Phylogenomics and biogeography of sawflies and woodwasps (Hymenoptera, Symphyta).

Phylogenomic approaches have recently helped elucidate various insect relationships, but large-scale comprehensive analyses on relationships within sawflies and woodwasps are still lacking. Here, we infer the relationships and long-term biogeographic history of these hymenopteran groups using a large dataset of 354 UCE loci collected from 385 species that represent all major lineages. Early Hymenoptera started diversifying during the Early Triassic âˆ¼249 Ma and spread all over the ancient supercontinent Pangaea. We recovered Xyeloidea as a monophyletic sister group to other Hymenoptera and Pamphilioidea as sister to Unicalcarida. Within the diverse family Tenthredinidae, our taxonomically and geographically expanded taxon sampling highlights the non-monophyly of several traditionally defined subfamilies. In addition, the recent removal of Athalia and related genera from the Tenthredinidae into the separate family Athaliidae is supported. The deep historical biogeography of the group is characterised by independent dispersals and re-colonisations between the northern (Laurasia) and southern (Gondwana) palaeocontinents. The breakup of these landmasses led to ancient vicariance in several Gondwanan lineages, while interchange across the Northern Hemisphere has continued until the Recent. The little-studied African sawfly fauna is likewise a diverse mixture of groups with varying routes of colonization. Our results reveal interesting parallels in the evolution and biogeography of early hymenopterans and other ancient insect groups.

Mitochondrial genome rearrangements and phylogenomics of the Hymenoptera (Insecta) using an expanded taxon sample.

The order Hymenoptera is one of the most species-rich insect orders, with more than 150,000 described extant species. Many hymenopteran insects have very different mitochondrial genome (mitogenome) organizations compared to the putative ancestral organization of insects. In this study, we sequenced 18 mitogenomes of representatives in the order Hymenoptera to increase taxonomic sampling. A total of 475 species were used in phylogenetic analyses, including 18 new mitogenomes and 457 existing mitogenomes. Using a site-heterogeneous model, Bayesian's inference from amino acid data yielded more resolved relationships among Hymenoptera than maximum-likelihood analysis and coalescent-based species analyses. The monophyly of Symphyta was not supported. The Xyeloidea was the earliest branching clade in the Hymenoptera. The Orussoidea was closely related to Apocrita. Within Apocrita, the Parasitoida was non-monophyletic. The monophyly of most Parasitoida superfamilies received strong support. The Proctotrupomorpha clade was supported in Bayesian's analysis. The Apoidea was monophyletic when excluding Ampulex compressa from consideration. The superfamilies Vespoidea and Chrysidoidea were found to be non-monophyletic. Comparisons of mitochondrial gene order revealed a higher frequency of gene rearrangement among lineages with a parasitoid lifestyle, particularly prominent in Chalcidoidea. The degree of gene rearrangement ranked second in specific taxa of Cynipoidea and Ichneumonoidea.

Cimbicidae (Hymenoptera, 'Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands.

We erect the Cenocimbicinae, a new subfamily of Cimbicidae (Hymenoptera, Symphyta), restricted to the Selandian Menat Formation of France, the oldest occurrence of the family, and the Ypresian Okanagan Highlands of far-western North America. We describe new taxa from the Okanagan Highlands: Allenbycimbex morrisae gen. et sp. nov. and Leptostigma n. gen. with seven new species: L. alaemacula n. sp., L. brevilatum n. sp., L. fasciatum n. sp., L. longiclava n. sp., L. longipallidum n. sp., L. longitenebricum n. sp., and L. proxivena n. sp. We revise the Cimbicidae from the Ypresian Green River Formation and the Priabonian Florissant Formation, both in Colorado, USA. The oldest fossil of a modern cimbicid subfamily appears with a single pachylostictine specimen in the Green River Formation, and all Cimbicidae are in modern subfamilies after the Ypresian (we did not examine one larva known from Priabonian Baltic amber). Pseudocimbex clavatus Rohwer 1908 from the Florissant Formation is not a cimbicid; we treat it as Tenthredinoidea incertae sedis. We transfer Cimbex vetusculus Cockerell to Floricimbex n. gen.

Trait-mediated responses of caterpillar communities to spongy moth outbreaks and subsequent tebufenozide treatments.

Outbreaks of the spongy moth Lymantria dispar can have devastating impacts on forest resources and ecosystems. Lepidoptera-specific insecticides, such as Bacillus thuringiensis var. kurstaki (BTK) and tebufenozide, are often deployed to prevent heavy defoliation of the forest canopy. While it has been suggested that using BTK poses less risk to non-target Lepidoptera than leaving an outbreak untreated, in situ testing of this assumption has been impeded by methodological challenges. The trade-offs between insecticide use and outbreaks have yet to be addressed for tebufenozide, which is believed to have stronger side effects than BTK. We investigated the short-term trade-offs between tebufenozide treatments and no-action strategies for the non-target herbivore community in forest canopies. Over 3 years, Lepidoptera and Symphyta larvae were sampled by canopy fogging in 48 oak stands in southeast Germany during and after a spongy moth outbreak. Half of the sites were treated with tebufenozide and changes in canopy cover were monitored. We contrasted the impacts of tebufenozide and defoliator outbreaks on the abundance, diversity, and functional structure of chewing herbivore communities. Tebufenozide treatments strongly reduced Lepidoptera up to 6 weeks after spraying. Populations gradually converged back to control levels after 2 years. Shelter-building species dominated caterpillar assemblages in treated plots in the post-spray weeks, while flight-dimorphic species were slow to recover and remained underrepresented in treated stands 2 years post-treatment. Spongy moth outbreaks had minor effects on leaf chewer communities. Summer Lepidoptera decreased only when severe defoliation occurred, whereas Symphyta declined 1 year after defoliation. Polyphagous species with only partial host plant overlap with the spongy moth were absent from heavily defoliated sites, suggesting greater sensitivity of generalists to defoliation-induced plant responses. These results demonstrate that both tebufenozide treatments and spongy moth outbreaks alter canopy herbivore communities. Tebufenozide had a stronger and longer lasting impact, but it was restricted to Lepidoptera, whereas the outbreak affected both Lepidoptera and Symphyta. These results are tied to the fact that only half of the outbreak sites experienced severe defoliation. This highlights the limited accuracy of current defoliation forecast methods, which are used as the basis for the decision to spray insecticides.

Cypress terpenes in sawfly larva of Susana cupressi (Hymenoptera: Symphyta: Tenthredinoidea).

Several sawfly species (Hymenoptera: Symphyta) possess larval stages with oesophageal diverticula in which plant compounds are sequestered and used for defence against predators. These organs are present in the larvae of Susana (Tenthredinidae) but remain poorly studied. Here, the aim was to analyse the diverticula extract of Susana cupressi by gas chromatography-mass spectrometry to better understand the ecology of this species. The foliage of the hostplant (Cupressus sempervirens), as well as the larval foregut, midgut, and haemolymph were also analysed. Complementary data were gathered by morphological observations, bioassays using ants, and genetic analyses to identify the studied Susana species. Altogether, 48 terpenes were identified, 30 being sesquiterpenes. The terpenes were generally detected in the foliage, but also in the diverticula, foregut, and midgut, whereas none of them in the haemolymph. The main compounds were alpha-cedrene, alpha-fenchene, alpha-pinene, alpha-terpinyl acetate, beta-myrcene, beta-pinene, cedrol, delta 3-carene, epi-bicyclosesquiphellandrene, germacrene D, limonene, sabinene, and terpinolene. The chemical profiles of these 13 compounds were significantly correlated between foliage-diverticula, diverticula-foregut and foregut-midgut, but not correlated for the three remaining possible comparisons. Alpha-pinene decreased and germacrene D increased from the foliage to the diverticula, which may reflect a specific sequestration of the latter terpene and its known deleterious effects on insects. We conclude that larvae of S. cupressi, similarly to those of diprionids, are well defended against predatory attacks by sequestering and regurgitating hostplant terpenes, including germacrene D.

The complete mitochondrial genome and novel gene arrangement in Nesodiprion zhejiangensis Zhou & Xiao (Hymenoptera: Diprionidae).

The complete mitochondrial genome (mitogenome) of the sawfly, Nesodiprion zhejiangensis Zhou & Xiao, was sequenced, assembled, and deposited in GenBank (Accession Number: OM501121). The 15,660 bp N. zhejiangensis mitogenome encodes for 2 ribosomal RNAs (rrnL and rrnS), 22 transfer RNAs (tRNAs), 13 protein-coding genes (PCGs), and an AT-rich region of 450 bp in length. The nucleotide composition is biased toward adenine and thymine (A + T = 81.8%). Each PCG is initiated by an ATN codon, except for cox2, which starts with a TTG. Of 13 PCGs, 9 have a TAA termination codon, while the remainder terminate with a TAG or a single T. All tRNAs have the classic cloverleaf structure, except for the dihydrouridine (DHU) arm of tRNAval, which forms a simple loop. There are 49 helices belonging to 6 domains in rrnL and 30 helices belonging to 4 domains in rrnS. In comparison to the ancestral architecture, N. zhejiangensis has the most rearranged mitogenome in Symphyta, in which rearrangement events of local inversion and transposition are identified in three gene clusters. Specifically, the main hotspot of gene rearrangement occurred between rrnS and trnY, and rearranged from rrnS-(AT-rich region)-I-Q-M-nd2-W-C-Y to rrnS-Q-W-C-nd2-I-M-(AT-rich region)-Y, involving a local inversion event of a large gene cluster and transposition events of some tRNAs. Transposition of trnA and trnR (rearranged from A-R to R-A) was observed at the nd3-nd5 gene junction while shuffling of trnP and trnT (rearranged from T-P to P-T) occurred at the nd4l-nd6 gene junction. While illegitimate inter-mtDNA recombination might explain the opposite orientations of transcription between rrnS and trnY, transposition events of tRNA in some gene blocks can be accounted for by the tandem duplication/random loss (TDRL) model. Our phylogenetic analysis suggests that N. zhejiangensis is closely related to congeneric species N. biremis and N. japonicus, which together form a sister lineage with the European pine sawfly, Neodiprion sertifer.

The complete mitochondrial genome of the woodwasp Euxiphydria potanini (Hymenoptera, Xiphydrioidea) and phylogenetic implications for symphytans.

The long-necked woodwasp superfamily Xiphydrioidea belongs to the suborder Symphyta (Hymenoptera). Here we newly characterize the complete mitochondrial genome of the South Korean Euxiphydria potanini (Xiphydriidae) using next-generation sequencing: 16,500 bp long with 84.27% A + T content and 37 typical mitochondrial genes including those encoding 13 PCGs, 2 rRNAs, 22 tRNAs, and one A + T rich region. We compare the patterns of symphytan mitochondrial gene arrangement with those of an ancestral insect form and found some synapomorphic rearrangements in phylogenetic context. We use a variety of nucleotide and amino acid sequence alignments (thirteen mtPCGs and/or eight nDNAs) alongside step-by-step exclusions of long-branched taxa to elucidate the phylogenetic position of Xiphydrioidea and phylogenetic relationships among the seven symphytan superfamilies, except for Anaxyeloidea of which no mtgenome was available. The monophyly of symphytan superfamilies (with weak support for Pamphilioidea), sister-group relationship of Xiphydrioidea and Cephoidea, and Symphyta being paraphyletic to Apocrita, etc. are consistently supported by maximum likelihood and Bayesian inference trees. We also discuss the problematic phylogenetic positions of Orussoidea and Siricoidea and propose a hypothetical scenario of morphological character transition during hymenopteran evolution based on morphological key characteristics, such as the cenchrus and the wasp-waist.

Two New Fossil Sawflies of Pamphiliidae (Hymenoptera: Symphyta) from the Mesozoic of Northeastern China.

Two new species of Pamphiliidae, Scabolyda latusa sp. nov. and Scabolyda tenuis sp. nov. are described and illustrated from the late Middle Jurassic Jiulongshan Formation and the Lower Cretaceous Yixian Formation of northeastern China, respectively. A new specimen of Scabolyda orientalis Wang, Rasnitsyn, Shih and Ren, 2014 with distinct male genitalia is documented for the first time. Based on the specimens with new and distinct structures of legs, antennae, and genitalia, the morphological characters of Scabolyda are supplemented: antenna with ca. 13-14 flagellomeres; fore leg with tibia without pre-apical spur; hind leg nearly 0.6 times as long as the body, hind tarsal claw without setae and its inner tooth not developed. In addition, the tarsal claw characteristics found in the new species may suggest Scabolyda has a closer relationship with Cephalciinae, rather than with Pamphiliinae.

New Fossil Xyelidae (Hymenoptera: Symphyta) from the Mesozoic of Northeastern China.

One new genus and species, Leptoxyela eximia gen. et sp. nov., and one new species, Scleroxyela cephalota sp. nov., are described and illustrated based on two well-preserved compression fossils from the Lower Cretaceous Yixian Formation and the Middle Jurassic Jiulongshan Formation of China, respectively. Leptoxyela eximia gen. et sp. nov. is placed in the tribe Angaridyelini, 1966 and Scleroxyela cephalota sp. nov. in the tribe Xyeleciini Benson, 1945; while both tribes are in the subfamily Macroxyelinae Ashmead, 1898 of Xyelidae. A key to the genera of Angaridyelini is provided. In addition, we investigated various angles between Rs+M and 1-Rs for known fossil species of Macroxyelinae, and we found the angle and the length of 1-Rs are correlated; however, we could not see any correlation between the angles and the fossil ages even within a tribe. Furthermore, based on Sc2 connected to R before Rs, the angle between Rs+M and 1-Rs, and the length of the first flagellomere, we believe that the tribe Ceroxyelini may have only one genus of Ceroxyela, and suggest that Isoxyela and Sinoxyela should be transferred to Gigantoxyelini.

The early Eocene Eourocerus anguliterreus gen. et sp. nov (Hymenoptera, Siricidae) from Republic, Washington.

A new fossil horntail wood wasp (Hymenoptera, Symphyta, Siricidae), Eourocerus anguliterreus gen. et sp. nov. from an early Eocene Okanagan Highlands locality at Republic, Washington, USA is described. Its forewing is most like that of the extant and fossil genus Urocerus Geoffroy, 1785. It is the third fossil siricid described from North America. We treat Xeris dorbnikensis Manukyan Smirnova, 2021 and Xeris sp. of Manukyan Smirnova (2021) as Urocerus dorbnikensis (Manukyan Smirnova, 2021), comb. nov. and Urocerus sp., both tentatively belonging to the genus, Urocerus klebsi Brues, 1926 as Xeris klebsi (Brues, 1926) comb. nov., and Afrotremex or Eriotremex sp. of Wedmann (1998) as a species of Eriotremex Benson, 1943. A key to extant and extinct genera of Siricinae by forewing characters is given.

The First Fossil Representatives of the Sawfly Genera Emphytus and Empria from the upper Miocene of France (Hymenoptera, Tenthredinidae).

Emphytusmiocenicus sp. nov., first fossil representative of this genus, is described from the upper Miocene of the diatom paleolake of Montagne d'Andance (Ardèche, France). Its placement is ensured by an in-depth comparison with all the extant and fossil genera of the subfamily Allantinae. The representatives of Emphytus are distributed in the Palearctic, Nearctic, and Oriental regions. Empria sammuti sp. nov., second representative of the latter genus, is described from the latest Miocene of the diatom paleolake of Sainte-Reine (Cantal, France). The placement of this new species is based on a detailed comparison with the extant genera of the tribe Empriini. The larvae of the extant Emphytus and Empria spp. are known to be phytophagous on angiosperm leaves of several families, all present as fossils in the taphocenoses of la Montagne d'Andance and Sainte-Reine. Emphytus miocenicus sp. nov. represents the oldest record of this genus and of its crown group, corroborating the estimate of a middle Eocene-middle Oligocene age for its stem group. Throughout our study, it appears that the first described fossil of the genus Empria, E. oligocaenica, from the Oligocene of Germany, needs to be revised and redescribed. It should preferably be treated until the revision as incertae sedis in Allantinae sensu lato.

Sex Pheromone of the Introduced Pine Sawfly, Diprion similis, Revisited to Define a Useful Monitoring Lure: Deviating Chiral Composition and Behavioural Responses Compared to Earlier Reports.

Extracts of Diprion similis females contained about 15 ng of the sex pheromone precursor 3,7-dimethylpentadecan-2-ol per female. After derivatisation with (S)-2-acetoxypropanoyl chloride, we found that the major stereoisomer in the extract was (2S,3R,7R)-3,7-dimethylpentadecan-2-ol. Small amounts of other stereoisomers of 3,7-dimethylpentadecan-2-ol were also identified in the extract, namely 1% of (2R,3S,7S), 0.3% (2R,3R,7R) and 0.4% of (2R,3R,7S). An unknown fifth substance showed a very similar spectrum to 3,7-dimethylpentadecan-2-ol, both in SIM and full scan mode. None of the earlier suggested behavioural synergistic isomers ((2S,3S,7S), (2S,3S,7R) and (2S,3R,7S)) were detected in the extracts. In field tests in Ontario, Canada, the earlier identified main pheromone component, viz. the propanoate of (2S,3R,7R)-3,7-dimethylpentadecan-2-ol, was tested alone and in combination with other stereoisomers, earlier reported to be synergistic. No synergistic effects were detected and the threo four-isomer blend was as attractive as the pure main compound. Thus, one of the few examples of a diprionid sawfly using more than one substance in its sex pheromone could not be confirmed. The results also suggest that monitoring programs can use the more easily synthesized threo-blend without losing efficiency. Furthermore, the study suggests that other diprionid pheromones may benefit from a reinvestigation, to clarify possible synergistic effects of stereoisomers.

Evidence for UV-green dichromacy in the basal hymenopteran Sirex noctilio (Siricidae).

A precondition for colour vision is the presence of at least two spectral types of photoreceptors in the eye. The order Hymenoptera is traditionally divided into the Apocrita (ants, bees, wasps) and the Symphyta (sawflies, woodwasps, horntails). Most apocritan species possess three different photoreceptor types. In contrast, physiological studies in the Symphyta have reported one to four photoreceptor types. To better understand the evolution of photoreceptor diversity in the Hymenoptera, we studied the Symphyta Sirex noctilio, which belongs to the superfamily Siricoidea, a closely related group of the Apocrita suborder. Our aim was to (i) identify the photoreceptor types of the compound eye by electroretinography (ERG), (ii) characterise the visual opsin genes of S. noctilio by genomic comparisons and phylogenetic analyses and (iii) analyse opsin mRNA expression. ERG measurements revealed two photoreceptor types in the compound eye, maximally sensitive to 527 and 364 nm. In addition, we identified three opsins in the genome, homologous to the hymenopteran green or long-wavelength sensitive (LW) LW1, LW2 and ultra-violet sensitive (UV) opsin genes. The LW1 and UV opsins were found to be expressed in the compound eyes, and LW2 and UV opsins in the ocelli. The lack of a blue or short-wavelength sensitive (SW) homologous opsin gene and a corresponding receptor suggests that S. noctilio is a UV-green dichromate.

Behavior and body size modulate the defense of toxin-containing sawfly larvae against ants.

The sawfly larvae of most Argidae and Pergidae (Hymenoptera: Symphyta) species contain toxic peptides, and these along with other traits contribute to their defense. However, the effectiveness of their defense strategy, especially against ants, remains poorly quantified. Here, five Arge species, A. berberidis, A. nigripes, A. ochropus, A. pagana, A. pullata, plus three Pergidae species, Lophyrotoma analis, Lophyrotoma zonalis, Philomastix macleaii, were tested in laboratory bioassays on ant workers mainly of Myrmica rubra. The experiments focused on short-term predator-prey interactions, sawfly survival rate after long-term interactions, and feeding deterrence of the sawfly hemolymph. The larvae of Arge species were generally surrounded by few ants, which rarely bit them, whereas larvae of Pergidae, especially P. macleaii, had more ants around with more biting. A detailed behavioral analysis of Arge-ant interactions revealed that larval body size and abdomen raising behavior were two determinants of ant responses. Another determinant may be the emission of a volatile secretion by non-eversible ventro-abdominal glands. The crude hemolymph of all tested species, the five Arge species and L. zonalis, was a strong feeding deterrent and remained active at a ten-fold dilution. Furthermore, the study revealed that the taxon-specific behavior of ants, sting or spray, impacted the survival of A. pagana but not the large body-sized A. pullata. The overall results suggest that the ability of Arge and Pergidae larvae to defend against ants is influenced by the body size and behavior of the larvae, as well as by chemicals.

The complete mitochondrial genome of Conaspidia wangi Wei, 2015 (Hymenoptera: Tenthredinidae) and its phylogenetic analysis.

The mitochondrial genome of Conaspidia wangi Wei, 2015 was described. The total length of the sequence was 15,924 bp. The overall A + T content was 80.4%. In comparison with the ancestral organization, trnG was reversed and translocated between the AT-rich region and trnQ, which was reported for the first time in Symphyta. The downstream gene order of the AT-rich region were thus arranged as trnG-trnQ-trnM-trnI.

Genomes of the willow-galling sawflies Euura lappo and Eupontania aestiva (Hymenoptera: Tenthredinidae): a resource for research on ecological speciation, adaptation, and gall induction.

Hymenoptera is a hyperdiverse insect order represented by over 153,000 different species. As many hymenopteran species perform various crucial roles for our environments, such as pollination, herbivory, and parasitism, they are of high economic and ecological importance. There are 99 hymenopteran genomes in the NCBI database, yet only five are representative of the paraphyletic suborder Symphyta (sawflies, woodwasps, and horntails), while the rest represent the suborder Apocrita (bees, wasps, and ants). Here, using a combination of 10X Genomics linked-read sequencing, Oxford Nanopore long-read technology, and Illumina short-read data, we assembled the genomes of two willow-galling sawflies (Hymenoptera: Tenthredinidae: Nematinae: Euurina): the bud-galling species Euura lappo and the leaf-galling species Eupontania aestiva. The final assembly for E. lappo is 259.85 Mbp in size, with a contig N50 of 209.0 kbp and a BUSCO score of 93.5%. The E. aestiva genome is 222.23 Mbp in size, with a contig N50 of 49.7 kbp and a 90.2% complete BUSCO score. De novo annotation of repetitive elements showed that 27.45% of the genome was composed of repetitive elements in E. lappo and 16.89% in E. aestiva, which is a marked increase compared to previously published hymenopteran genomes. The genomes presented here provide a resource for inferring phylogenetic relationships among basal hymenopterans, comparative studies on host-related genomic adaptation in plant-feeding insects, and research on the mechanisms of plant manipulation by gall-inducing insects.

Taxonomic and biological notes on Pristiphora ribisi Togashi, 1990, (Hymenoptera, Tenthredinidae) injurious to gooseberries and currants.

Taxonomic and biological notes are given for the East Asian sawfly Pristiphora ribisi Togashi, 1990, injurious to Ribes. The following new synonyms of this species are proposed: Pristiphora (Pristiphora) anivskiensis Haris, 2006, syn. nov., Pristiphora (Pristiphora) nigrocoreana Haris Zsolnai, 2007, syn. nov. and Pristiphora grossulariae Anon., 1912 [a primary homonym of Pristiphora grossulariae Walsh, 1866], syn. nov. The lectotype of Pristiphora grossulariae Anon., 1912 is designated. Redescriptions of the adult and immature stages are given. The male is described for the first time. The life history and host plants are summarized. Its close relative, Pristiphora appendiculata (Hartig, 1837), is excluded from the fauna of Japan.

Some sawfly larvae survive predator-prey interactions with pentatomid Picromerus bidens.

Most Asopinae stinkbugs (Hemiptera: Pentatomidae) prey on other insects, including sawfly larvae (Hymenoptera: Symphyta). Sawfly larvae of the Argidae and Pergidae contain toxic peptides, but whether they are defended against stinkbugs remains poorly studied. A literature survey indicates that no publication is devoted to laboratory tests specifically using these sawflies against stinkbugs. Here, laboratory bioassays were made with the stinkbug Picromerus bidens and four sawfly species at last larval instars: Arge ochropus (Argidae), Arge pagana (also tested at medium instars), Lophyrotoma zonalis (Pergidae), and Allantus rufocinctus (Tenthredinidae). Following 24 h of possible predator-prey interactions, no larvae of A. rufocinctus survived, whereas most or all larvae of the other sawfly species did survive and were still alive 48 h later. When feeding on an argid or pergid larva, the feeding periods lasted on average 6-20 s only, some bugs removing their rostrum and abruptly backing away. Full-grown larvae of A. pagana were attacked less than younger ones. It is likely that the tested Argidae and Pergidae are well defended against P. bidens by potent, internal antifeedants, while defensive body movements combined with a large body size play a secondary role.

Assessing temporal dynamics of predation and effectiveness of caterpillar visual defense using sawfly larval color and resting posture as a model.

Caterpillars (Lepidoptera and Symphyta larvae) employ diverse visual defensive tactics, and effectiveness of such tactics may be highly dynamic across time due to seasonal changes in the predator assemblages and their preferences. However, this has rarely been studied especially in tropical regions. Here we assessed temporal changes in the defensive value of caterpillar color and shape, using six types of plasticine dummy caterpillars: three colors (green, black, and white) × two shapes (curled and straight). These dummy caterpillars were deployed five times over different seasons in tropical forests of Xishuangbanna (China) and, as a comparison, twice in a temperate forest of Hirosaki (Japan). The colors and shapes of dummy caterpillars simulate visual traits of black sawfly larvae which take the curled resting posture in tropical rainforests of Xishuangbanna, apparently masquerading excrements commonly found on plants, while in Hirosaki there is no black-curled sawfly larvae and few excrements on plants. We found no significant effects of caterpillar colors or shapes on predation in Hirosaki. In contrast, black and curled caterpillars received significantly lower predation by birds in Xishuangbanna constantly across time. However, we were unable to provide evidence that the black-curled sawfly larvae are masquerading as excrements. Shapes of the dummy caterpillars also affected the predation by ants and parasitoid wasps at certain times. This is the first report on ecological function of the curled posture of sawfly larvae, and we demonstrated the importance to assess the temporal dynamics of predation and effectiveness of defensive tactics in tropical forests.

Caliroa slug sawflies of Japan (Hymenoptera, Tenthredinidae).

Japanese species of Caliroa (Hymenoptera, Tenthredinidae) are the following 13 species, five of which are described as new: C. aizankei Hara, sp. nov. from Hokkaido, C. bibaiensis Hara, sp. nov. from Hokkaido, C. cerasi (Linné, 1758), C. ibukii Hara, sp. nov. from Honshu, C. matsumotonis (Harukawa, 1919), C. nara Hara, 2011, C. nire Hara, sp. nov. from Hokkaido, C. oishii (Takeuchi, 1933), C. ouensis Hara, sp. nov. from Honshu, C. staphyleae Oishi, 1961, C. vaccini Okutani, 1965, C. varipes (Klug, 1816) and C. zelkovae Oishi, 1961. The following two new synonymies are proposed: C. quercivora Togashi, 1999, syn. nov. with C. oishii; C. sumomovora Togashi Oishi, 1978, syn. nov. with C. zelkovae. Lectotypes are designated for C. staphyleae Oishi, 1961 and C. zelkovae Oishi, 1961. Keys to adults and larvae are given. Caliroa staphyleae and C. zelkovae are newly recorded from Hokkaido. New host records are given for C. cerasi and C. zelkovae. Caliroa annulipes (Klug, 1816) is excluded from the fauna of Japan. An additional description of C. annulipes is given. The generic characters are also discussed.