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Larval thoracic legs and prolegs of Panorpidae. (A) and (E) Dicerapanorpa magna; (B) and (F) Cerapanorpa nanwutaina; (C) and (G) Panorpa curva; (D) and (H) Neopanorpa lipingensis; (A-D) prothoracic legs, arrows point to tibial lobes; (E-H) prolegs, arrow heads point to morphologically reduced prolegs on the anterior four abdominal segments. Scale bars: (A-D) = 40 μm; (E)-(H) = 200 μm.

Larval thoracic legs and prolegs of Panorpidae. (A) and (E) Dicerapanorpa magna; (B) and (F) Cerapanorpa nanwutaina; (C) and (G) Panorpa curva; (D) and (H) Neopanorpa lipingensis; (A-D) prothoracic legs, arrows point to tibial lobes; (E-H) prolegs, arrow heads point to morphologically reduced prolegs on the anterior four abdominal segments. Scale bars: (A-D) = 40 μm; (E)-(H) = 200 μm.

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Article
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Insects are the most diverse group of organisms in the world, but how this diversity was achieved is still a disputable and unsatisfactorily resolved issue. In this paper, we investigated the correlations of habitat preferences and morphological traits in larval Panorpidae in the phylogenetic context to unravel the driving forces underlying the evo...

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Context 1
... thoracic legs are four-segmented, each consisting of a stout coxa, a cylindrical elongate femur, a slender tibia, and a short pointed hirsute tarsus. A remarkable triangular tibial lobe is present posteromesally on the tibiae of Dicerapanorpa, Cerapanorpa, and Panorpa ( Fig. 3A-C), but absent in Neopanorpa (Fig. ...
Context 2
... thoracic legs are four-segmented, each consisting of a stout coxa, a cylindrical elongate femur, a slender tibia, and a short pointed hirsute tarsus. A remarkable triangular tibial lobe is present posteromesally on the tibiae of Dicerapanorpa, Cerapanorpa, and Panorpa ( Fig. 3A-C), but absent in Neopanorpa (Fig. ...
Context 3
... unsegmented ventral prolegs are present on the first eight abdominal segments. These prolegs are well developed in Dicerapanorpa, Cerapanorpa, and Panorpa ( Fig. 3E-G), but remarkably reduced on the anterior four pairs in Neopanorpa (Fig. ...
Context 4
... unsegmented ventral prolegs are present on the first eight abdominal segments. These prolegs are well developed in Dicerapanorpa, Cerapanorpa, and Panorpa ( Fig. 3E-G), but remarkably reduced on the anterior four pairs in Neopanorpa (Fig. ...
Context 5
... dorsal annulated processes are very similar between the species of each genus, but vary in length among the genera. These dorsal processes are very prominent in Dicerapanorpa, median-sized in Cerapanorpa, and relatively shorter in Panorpa and Neopanorpa (Fig. ...

Citations

... Detritivorous larvae like maggots contribute to nutrient recycling by decomposing organic matter while soil-dwelling larvae like those of lacewings and antlions are adept ambush predators with specialized morphological features for hunting. This wide range of larval forms underscores the intricate adaptations of insects to diverse habitats and ecological niches, highlighting their significant contributions to ecosystem dynamics and biodiversity within the insect realm ( Jiang et al., 2019). ...
... Both Neopanorpa and Panorpa larvae are euedaphic, living in the soil most of their life time. However, Panorpa larvae emerge to the surface, likely to forage at night, to a significantly greater degree than Neopanorpa [75]. We speculate that Panorpa larvae would have a much greater opportunity for survival at higher elevations and also during cooler seasons with less competition for food as well as less predation, where, and when, ants are less abundant. ...
... This phenomenon is hypothesized to be linked to the biological characteristics of the larval stage in Panorpidae scorpionflies. Specifically, Panorpidae are known to deposit their eggs in moist soil [18], and their larvae exhibit reduced sclerotization along with specialized morphological traits [75], adaptations that are conducive to surviving and foraging within humid soil environments. Similar adaptations, associated with diminished desiccation tolerance, have been observed in other insects, such as termites [76]. ...
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Ecological analyses of the small and lesser-known insect order Mecoptera in Thailand are presented. Specimens were collected monthly over a period of 12 consecutive months, using both Malaise and pan traps, from 29 sampling sites located in 18 national parks throughout Thailand. A total of 21 species in four genera were identified from 797 specimens, including Panorpa (1 species), Neopanorpa (18 species), Bittacus (1 species), and Terrobittacus (1 species), with the latter genus representing a new genus record to Thailand. Neopanorpa harmandi, N. siamensis, N. byersi, and N. malaisei were the most abundant species, representing 27.4%, 11.3%, 10.3% and 8.8% of the total specimens, respectively. The species with the highest frequency, as indicated by the high percentage of species occurrence (%SO), was N. siamensis (51%), followed by N. byersi (34%), N. harmandi (34%), N. spatulata (27%), and N. inchoata (27%). Eleven species (52%) exhibited specific regional occurrences. N. tuberosa and N. siamensis had the widest distribution, being found in almost all regions except for western and southern regions for the first and second species, respectively. The seasonal species richness of Mecoptera was high during the rainy season in the northern, northeastern, central, eastern, and western regions, with the highest richness observed in July (15 species), followed by the hot (10 species) and cold seasons (7 species), while there was no significant difference in species richness between seasons in the southern region. Multiple regression models revealed a negative association between species richness and abundance of Mecoptera with both elevation and temperature, and a positive association between rainfall and species evenness. It is predicted that climatic changes will have a detrimental effect on the mecopteran community. The results of this study enhance the understanding of the ecological aspects of Mecoptera, offering crucial insights into its biodiversity and distribution, which are vital for conservation and forest management.
... However, the information of pre-imaginal stages is quite rare. It is estimated that only about 2% of holometabolous larvae are well known (Newton 1990, Jiang et al. 2019. Extant scorpionflies with nine families have three general types of larvae: eruciform (caterpillar-shaped) larvae, characteristic of two small relict families from Australia (Apteropanorpidae and Choristidae), and two of the largest and most species-rich contemporary families (Panorpidae and Bittacidae); scarabaeiform larvae (Boreidae and Panorpodidae); larvae of Nannochoristidae are the only ones that are aquatic, and represent a completely different type of morphology. ...
... The Eocene larva is without a doubt specific for Panorpidae, based on the form of the labial palp, as well as the shape of the dorsal annulated processes and the ventral prolegs of the abdominal segments (Cai and Hua 2009, Chen and Hua 2011, Jiang and Hua 2013, 2015a, Ma et al. 2014, Jiang et al. 2019, Liu et al. 2021). This highly numerous and diverse family includes 10 extant genera and one exclusively fossil. ...
... The descriptions and graphical documentation of larvae in Furcatopanorpa Megapanorpa Wang andHua, 2018 are rare and superficial Hua 2011, Wang andHua 2018). Recent researches have demonstrated significant morphological variations among larvae of various panorpids, particularly in terms of head capsules, thoracic legs, abdominal prolegs, spiracles, and chaetotaxy ( Jiang et al. 2019). The larvae of Neopanorpa van der Weele, 1909(van der Weele 1909 are the most distinct among extant Panorpidae, which are distinguished by the presence of the furrows on the head capsule, the shape of the antenna, and the absence of a tibial lobe Hua 2011, Jiang andHua 2015b). ...
Article
So far, there has been no information of the pre-imaginal stages in the well-preserved fossil record of the Mecoptera. The first and well-preserved mecopteran larva was discovered in Eocene Baltic amber. The application of synchrotron X-ray microtomography enabled the reconstruction of the body structure with high accuracy, providing a comprehensive set of morphological data that classical stereoscopic microscopy could not capture. The larva is eruciform, with distinct segmentation of the body. All the most important morphological structures are documented, including the chaetotaxy system. Shape of antenna and annulated processes on the abdominal segments point to the third/four stage of development. There is no doubt that the larva belongs to Panorpidae, the most abundant family of extant Mecoptera. The morphological characters point to the closest affinity to larvae of Cerapanorpa and Panorpa but the fossil larva cannot be assigned to any extant genus of Panorpidae. Based on the results of synchrotron X-ray microtomography, it is possible to discuss the assignement of this larva to a lower systematic level than the family, and draw conclusions about the nature of the habitat preferences of Eocene panorpid.
... The Mecoptera lies in the superorder Antliophora in the Holometabola and consists of nine extant families (Byers & Thornhill, 1983;Misof et al., 2014;Willman, 1987). The abdominal prolegs of the larvae exhibit a considerable diversity among these families, with a wide range of variation in both segmental arrangement and number (Table S1; Jiang et al., 2019). The larvae are eruciform in Panorpidae, Bittacidae, Choristidae and Apteropanorpidae, possessing eight pairs of prolegs on abdominal segments 1-8 (A1-A8), but are scarabaeiform in Panorpodidae, which bear several conspicuous unpaired lobe-like processes on the mid-ventral line of A3-A6 or A2-A8 (Byers, 1997;Evans, 1942;Jiang & Hua, 2015;Liu et al., 2021;Suzuki, 1990). ...
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The abdominal appendages of larval insects have a complex evolutionary history of gain and loss, but the regulatory mechanisms underlying the abdominal appendage development remain largely unclear. Here, we investigated the embryogenesis of abdominal prolegs in the scorpionfly Panorpa liui Hua (Mecoptera: Panorpidae) using in situ hybridization and parental RNA interference. The results show that RNAi‐mediated knockdown of Ultrabithorax ( Ubx ) led to a homeotic transformation of the first abdominal segment (A1) into the third thoracic segment (T3) and changed the distributions of the downstream target Distal‐less ( Dll ) expression but did not affect the expression levels of Dll . Knockdown of abdominal‐A ( abd‐A ) resulted in malformed segments, abnormal prolegs and disrupted Dll expression. The results demonstrate that the gene Ubx maintains an ancestral role of modulating A1 appendage fate without preventing Dll initiation, and a secondary adaptation of abd‐A evolves the ability to specify abdominal segments and proleg identity. We conclude that changes in abdominal Hox gene expression and their target genes regulate abdominal appendage morphology during the evolutionary course of holometabolous larvae.
... On one hand, larvae are considered feeding devices that turn smaller embryos into larger individuals (Zacharuk and Shields 1991;Hall and Wake 1999). On the other hand, larvae exhibit diverse relationships with host plants, natural enemies and ecological factors in their relatively longer living periods (Ruxton et al. 2005;Jiang et al. 2019). In fact, in peculiar situations, larvae are more likely to attract attention from biologists than are the corresponding adults, even in the most famous insect lineage, butterflies (Igrashi 1984;Stehr 1987;Hasenfuss 1999;Casagrande and Mielke 2005). ...
Article
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Papilionidae, swallow-tailed butterflies, attract the attention of biologists for the importance of their ecological and evolutionary study. Papilionidae are peculiar among Lepidoptera for their larvae bearing a Y-shaped eversible osmeterium on the prothorax. However, morphological study of the larvae of Papilionidae, especially in terms of the osmeteria, are far from satisfactory. In this study, larval ultramorphology of Sericinus montela was described using scanning electron microscopy after a novel method of sample preservation. Larvae of S. montela are peculiar for the 15 teeth on the incisor, the triordinal mesoseries plus biordinal lateroseries of crochets on the anal prolegs, the urticating acanthae on secondary setae, and the micro openings and surrounding microtrichia on the osmeterium. The method of sample preservation, larval morphological characters and correlated defensive strategies are compared and briefly discussed.
... Certain mechanisms have been developed in the larvae to reduce competitive relations. For example, in the soil, Panorpidae larvae have not only different spatial niches (they are kept in various soil layers), but also are separated by the time of their activity (Jiang et al., 2019). ...
Article
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The study of the insect distribution in geographical areas is relevant since it is important in terms of understanding the global trend of biodiversity decline. The paper presents the results of a study on the distribution of six species of Panorpidae (Mecoptera), carried out in 2008, 2009, 2011, 2015, 2017–2020. One part of data was collected by the authors. Other material was provided by colleagues from 11 regions in Russia. In European Russia, six species of Panorpa are reliably known, namely Panorpa alpina, P. cognata, P. communis, P. germanica, P. hybrida, and P. vulgaris. The most common and frequently encountered species are P. communis (in 21 regions), P. hybrida (in 12 regions), P. vulgaris (in 11 regions), and P. cognata (in 11 regions). It is assumed that all studied species can be found in other regions of European Russia as a result of further investigations. Among the studied species, P. alpina and P. germanica are the rarest species, recorded from two and one regions, respectively. Panorpa vulgaris was found for the first time in Russia.
... Panorpids have eruciform larvae with compound eyes and bear long setae that tend to trap soil. They have similar biology with four instars of larvae dwelling in the soil and feeding on debris (Mampe & Neunzig 1965, Boese 1973 were found to be nocturnal coming to the surface at night to feed, while only a comparatively small fraction of Neopanorpa were found to venture to the surface at night (Jiang, et al. 2019). We presume this behavior might be an adaptation to competition with ants, which dominate tropical regions. ...
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The first Panorpa was recently discovered in Thailand even though the genus had not been thought tooccur south of the Himalaya. We now add a second new species, Panorpa apscisacera n. sp., also fromNorthern Thailand, which has nearly immaculate pale-yellow wings with only a pronounced stigma. The malehas a blunt, vestigial bifid anal horn, and thin, bare ventral parameres. Additionally, we describe Neopanorpaappendicema n. sp. with mesad-curled male hypovalve apices and large, basomesal lobes, a relatively largespecies, N. inchoata n. sp., with overlapping elliptical hypovalves and incomplete apical bands, N. setosiloban. sp. with huge bristly lobes on the mesal margin of the male gonostyles, and N. mandangensis n. sp.with a narrow 3rd tergum process extending to the posterior margin of the 4th tergum. Neopanorpa nielseniByers, 1965, previously known from northern Vietnam and southern China, has recently been discovered innorthern Thailand. Bittacus leptocaudus Byers, 1965, known from a lone male has been rediscovered andlocality details and photo of the habitat provided. These bring the number of species of Mecoptera occurringin Thailand to 22 (19 Neopanorpa, two Panorpa, and one Bittacus) and the number of Mecoptera speciesknown from Indochina to 56 (48 Neopanorpa, three Panorpa, 4 Bittacus, and one Bicaubittacus). A key tothe species of Mecoptera in Thailand with illustrations is provided. DNA was extracted from specimens andthe DNA Barcode, mitochondrial cytochrome oxidase I (COI) gene fragment sequenced and analyzed for 19of the 22 species of Thai Mecoptera. The results are figured as a Neighbor-Joining tree.
... Occasionally, they also feed on fresh vegetative materials such as nectar, petals, and pollen (Palmer 2010). Larvae generally remain in the soil, although some species spend their larval stage on the surface of the soil (Jiang et al. 2019). Most adults have an elongated head with slender, chewing mouthparts near the tip of a stout beak. ...
Article
The first annotated checklist of Mecoptera of Ukraine is presented. Altogether, 11 species of scorpionflies are known with a confirmed or probable occurrence in Ukraine. This list is based on previously published data, as well as on recently collected materials and specimens housed in the collection of the State Museum of Natural History NAS Ukraine, Lviv. This collection belongs to the oldest one in Ukraine and comprises specimens collected by Józef Dziędzielewicz, the founder of mecopteran investigations in Eastern Europe in the 19 th and 20 th centuries. For each listed species, its distribution in Ukraine is given together with notes on the history of investigations, and synonymies of the respective taxa. Detailed information on the distribution of selected species from the Red Book of Ukraine is presented.
... Morphological variety in insects occurs as a result of environmental variables such as food and climatic availability, geographic position, and the existence of natural enemies, as well as the process of natural selection [29]. Morphological diversity occurs mostly in the shape and size of bodily organs in insects [30]. ...
Article
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Spodoptera exigua (Hübner, 1808) (Lepidoptera: Noctuidae) is a significant agricultural crop pest in Indonesia, causing significant economic losses in recent years. This species’ ability to survive on a wide variety of host plants provides an adaptive advantage for survival in the environment, which is facilitated by its high mobility, fecundity, and capability to acquire resistance to a broad spectrum of chemical pesticides. It is well-established that knowledge of diversity and evolutionary origins facilitate the development of pest management strategies. In the present study, we report the morphology, diversity and phylogeny analysis of S. exigua from North Sulawesi, Indonesia. The specimen from Rurukan have a body size and other segments that are longer than in Langowan and Modoinding. Dendrogram analysis shows that the similarity distance based on morphology ranges from 1-25%, which forms four clusters, where the specimen from Rurukan is separated from the rest of the specimens. The phylogeny of S. exigua from North Sulawesi, Indonesia, based on CO1 (Cytochrome c oxidase subunit 1) gene fragment, which is juxtaposed with CO1 data of the allied species from many geographical locations. A total of twenty-five isolates representing Indonesia, Japan, Germany, Thailand, India, UK, USA, Spain and Australia were compared. Nineteen sequences of S. exigua retrieved from GenBank were selected as references based on previous published phylogenic trees. The twenty-four isolates were scattered in two distinct clades indicating S. exigua is polyphyletic, but S. exigua from North Sulawesi, Indonesia is monophyletic.
... The eruciform larvae of Panorpidae have been described in detail in five genera: Panorpa Linnaeus, 1758(Miyak e 1912Steiner 1930;Byers & Thornhill 1983); Neopanorpa van der Weele, 1909 (Yie 1951 (Ma et al., 2014); and Cerapanorpa Gao, Ma & Hua, 2016(Jiang & Hua 2015b. The larval morphology is diverse among genera (Bierbrodt 1942;Yie 1951;Boese 1973;Cai & Hua 2009a;Jiang et al., 2019;, and varies among species of Panorpa (Mampe & Neunzig 1965;Cai & Hua 2009b;Jiang & Hua 2013. The larvae of Panorpa vary in head morphology and cephalic chaetotaxy among Panorpa qinglingensis Chou & Ran in Chou et al., 1981 (the Panorpa davidi group), P. liui Hua, 1997 (the Panorpa amurensis group) and Panorpa macrostyla Hua, 1998 (the Panorpa kongosana group) (Cai & Hua 2009b;Jiang & Hua 2013. ...
... 2200e2500 m), Kunming, Yunnan Province, southwestern China from late August to early September in 2017. Adults were reared in pairs in plastic jars covered with a piece of gauze under semi-natural conditions (Liu & Hua 2017;Jiang et al., 2019). The jars were filled with humid soil of approximately 5 cm in depth for females to deposit eggs inside. ...
... The dorsal protuberances on the larval abdominal segments in Panorpidae are furnished with annulated processes, a noticeable character in Mecoptera (Brauer 1863;Miyak e 1912;Grell 1938;Byers 1987;Cai & Hua 2009a;Chen & Hua 2011). These processes vary in length and morphology among genera related to habitat divergence in the Panorpidae (Jiang & Hua 2015b;Jiang et al., 2019). The annulated processes are greatly shortened in the euedaphic larvae of Neopanorpa, but are extremely elongate in the epedaphic larvae of Dicerapanorpa and medium-sized in Cerapanorpa (Ma et al., 2014;Jiang & Hua 2015b). ...
Article
The first-instar larvae of the scorpionflies Panorpa kunmingensis Fu and Hua, 2009 and Panorpa changbaishana Hua, 1998 were compared using scanning electron microscopy. The larvae of both species are eruciform and similar in gross morphology, each bearing three pairs of thoracic legs and eight pairs of abdominal prolegs. The head is more flattened and widened in P. changbaishana than in P. kunmingensis. Cephalic setae are mostly setiform in P. kunmingensis, but are apically expanded in some setae of P. changbaishana. Annulated processes are more prominent in P. changbaishana than in P. kunmingensis. The potential value of larval characters in phylogenetic analyses is briefly discussed in the Panorpidae.