae and adults of Parastasia ferrieri had been discovered in Liaoning Province of northeastern China from the Palearctic realm. The third-instar larvae of P. ferrieri were described using light and scanning electron microscopy in order to discover more morphological characters for larval taxonomy. The larvae of P. ferrieri exhibit remarkable features, including four protuberances on labrum, no helus on epi- pharynx, two scissorial teeth on each mandible, five stridulatory teeth plus a blunt protuberance on each maxilla, and the obtuse claws on the thoracic legs. The correlation between morphological features and feeding habits is briefly discussed. Key Words saproxylic, shinning leaf chafer, SEM, ultramorphology, white grub Introduction Rutelinae, commonly known as shining leaf chafers, are so named due to the bright colors of most of their species and their adults feeding on plant leaves (Jameson and Rat- cliffe 2002). Rutelinae encompass more than 4,000 spe- cies distributed among 235 genera (Krajctik 2007; Dietz et al. 2023). These species are further categorized into sev- en tribes: Adoretini, Alvarengiini, Anatistini, Anomalini, Anoplognathini, Geniatini, and Rutelini (Bouchard et al. 2011). On tribal level, in fact, Rutelinae exhibit great diver- sity on adult morphology, geographical distribution, diur- nal/nocturnal rhythms, phototaxis, or even larval feeding habits (Frew et al. 2016; Slipinski and Lawrence 2019). Larvae of Rutelinae are commonly referred to as white grubs, exhibiting diverse feeding and living habits at the tribal level (Ritcher 1966; Johnson and Rasmann 2015; Frew et al. 2016). The larvae are described as root-feeding in some Anoplognathini, Geniatini, Adore- tini and most of the Anomalini species (Habeck 1963; Ritcher 1966; McQuillan 1985; Fuhrmann 2013; Fang et al. 2018). However, the larval feeding habits remain generally unknown in Anatistini and Alvarengiini (Par- do-Locarno et al. 2006; Fuhrmann 2013; Rodrigues et al. 2017). Rutelini larvae predominantly exhibit saprox- ylic habits, characterized by their consumption of de- caying wood, vegetation, roots, or other organic matter (Ritcher 1948), distinguishing them from the majority of other root-attacking larvae in the Anomalini (Ritcher 1966; Zhang 1984). It is unwise to generalize the ecolog- ical role of a specific Rutelinae species until their larvae have been accurately identified (Slipinski and Lawrence 2019). Unfortunately, larval identification has been prov- en to be extremely challenging, with fewer than 2% of all known species having been described (Newton 1990; Sipek 2010; Lawrence et al. 2011). The Rutelini encompass approximately a thousand species assigned into 93 genera, with a primary presence in the New World (Jameson 1997; Slipinski and Law- rence 2019). They exhibit an array of distinctive morpho- logical features, including enlarged mandibles, prominent thoracic horns, expanded hindlegs, and striking metallic coloration (Jameson and Ratcliffe 2002). Notably, Old World Rutelini typically display significant sexual dimor- Copyright Xiao- Yu Sun et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 186 phism, with male adults possessing well-developed tho- racic horns or mandibles (Jameson 1997; Slipinski and Lawrence 2019). Regarding their immature stages, the larval morphology of Rutelini has been described for a limited subset, totaling 39 species in 24 genera (Ritcher 1948, 1966; Jameson and Moron 2001; Albertoni et al. 2014; Carvalho et al. 2019; Barria et al. 2020; Barria et al. 2021; Lugo-Garcia et al. 2023). Parastasia is one of the largest genera within Rutelini, comprising approximately 105 species worldwide (Wada 2015; Zhao 2019; Hongsuwong et al. 2022). However, un- like many other Rutelini beetles, the adults of Parastasia do not display enlarged structures or vibrant metallic coloration (Zhao 2019). Recently, both larvae and adults of Parastasia Jerrieri were observed in Liaoning Province of northeastern China, belonging to the Palearctic realm, similar to the pre- vious records in Korea (Kim 2014). Larvae of P. brevipes have been morphologically described and reported to feed on dead wood (Ritcher 1966). However, most of the other larvae of Parastasia are not adequately described hitherto. In this study, third instar larvae of P. ferrieri were ob- tained through rearing. Their morphology is described us- ing light and scanning electron microscopy in order to bet- ter understand the morphological diversity within the group and help with the identification of larvae in this genus. Materials and methods Insect collection and rearing Larvae of P. ferrieri were collected from Qipanshan For- est Park in Shenyang City, Liaoning Province of north- eastern China, in late October 2019. A total of 13 adult P. ferrieri beetles (Fig. 1A, B) were obtained through rearing in the following May. Paired adults were kept in plastic boxes filled with moist, fermented sawdust (Bee- tle-Password Company, Shenyang, China), and a decayed wood log off-cut was provided to facilitate potential bor- ing and egg-laying. Third instar larvae were collected from the sawdust in the following September. Light and scanning electron microscopy To conduct morphological observations, a total number of ten larvae were fixed in Dietrich’s solution (formalin: 95% ethanol: glacial acetic acid: distilled water = 6: 15: 1: 80, v/v), which was heated up to 70 °C and then left to cool naturally for 12 h under a fume hood before being preserved in 75% ethanol (Jiang and Hua 2015). Photographs were captured using a SONY ILCE-7RM4 digital camera. Scanning electron microscopy (SEM) was employed to examine third instar larvae. These larvae were dissected and examined in 75% ethanol using a Leica EZ4HD Stereoscopic Zoom Microscope. After a two-min- ute ultrasonic cleaning and two rinses in 75% ethanol, they were prepared for SEM. Dissected organs underwent seri- al dehydration using graded ethanol, followed by replace- dez.pensoft.net Xiao-Yu Sun et al.: Larval morphology of Parastasia ferrieri ment with tertiary butanol. They were then subjected to freeze-drying for 3 hours, sputter-coated with gold, and examined under a Hitachi S—3400N scanning electron mi- croscope (Hitachi, Tokyo, Japan) at 5 kV. Nomenclature for larval morphology follows (Ritcher 1966). Voucher specimens of both adults (Fig. 1 A—E) and lar- vae (Fig. 1F) were deposited at the Entomological Muse- um of Shenyang Agricultural University (SYAU). Results General larval morphology Larvae are of typically scarabaeiform shape, bearing three pairs of thoracic legs on the C-shaped body (Fig. 1F). The larval trunk 1s generally white in addition to the yellowish head capsule, thoracic legs and spiracles. The prothoracic spiracles are C-shaped, 0.41+0.05 mm (N = 20) in length. Spiracles on the anterior six abdominal segments are simi- lar in sizes, approximate 0.22 + 0.05 mm (N= 20) in length. Whereas, the spiracles on the seventh and eighth abdomi- nal segments are comparative larger, 0.27 + 0.05 mm (N= 20) and 0.28 + 0.05 mm (N = 20) respectively. Head The head capsules are 3.1 + 0.15 mm (N = 10) in width. The larval head displays symmetrical adornment, boast- ing a total of 14 pairs of setae. These include two pairs aligned vertically on the clypeus, five pairs in the frontal region (comprising two pairs of posterior frontal setae and three pairs of anterior frontal setae), three pairs on the vertex, and four pairs in the genal area (Fig. 2A). The antenna is elongated and slender, comprising four segments, with the second segment being the longest (Fig. 2C). The basal two segments of the antenna are adorned with one and four setae, respectively. The third segment is smooth, featuring two to four oval dorsal sensory spots. The distal segment of the antenna of conical shape with seven sensilla basiconica at its apex (Fig. 2D). Mouthparts The mouthparts are of a biting-chewing type, consisting of a labrum, a pair of mandibles, and a maxilla-labia complex. Labrum exhibits symmetry and 1s slightly wider than it is long. The outer surface of the labrum displays sym- metrical features, including four prominent protuberanc- es and seven pairs of setae (Fig. 2B). The distal margin of the labrum is equipped with numerous sensory setae pointed distally (Fig. 3A). Epipharynx, membranous, situated on inner surfaces of labrum and clypeus. Epipharynx is further divided into distinct functional areas (Fig. 3A), including a heptomeron at the apex, a pair of plegmatium on the lateral margin, nesium in the central portion, and glabrous gymnoparia on Dtsch. Entomol. Z. 71 (1) 2024, 185-192 187 A B — 0.5 mm “ 0.5mm z= 0.5mm 2.0 mm Figure 1. Adults and a larva of Parastasia ferrieri. A. Male adult, dorsal view; B. Male adult, ventral view; C. Male genitalia, dorsal view; D. Male genitalia, lateral view; E. Male genitalia, ventral view; F. Third instar larva. Figure 2. Larval head of Parastasia ferrieri. A. Head; B. Labrum; C. Antenna; D. Sensilla on the apex of antenna. AFS, anterior frontal seta; AT, antenna; CLP, clypeus; DES, dorsoepicranial setae; ES, epicranial stem; F, frons; FCS, frontoclypeal suture; LB, labrum; LP, labral protuberance; PFS, posterior frontal seta; SB, sensillum basiconicum. dez.pensoft.net 188 Figure 3. Larval epipharynx of Parastasia ferrieri. A. Epipharynx; B. Magnification of haptomerum; C. Magnification of ple- Xiao-Yu Sun et al.: Larval morphology of Parastasia ferrieri _— gma; D. Basal part of the epipharynx. ACP, acanthoparia; GP, gymnoparia; PL, plegma; PE, pedium; SP, sclerotized plate; S, sensillum; Z, zygum. both sides (Fig. 3A). The heptomeron exhibits a slight cur- vature and is composed of four minute sensilla basiconica, lacking helus (Fig. 3B). The plegmatium consists of eight acanthoparia, intricately intertwined with an equal number of plegma (Fig. 3C). The chaetoparia displays asymme- try and is composed of numerous sensilla chaetica, with a higher density on the right side than on the left. Adjacent to the right acanthoparia, the sensory nesium is situated, fea- turing four micro sensilla basiconica at its apex (Fig. 3D). The paired mandibles are heavily sclerotized, bear- ing two anterodorsal setae and a row of 11 setae on the lateral surface (Fig. 4A, B). The paired mandibles each bear a shuttle-shaped stridulatory area on ventral surface (Fig. 4C, D). The mandibular incisor is equipped with two apical teeth curved inward. The molar region exhib- its asymmetry, featuring a prominent molar tooth accom- panied by an acia on the left and a group of four ridged molar teeth on the right (Fig. 4C, D). The ventral process is relatively narrow on the left mandible (Fig. 4D), while it is generally wider on the right mandible (Fig. 4C). The maxillae, labia, and hypopharynx are fused to- gether to form a structural complex (Fig. 4E). Each of the paired maxillae comprises a cardo, a stipes, a maxillary palp, and a lobe fused from the galea and lacinia (Fig. 4E). The maxillary stridulatory area comprises a row of five stridulatory teeth and an anterior truncate process (Fig. AF). The maxillary palp consists of four segments, with dez.pensoft.net the longest distal segment bearing a digitiform organ on its lateral surface and ten sensilla basiconica on its apex (Fig. AE). The labia comprise a mentum, a prementum, a pair of two-segmented labial palps, and a sensory glossa (Fig. AE). Dorsally, the glossa is adorned with numerous sen- silla (Fig. 4E). The hypopharynx 1s specialized, forming a hypopharyngeal sclerome that bears a sclerotized truncate process, two tufts of microtrichia on the left, and a pair of membranous lobes on the lateral margin (Fig. 4F). Thoracic legs Each of the thoracic legs is composed of five segments: coxa, trochanter, femur, tibiotarsus, and a distal single claw (see Fig. 5A, B). The coxa exhibits a long and slen- der shape. The trochanter possesses a slight curvature. The femur is covered with numerous setae on its surface. The tibiotarsus is originally a fusion of the tibia and tarsus, and it bears medium-sized setae on its dorsal, ventral, and lateral sides. The distal claw is heavily sclerotized, fea- turing an obtuse distal end and bearing three short setae. The respiratory plate is composed of numerous minute openings that encircle the oval bulla, along with a slightly curved spiracular slit (refer to Fig. 5B). The prothoracic spiracles are slightly larger than the abdominal ones. The abdominal spiracles are similar in size (see Fig. 1). Dtsch. Entomol. Z. 71 (1) 2024, 185-192 189 —, 4 47 200 um : on oe Figure 4. Mandibles and maxillae of Parastasia ferrieri. A. Left mandible, dorsal surface; b Right mandible, dorsal surface; C. Right mandible, ventral surface; D. Left mandible, ventral surface; E. Maxillae and labia, ventral surface, insert showing the magnification of maxillary palpus; F. Maxilla, labia, and hypopharynx, dorsal surface. AC, acia; CAR, cardo; DC, dorsal carina; DO, digitiform organ; GL, glossa; HS, hypopharyngeal sclerome; IC, incisor; LL, lateral lobe; LP, labial palpus; M, mola; MP, max- illary palpus; PC, penicillus; PM, postmentum; PR, precoila; PRM, prementum; SC, scobis; ST, stridulatory teeth; STA, stridulatory area; SP, stipes; TP, truncate process; VP, ventral process. Raster The raster is furnished with paralleled palidia, each com- posed of a longitudinal patch of mesal directed pali, sur- rounding a prominent septula. The palidia are submerged in a large number of setae, which are slightly longer and distributed at the lateral region. Discussion In this study, larvae of P. ferrieri were described using scanning electron microscopy for the first time. The larvae of P. ferrieri are remarkable for the following structures: bearing four protuberances on labrum; no helus on epi- pharynx; two scissorial teeth on each mandible; five strid- ulatory teeth and a blunt protuberance on each maxilla; ob- tuse claws on thoracic legs, and 12 pairs of pali on raster. By the combination of these characters, larvae of P. ferri- eri can be readily distinguished from most of the other lar- vae in Rutelini (Ritcher 1948, 1966; Jameson and Moron 2001; Albertoni et al. 2014; Carvalho et al. 2019; Barria et al. 2020; Barria et al. 2021; Lugo-Garcia et al. 2023). The labrum exhibits a wide range of morphological features within families or subfamilies of Scarabaeidae (Ritcher 1966; Grebennikov and Scholtz 2004). The labrum is typically fan-shaped in Passalidae and Luca- nidae (Hayes 1929; Qu et al. 2019), trilobed in Aphodi- inae, Scarabaeinae, some Cetoniinae, and Pleocomidae (Grebennikov and Scholtz 2004; Li et al. 2019; Dong et al. 2020), or bearing an apical protuberance in some Seri- cinae (Sipek and Ahrens 2011). In the genus Apogonia (Melolonthinae), the labrum forms a dorsal ridge (Jia et al. 2023), while in some Hybosoridae, the labrum possesses serrations (Grebennikov and Scholtz 2004; Grebennikov et al. 2004). The labrum has varying numbers of setae or display glabrous, wrinkled, or humped dorsal surfaces in different lineages (Qu et al. 2019; Jia et al. 2020; Jia et al. 2021; Jia et al. 2023). In this study, the larval labrum of P. ferrieri is atypical for the presence of four protuberances on its dorsal surface, which is not mentioned in the con- generic larvae of P. brevipes (Ritcher 1966). Previous comprehensive studies (Ritcher 1966; Zhang 1984; Sawada 1991) have indicated that heli on the epi- pharynx often serve as valuable taxonomic characteris- tics for larval identifications in Scarabaeidae (Fang et al. 2018; Jia et al. 2021). In Rutelinae, the larval epipharynx is equipped with two to four heli in Anomalini (Mico et al. 2003), six to eight in Anoplognathini (Neita-Moreno and Moron 2017), six to nine in Adoretini (Fang et al. 2018), dez.pensoft.net 190 Xiao-Yu Sun et al.: Larval morphology of Parastasia ferrieri Figure 5. Thoracic legs, spiracles and raster of Parastasia ferrieri. A. Prothoracic leg, lateral view, B. Prothoracic spiracle; C. Raster; D. Magnification of the anterior palidia; E. Magnification of the hamate seta. BU, bulla; CL, claw; CX, coxa; FM, femur; RSP, respiratory plate; SS, spiracular slit; TT, tibiotarsus; TR, trochanter. or entirely absent in Geniatini and Rutelini (Jameson and Moron 2001; Fuhrmann 2013). In this study, the larval epipharynx of P. ferrieri is devoid of heli, consistent with previous descriptions in Rutelini (Ritcher 1966; Jameson and Moron 2001; Barria et al. 2020; Barria et al. 2021; Lugo-Garcia et al. 2023). Mandibles, being the most heavily sclerotized struc- tures on mouthparts, are structurally correlated with feeding habits (Zhang 1984). The larval mandibular in- cisor typically varies: straight and apically sharp in some wood-consuming species of Lucanidae and Passalidae (Hayes 1929; Katovich and Kriska 2002; Richards and Spencer 2014; Qu et al. 2019), sharp but inwardly curved in dung decomposers within Aphodiinae and Scarabaein- ae (Jerath 1960; Edmonds and Halffter 1978; Frolov 2009; Li et al. 2019), or blade-like and specialized to form a cutting edge in phytophagous Melolonthinae (Jia et al. 2020; Jia et al. 2021; Jia et al. 2023). Within Rutelinae, the mandibular incisors are typically blade-like in some phytophagous larvae of Anomalini and Adoretini (Mico et al. 2003; Fang et al. 2018), and are apically sharp and curved in some saproxylic larvae of Rutelini (Jameson and Moron 2001). Within Rutelini, larval mandibles are usually asymmetric, equipped with three teeth on the left and two teeth on the right (Jameson and Moron 2001; Al- bertoni et al. 2014; Carvalho et al. 2019). In this study, dez.pensoft.net however, the larval mandibular incisor is symmetric, each bearing two scissorial teeth, similar to the previously de- scribed larvae of P. brevipes (Ritcher 1966). The claws of thoracic legs show considerable morpho- logical diversity across lineages (Zhang 1984). In most species of Melolonthinae and Cetoniinae, the claw is typi- cally curved, sharp, and adorned with paired setae (Sousa et al. 2018; Jia et al. 2020; Jia et al. 2021; Jia et al. 2023). Conversely, it 1s usually blunt in some wood-decompos- ing species within Lucanidae (Katovich and Kriska 2002; Qu et al. 2019) and remnant in certain dung-feeding lar- vae of Geotrupidae (Grebennikov and Scholtz 2004). In Rutelinae, thoracic claws are typically sharp in Adoretini (Fang et al. 2018), Geniatini (Pardo-Locarno et al. 2006; Fuhrmann 2013), and Anomalini (Mic6 and Galante 2005). However, they display morphological heterogene- ity among the pro-, meso-, or metathoracic legs in some species of Rutelini (Carvalho et al. 2019). In this study, the claws of P. ferrieri are generally blunt on all three pairs of thoracic legs, congruent with other saproxylic larvae in Lucanidae (Richards and Spencer 2014; Qu et al. 2019). Rutelinae usually attract attention due to their exquisite adult appearances or the economic losses caused by their larval stages (Jameson and Ratcliffe 2002). In the Pale- arctic realm, particularly in northeastern China, larvae of Rutelinae are frequently recognized as agricultural pests, Dtsch. Entomol. Z. 71 (1) 2024, 185-192 because they mostly belong to the phytophagous Anoma- lini or Adoretini (Zhang 1984; Sawada 1991). This scar- city record of P. ferrieri may be attributed to their small, nocturnal adults, or the fact that their larvae never attack living organs of crops or trees. Given their significance of biogeographical distribution, the larvae of P. ferrieri warrant increased attention for conservation purposes. Author contributions Conceived and designed the experiments: LJ, XYS. Per- formed the experiments: XYS, XMD. Analyzed the data: XYS and LJ. Wrote the paper: XYS, XMD and LJ. Acknowledgements We are grateful to Ming-Zhi Zhao and Zi Shan for their kind help during our larval collecting and rearing periods. Our special thanks go to Dr. Yuan-Yuan Lu and Dr. Valentina Filippini for their valuable suggestions on our earlier draft. This research was financially supported by the National Natural Science Foundation of China (grant no. 32370470 and 31702036), China Postdoctoral Science Foundation (grant no. 2020M680982), Natural Science Foundation of Liaoning Province (2021-MS-230), Scientific Research Project of Education Department of Liaoning Province (LJKZ0641), Science and Technology Planning Project of Liaoning Province (grant no. 1618214601077) and Sci- entific Research Foundation for the Introduced Talent of Shenyang Agricultural University (grant no. 880417008). References Albertoni FF, Fuhrmann J, Ide S (2014) Lagochile emarginata (Gyllenhal): Morphology of immature and imago, and biological records (Coleoptera, Scarabaeidae, Rutelinae). 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