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Nemoptera sinuata (Neuroptera: Nemopteridae). 1 – visiting a flower of Achillea coarctata (Asteraceae) (June 2006, Struma Valley, Bulgaria); 2 – feeding, accumulation of pollen on the frontal part of head; arrow. Inflorescence of Achillea coarctata composed of numerous single flowers.
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The mouthparts of the spoon-winged lacewing Nemoptera sinuata are adapted for the uptake of pollen and nectar. Form and function of the mouthparts are described, and the technique of food uptake is discussed in context with flower-visiting behaviour and floral architecture of the preferred flowers. The maxillae are the main organs for food uptake....
Citations
... It is interesting that the diversification within the SA group is confined to the Afrotropical region, whereas the diversification of the LA group involves multiple dispersals out of Africa. In terms of the biological traits, Nemopterinae are known to be thermophilic, pollenophagous in adults, and sandysoil dwelling near the roots of desert plants as larvae (Mansell 1973, 1996, Monserrat and Martínez 1995, Krenn et al. 2008, Miller and Stange 2012. As assumed by Sole et al. (2013), the high species diversity of the African Nemopterinae might have co-evolved with Ruschioideae (Caryophyllales, Aizoaceae) that have greatest species diversity and endemism from southern Africa, and the rareness of these plants in the other continents might hamper the diversification of spoon-winged lacewings beyond the Afrotropical region. ...
The spoon-winged lacewings (Neuroptera: Nemopteridae: Nemopterinae) are a group of charismatic insects with morphological and biological specializations. Among the known 105 species of Nemopterinae worldwide, only one species, namely Nemopistha sinica Yang, 1986, is recorded from East Asia. However, the morphology, taxonomic status, and evolutionary history of this rare species are poorly known. Here, we present a systematic revision of the Chinese Nemopterinae and establish a new genus, Sinonemoptera, that comprises Sinonemoptera sinica (Yang, 1986) comb. nov. from western Yunnan and a new species, Sinonemoptera tibetana sp. nov., from southeastern Tibet. Based on the phylogeny of Nemopterinae combining morphological and molecular evidence, Nemopterinae are divided into two major clades by the length of the adult abdomen, and Sinonemoptera gen. nov. together with some Afrotropical genera constitute a monophyletic lineage characterized by a long abdomen. Our results suggest a Late Cretaceous African origin and three Tertiary transcontinental dispersals in shaping the global distribution of Nemopterinae. Our ecological niche modelling demonstrates the specific requirement for warm and dry habitats in nemopterines and highlights the urgent need for protection of the savannah-like habitat along the Nujiang valley for the Chinese Nemopterinae.
... In addition, even though sunn hemp flowers have nectaries (Meagher et al. 2019) that could attract adult C. externa for nectar drinking to obtain carbohydrates, a large amount of its pollen was consumed by females and males. To our knowledge, the flower-visiting behavior and functional mode of the mouthparts of crepuscular and nocturnal adult C. externa when consuming pollen through flower visitation have not been published thus far; however, these aspects were described for adults of the spoon-winged lacewing, Nemoptera sinuata Olivier (Neuroptera: Nemopteridae), which is a diurnal flower visitor and an obligate pollen feeder (Krenn et al. 2008). Studies on pollen manipulation by adult C. externa are limited by its crepuscular and nocturnal behavior. ...
Pollen is a food source for adult Chrysoperla externa (Hagen), whose larvae are biocontrol agents against pests. However, adults may face challenges in foraging for pollen due to differences in pollen accessibility and variability in pollen morphology and chemistry. In the laboratory, we investigated the ability of adult C. externa to consume pollen from flowers of Cajanus cajan, Canavalia ensiformis, Crotalaria juncea, Flemingia macrophylla, Avena strigosa, Pennisetum glaucum, Sorghum bicolor, and Zea mays, and we explored whether adults chose any of these pollens based on their quantitative and qualitative features. Cajanus cajan and F. macrophylla pollen were the only ones not consumed by adults when confined to flowers. Pollen removed from the preanthesis buds was offered simultaneously for 24 and 48 h. In both periods, adults consumed more of the medium-sized P. glaucum (with the second largest exine thickness) and large-sized Z. mays (with the thinnest exine) pollen, even though they had significantly less crude protein than Fabaceae pollen, whose sizes varied from medium (C. juncea, with the thickest exine) to large (C. ensiformis, whose exine thickness was equal to that of P. glaucum). Overall, adults consumed more Poaceae pollen than Fabaceae pollen, but the palynological features and the protein contents did not affect this choice. Our results highlighted that C. juncea, P. glaucum, S. bicolor and Z. mays are good pollen sources for adult C. externa and should be considered promising candidates in the selection of insectary plants to deploy in biocontrol programs aimed at the conservation of this lacewing.
... The recent finding of Myanmar amber nemopterids represents another notable case for understanding the early evolution of pollinating behaviour in lacewings. Extant nemopterids are a group of flower-visiting insects, possessing a long rostrum modified from the elongation of mandibulate mouthparts in adaption to pollen-feeding habits (Tjeder, 1967;Krenn et al., 2008). The plant hosts of extant nemopterids comprise a broad range of angiosperms as well as a few gymnosperms (Tjeder, 1967;Monserrat, 1983). ...
The palaeodiversity of Neuropterida was extraordinarily rich during the Cretaceous in northern Myanmar. Owing to the sharply increased studies on the Myanmar amber Neuropterida, a total of 108 genera and 135 species in 23 families of three orders have been recorded. Here we provide a review of the present knowledge on the systematics, phylogeny, and palaeoecology of Neuropterida from the mid-Cretaceous (lowermost Cenomanian) of Myanmar. Cretadilarinae Makarkin, 2017 is here moved to Sisyridae, being closely related to the extinct sisyrid subfamily Paradoxosisyrinae Makarkin, 2016. A catalogue and a faunal analysis on Neuropterida recorded from world amber deposits are provided. The Myanmar amber Neuropterida highlights the Cretaceous diversity of this order and provides significant fossil evidence to infer the evolutionary history of this archaic and charismatic insect group.
... In Nemopteridae, showing the greatest number of adaptations to palynivory, the elongate maxillae participate in pollen collection while the mandibles have lost their chewing function and are used for removing the pollen stuck to the other mouthparts. However, even spoonwings have not lost any elements of the standard chewing apparatus (Krenn et al., 2008). ...
The paper overviews the fossil record of insects with long mouthparts and rostra adapted to feeding on floral nectar and pollination drops of extinct gymnosperms. The presence of suctorial mouthparts is demonstrated for the first time for the Permian mecopterans Permochoristidae and Permotanyderidae. The long-proboscid scorpionflies Mesopsychidae are recorded for the first time from the Upper Jurassic of Kazakhstan. A new finding of a detached head of a long-proboscid nectar-feeding brachyceran fly is reported from the Lower Cretaceous of Transbaikalia. Three major radiations of long-proboscid nectar feeders are identified: the Paleozoic, the Mesozoic, and the Cenozoic one; they were related to the Paleozoic seed ferns, the Bennettitales and other Mesozoic entomophilous gymnosperms, and the flowering plants, respectively. The earliest long-proboscid nectar feeders, found in the Lower Permian deposits of the Cis-Urals, belong to Protomeropidae (stem-Amphiesmenoptera). The few other Paleozoic insects specialized to nectarivory probably also included some long-proboscid Permochoristidae. The diversity of long-proboscid nectar feeders shows a dramatic increase since the Middle Jurassic. About 70 Mesozoic species with preserved long mouthparts and rostra are known to date; they belong to 12 families and 3 orders (Mecoptera, Neuroptera, and Diptera) and can be clustered into three morphogroups. With the beginning of the Cenozoic the long-proboscid Mecoptera and Neuroptera were supplanted by Hymenoptera and Lepidoptera in the nectar-feeding niche, while Diptera on the whole retained this specialization. Considerable abundance of long-proboscid nectar feeders before the appearance of flowers with hidden nectar indicates that complex pollination systems first evolved in gymnosperms. Therefore, insect pollination cannot be considered the key novelty in flowering plants crucial for their evolutionary success.
... In Myrmeleontiformia, Hemerobiidae (Tjeder 1961) and Mantispidae (Ferris 1940;Lucchese 1956;Poivre 1978Poivre , 1981, the area between the molar process and the incisivus is prolonged and forms a distinct cutting edge (Fig. 11.9g-j). The mandibles are usually asymmetric in Neuroptera, with exceptions in phytophagous forms such as the ithonid Ithone fusca (Shepard 1967), the chrysopid Pseudomallada prasinus (Stelzl 1992) and Nemopteridae (Acker 1958;Tjeder 1967;Krenn et al. 2008). ...
... Pollen is a major food source for the adults of Nemopteridae (Fig. 11.7) and Crocidae, as well as some species of Chrysopidae, Berothidae, Sisyridae and Myrmeleontidae (Pupedis 1987;Devetak and Klokočovnik 2016). The process of pollen-feeding was studied in detail in Nemoptera (Krenn et al. 2008): in these flower-visiting lacewings the brush-shaped setaceous laciniae function as the main pollen-collecting organ. The maxillary structures are elongated, being folded beneath the head in resting position and extended forward during feeding. ...
... In contrast to the labium, the maxillae are well developed and the cardo, stipes, lacinia and galea are elongated and folded sideways at the cardostipital joint (Fig. 11.10c), so that they can be extended forwards with the respective muscular contraction. A similar formation, though with a downward folding at the cardo-stipital joint, is present in adult Nemopteridae (Krenn et al. 2008), where the movement of the maxillae serves to collect nectar and pollen from flowers. Accordingly, a palyno-glycophagous feeding as in Nemopteridae seems to be the most plausible hypothesis. ...
The Neuroptera are highly heterogeneous endopterygote insects. While their relatives Megaloptera and Raphidioptera have biting mouthparts also in their larval stage, the larvae of Neuroptera are characterized by conspicuous sucking jaws that are used to imbibe fluids, mostly the haemolymph of prey. They comprise a mandibular and a maxillary part and can be curved or straight, long or short. In the pupal stages, a transformation from the larval sucking to adult biting and chewing mouthparts takes place. The development during metamorphosis indicates that the larval maxillary stylet contains the Anlagen of different parts of the adult maxilla and that the larval mandibular stylet is a lateral outgrowth of the mandible. The mouthparts of extant adult Neuroptera are of the biting and chewing functional type, whereas from the Mesozoic era forms with siphonate mouthparts are also known. Various food sources are used in larvae and in particular in adult Neuroptera. Morphological adaptations of the mouthparts of adult Neuroptera to the feeding on honeydew, pollen and arthropods are described in several examples. New hypotheses on the diet of adult Nevrorthidae and Dilaridae are presented.
... Adult nemopterids are obligate nectar and pollen feeders with specially adapted elongate mouthparts (Popov 1963;Picker 1987;Monserrat & Martinez 1995;Mansell 1996;Monserrat 1996;Krenn et al. 2005, Krenn et al. 2008. These modifications are derived traits that have evolved from predatory biting and chewing mouthparts to the present form and function (Krenn et al. 2008). ...
... Adult nemopterids are obligate nectar and pollen feeders with specially adapted elongate mouthparts (Popov 1963;Picker 1987;Monserrat & Martinez 1995;Mansell 1996;Monserrat 1996;Krenn et al. 2005, Krenn et al. 2008. These modifications are derived traits that have evolved from predatory biting and chewing mouthparts to the present form and function (Krenn et al. 2008). Unlike the adults, larvae of Nemopteridae are all specialised predators, with elongate mandibles and maxillary laciniae adapted for piercing and ingesting the internal tissues of small arthropods (Popov 1963(Popov , 1973. ...
The southern African genera Nemopterella Banks, 1910 and Nemia Navás, 1915 (Neuroptera: Nemopteridae: Nemopterinae) are revised. Nemopterella is split into three genera: Nemopterella sensu stricto with type species Nemopteryx africana Leach, 1815 (= Nemopterella africana), Afroptera gen. nov., with type species Nemopterella munroi Tjeder, 1967, and the monotypic genus Siccanda gen. nov., with type species Nemopterella arenaria Tjeder, 1967. Eight new species are described in Afroptera gen. nov.: A. acuta Abdalla & Mansell sp. nov., A. alba Mansell & Abdalla sp. nov., A. brinkmani Abdalla & Mansell sp. nov., A. balli Abdalla & Mansell sp. nov., A. cylindrata Abdalla & Mansell sp. nov., A. folia Abdalla & Mansell sp. nov., A. koranna Mansell & Abdalla sp. nov., A. maraisi Abdalla & Mansell sp. nov., as well as two new species in the genus Nemopterella: N. kabas Mansell & Abdalla sp. nov., and N.cedrus Mansell & Abdalla sp. nov.
... In fact, the modifications of adult mouthparts between extant Crocinae and Nemopterinae are also generally similar (Tjeder, 1967), indicating possible analogous feeding behaviour. Nemopterinae are better documented for their flower-visiting behaviour than Crocinae, which lack direct evidence of flower-visiting (Monserrat, 1996;Krenn et al., 2008). However, the similar mouthpart structures and detection of pollen in the gut content in both Crocinae and Nemopterinae suggest that crocine adults are also flower visitors. ...
... (1) Araneae (N = 17, Table 1), which scarcely moved and were generally predators, presumably waiting for prey; (2) Hemiptera (N = 93), which also stayed on the peduncles and rarely moved, suggesting that they were merely herbivores of Lindera flowers; and (3) Neuroptera (N = 1), which fed on pollen and were potentially pollinators of other plant species (Krenn et al. 2008) (however, we found only one individual moving on Lindera flowers (Table 1)). These three arthropod orders may be foragers or herbivores of Lindera species, rather than potential pollinators. ...
Two Lindera species (Lindera praecox and Lindera triloba) are distributed sympatrically along the Pacific Ocean side of the main island of Japan. They are similar, but not identical, in habitat preference and flowering season. Slight differences in these traits have been proposed, but a comparative study on the reproductive ecology of these two species has not yet been conducted. To reveal the interspecies differences between these two Lindera spp. and characterize the life historical characteristics related to their reproduction, we assessed differences in floral visitors and reproductive ecology (fruit set, seed and pulp weights, and pollen limitation) between species and sexes over a period of 3 years. A total of 39 species (458 individuals) of floral visitors, approximately 70% of which comprised three Coleoptera species, were collected from plants of both species and sexes. The number of insect species that visited the late-flowering species, L. triloba, was significantly higher than the number that visited the early-flowering species, L. praecox. However, the numbers of visitors were not significantly different between species. The fruit set was higher in L. triloba than in L. praecox, but L. praecox pulps were heavier than those of L. triloba. Our study revealed interspecies differences in floral visitors and reproductive success between two species and indicate the possible differences of reproductive strategy of these two Lindera spp.
... These palps are densely covered with long setae, and there is an ovoid sensory area in the terminal palpomeres (Fig. 2h, i and Supplementary Fig. 2b). Such features suggest that these palps Tables 1 and 2 for details were probably used to probe for nectar or pollen 33,34 . Furthermore, these palps are also present in three well-preserved kalligrammatid specimens from the Jurassic and Cretaceous of China ( Supplementary Fig. 10) as well as in a specimen from Central Asia 20 , suggesting that both maxillary and labial palps were present in most or even all kalligrammatids. ...
Niche diversity of pollinating insects plays a vital role in maintaining extant terrestrial ecosystems. A key dimension of pollination niches refers to the insect proboscis length that commonly matches the floral tube length. Here we describe new kalligrammatid lacewings (an iconic Mesozoic pollinating insect lineage) from late Cretaceous Burmese amber and Mesozoic sediments in China. Kalligrammatids display complex configurations of elongate mouthpart elements consisting of well-developed maxillae, labium and their palps. The mouthpart lengths vary among species, from 0.6 to 18.0 mm, suggesting corresponding variability in the floral tube lengths of Mesozoic plants. With the diversification of pollinating habits, the kalligrammatids presented highly divergent traits related to chemical communication and defence mechanisms. Together with other Mesozoic long-proboscid insects, these fossils not only reveal the high niche diversity of Mesozoic pollinating insects but also highlight the diversity of Mesozoic pollinator-dependent plants prior to the rise of angiosperms.
... The externally discernible sutures delimiting the hypostomal bridge laterally (Zimmermann et al., 2009) are represented internally as two lateral hypostomal ridges. Hypostomal bridges are not an unusual formation for Neuroptera, smaller hypostomal bridges are also present in some species of the families Mantispidae (Ferris, 1940;Shepard, 1967: postgenal bridge), Nemopteridae (Acker, 1958;Krenn et al., 2008: Fig. 7) and Chrysopidae (Morse, 1931: described as "gula"; see Randolf et al., 2013). Anteriorly protruding processes, such as those of the hypostomal bridge in Coniopteryx that serve as an articulation site for the cardines, are also present in Micromus posticus, Sympherobius barberi and Plega dactylota, but are formed by the postgenae (Shepard, 1967). ...
External and internal head structures of adult Coniopteryx pygmaea Enderlein, 1906, one of the smallest known lacewings, are described in detail for the first time. Possible effects of miniaturization and two hypotheses on the phylogenetic position of Coniopterygidae are evaluated and compared with data from literature. Several convergent modifications in C. pygmaea and other miniaturized insect species are outlined, e.g., a relative increase in the size of the brain, simplification of the tracheal system with respect to the number of tracheae, and reduction of the number of ommatidia and diameter of the facets. Further, the ocular ridge is bell-shaped and countersunk into the head capsule. The cuticle is weakly sclerotized and equipped with wax glands which are unique in Neuroptera. The total number of muscles is not affected by miniaturization. The phylogenetic analysis yields Coniopterygidae as sistergroup to the dilarid clade based on one larval character, the shape of the stylets. The enforced basal position of Coniopterygidae is supported by one disputable synapomorphy of the remaining Neuroptera, the presence of paraglossae in adults.
