journal of hymenoptera research

Thank you for visiting nature.com. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

• View all journals
• My Account Login
• Explore content
• About the journal
• Publish with us
• Sign up for alerts
• Open access
• Published: 03 March 2023

Key innovations and the diversification of Hymenoptera

• Bonnie B. Blaimer   ORCID: orcid.org/0000-0002-8961-9998 1 , 2 ,
• Bernardo F. Santos   ORCID: orcid.org/0000-0002-2634-3066 1 , 2 ,
• Astrid Cruaud   ORCID: orcid.org/0000-0001-8932-4199 3 ,
• Michael W. Gates   ORCID: orcid.org/0000-0002-5760-1371 4 ,
• Robert R. Kula   ORCID: orcid.org/0000-0002-1516-0360 4 ,
• István Mikó   ORCID: orcid.org/0000-0001-9719-0215 5 ,
• Jean-Yves Rasplus   ORCID: orcid.org/0000-0001-8614-6665 3 ,
• David R. Smith   ORCID: orcid.org/0000-0001-8502-9061 4 ,
• Elijah J. Talamas   ORCID: orcid.org/0000-0002-1048-6345 6 ,
• Seán G. Brady   ORCID: orcid.org/0000-0003-0468-940X 2 &
• Matthew L. Buffington   ORCID: orcid.org/0000-0003-1900-3861 4  

Nature Communications volume  14 , Article number:  1212 ( 2023 ) Cite this article

20k Accesses

27 Citations

117 Altmetric

Metrics details

• Comparative genomics
• Phylogenomics

The order Hymenoptera (wasps, ants, sawflies, and bees) represents one of the most diverse animal lineages, but whether specific key innovations have contributed to its diversification is still unknown. We assembled the largest time-calibrated phylogeny of Hymenoptera to date and investigated the origin and possible correlation of particular morphological and behavioral innovations with diversification in the order: the wasp waist of Apocrita; the stinger of Aculeata; parasitoidism, a specialized form of carnivory; and secondary phytophagy, a reversal to plant-feeding. Here, we show that parasitoidism has been the dominant strategy since the Late Triassic in Hymenoptera, but was not an immediate driver of diversification. Instead, transitions to secondary phytophagy (from parasitoidism) had a major influence on diversification rate in Hymenoptera. Support for the stinger and the wasp waist as key innovations remains equivocal, but these traits may have laid the anatomical and behavioral foundations for adaptations more directly associated with diversification.

Similar content being viewed by others

Harrison’s rule corroborated for the body size of cleptoparasitic cuckoo bees (Hymenoptera: Apidae: Nomadinae) and their hosts

The angiosperm radiation played a dual role in the diversification of insects and insect pollinators

Quantitative analysis of lacewing larvae over more than 100 million years reveals a complex pattern of loss of morphological diversity

Introduction.

The question of why some groups of organisms have diversified more than others has fascinated biologists since the early days of phylogenetics. The success of particular clades has often been attributed to the evolution of novel traits or key innovations conferring an evolutionary advantage 1 , 2 , and linking these traits to changes in diversification rates estimated from phylogenies has been a major goal in macroevolutionary research 3 , 4 , 5 , 6 . The concept of what defines a key innovation has been fluid 1 , 2 , 7 . Originally developed to describe traits that facilitate the radiation of a clade into new adaptive zones, it has more recently been generalized in macroevolutionary studies to describe an evolutionary change in a trait leading to increased species diversification in the clade that possesses this trait 1 , 6 , 8 . This broader definition has appeal due to its simplicity with regard to hypothesis testing; however, the idea of one trait single-handedly influencing survival and diversification has been criticized as overly simplistic 7 . More nuanced concepts and terminology have therefore been suggested for more complex evolutionary scenarios involving multiple traits 9 , 10 , 11 .

Much speculation and numerous hypotheses have attempted to account for the diversity of insects, summarized by Mayhew 12 . In particular, several morphological, physiological, and behavioral traits have been suggested as key innovations promoting the early diversification of insects and greatly influencing their success. These traits include the evolution of insect flight, complete metamorphosis, and phytophagy 6 , 8 , 13 , 14 , 15 , 16 . The Hymenoptera, which comprise the ants, bees, wasps, and sawflies, is one of the most species-rich and abundant insect orders 17 , 18 , possessing a remarkable diversity of life histories and morphological adaptations, some of which set records among the insects. For example, the smallest insect on Earth is a fairy wasp of the genus Dicopomorpha (Mymaridae) 19 , while the longest egg-laying organ (the ovipositor, measured in absolute size) occurs in Darwin wasps of the genus Megarhyssa (Ichneumonidae) 20 . These two extreme examples have a major life history strategy in common: they are both parasitoids, carnivores that complete their entire life cycle feeding on just one individual prey item, the host 21 . However, they are adapted to very different hosts: Dicopomorpha is an egg parasitoid of bark lice (Psocodea: Lepidopsocidae) 22 , while Megarhyssa is a parasitoid of wood-boring horntail larvae (Hymenoptera: Siricidae) 20 . A suite of morphological, physiological, and genetic adaptations provides the means for Hymenoptera to exploit a myriad of host niches 23 . In fact, about 70% of all described hymenopterans are parasitoids, while the other 30% are phytophages, such as leaf-feeding and wood-boring sawflies, gall-inducing wasps, and pollen-collecting bees, or predators, such as many social wasps. Among the insects, Hymenoptera is unique in that they comprise 75–80% of all described parasitoid insect species 24 , 25 . Parasitoidism and its associated features may be key drivers that explain diversification in Hymenoptera, as the adaptation to different host species and, therefore, niche subdivision may have resulted in increased speciation rates in parasitoids 26 .

Two other defining morphological features within the early evolution of Hymenoptera are putative key innovations. First, in the lineages of Hymenoptera comprising the grade of taxa commonly referred to as the “sawflies and wood wasps” (“Symphyta”), the abdomen is broadly attached to the thorax, which is the plesiomorphic ground plan for Hymenoptera 27 , 28 . In contrast, in apocritan wasps (suborder Apocrita), which comprise the Aculeata and numerous other primarily parasitoid lineages, the first abdominal tergum is fused with the metathorax forming the propodeum. The thorax and propodeum together form the mesosoma, which is connected to the remainder of the abdomen (the metasoma) via a constricted articulation 29 . It is this narrow wasp waist that gives female apocritan wasps the flexibility to bend their metasoma, enabling them to insert their ovipositor or stinger into a substrate, prey item, or enemy 29 , and thus potentially facilitating the success of both parasitoidism and stinging in apocritan wasps. Second, in the Aculeata, the female ovipositor is modified from an egg-laying to a stinging apparatus (the stinger) 30 . This group of stinging wasps also includes all social and colony-forming lineages of Hymenoptera, such as ants and certain groups of bees and wasps. In social hymenopterans, the stinger enables the defense of a nest and its brood, and possibly was an important factor in the diversification of Aculeata, as the increased defense system may have substantially decreased the risk of extinction in this clade.

While carnivory and parasitoidism are pervasive across the apocritan wasps, extant members of the sawfly and woodwasp lineages are generally phytophagous. This feeding strategy is therefore considered ancestral in Hymenoptera 14 . However, phytophagy has secondarily evolved in several groups of apocritan wasps, for example, in pollen-collecting bees or gall-inducing cynipoid wasps. Arguably, these transitions to a secondarily phytophagous life strategy were additional major innovations in Hymenoptera as they represent the formation of new niches (e.g., use of plant galls and pollen) and not reversals to a symphytan-style phytophagy. All four innovations—parasitoidism, the wasp waist, the stinger, and secondary phytophagy—could thus potentially be catalytic in the diversification of Hymenoptera 26 , 27 , 29 , 31 , 32 , 33 , yet they have never been analyzed across a phylogeny of the entire order in a macroevolutionary framework.

Phylogenetic relationships among major hymenopteran lineages have been the focus of much scrutiny over the last decade, based on either morphology or molecular data, or both 29 , 33 , 34 , 35 , 36 , 37 , 38 , 39 , but even the most recent phylogenomic analyses 36 , 38 , 40 were not able to provide clarity on the placement of some superfamily-level lineages, mainly due to sampling bias toward aculeate wasps. In this study, we infer a robust and balanced family-level phylogenetic framework from 765 taxa representing 94 families in Hymenoptera to study the early diversification and evolution of the four putative key innovations in Hymenoptera. Phylogenomic analyses are based on ~1,100 loci of ultraconserved elements (UCE), a group of markers widely used across higher-level groups of insects 38 , 41 , 42 , largely generated de novo by us. From these data, we estimate the most extensive phylogeny for Hymenoptera to date and establish support for hypotheses of superfamily-level relationships. Time-calibrated phylogenies infer the early evolution of parasitoidism in the order and allow us to investigate whether the four innovations outlined above are associated with the diversification dynamics of Hymenoptera by using macroevolutionary models. We discuss our results within a broader framework of insect evolution and diversification and give a perspective of future avenues for investigation.

Phylogenetic inference and major relationships within Hymenoptera

Our phylogenetic results are covered in detail in the Supplementary Discussion. In brief, to infer a robust phylogeny for Hymenoptera, we assembled our data set in several arrangements with varying levels of missing data and phylogenetic information content. We first filtered all captured UCE loci for taxon completeness, retaining a 50%, 60%, and 70% complete matrix (nuc-50% = 1118 loci, nuc-60% = 765 loci and nuc-70% = 446 loci) for further analyses. UCE loci were also matched to a reference protein database, and a 50% taxon-complete set (=324 loci) of protein-coding loci was created, which was further analyzed as nucleotide (prot-NUC) and amino acid (prot-AA) data set. These five matrices (nuc-50%, nuc-60%, nuc-70%, prot-AA, prot-NUC) were then analyzed via partitioned and unpartitioned analyses in a maximum likelihood framework. Supplementary Fig.  1 provides a flowchart overview of analyses; descriptions of data characteristics and filtering can be found in Supplementary Data  1 – 5 and in the Supplementary Methods, together with additional analyses that explored the impact of GC content on phylogenetic inference.

We recovered full support for two subdivisions within Apocrita, consisting of (1) Proctotrupomorpha, a clade including the superfamilies Chalcidoidea, Mymarommatoidea, Diaprioidea, Proctotrupoidea, Cynipoidea and Platygastroidea (ultrafast bootstrap support (ufBS) = 95–100, across the main analyses summarized in Fig.  1 and Supplementary Fig.  2 ), and (2) a clade consisting of Aculeata and the former “Evaniomorpha” (itself a grade). All analyses recover Orussoidea as a sister to Apocrita with ufBS=100 (Fig.  1 and Supplementary Fig.  2 ), corroborating Vespina (sensu Rasnitsyn 43 ). Our nucleotide data sets recovered Tenthredinoidea as the sister group to all other Hymenoptera (Fig.  1 and Supplementary Fig.  2 ), a unique result compared to previous studies. In the protein-coding, amino acid translated matrix, the sister group to all other Hymenoptera was Tenthredinoidea + (Xyeloidea + Pamphilioidea), similar to previous results using transcriptomes 36 . Full phylogenetic trees are shown in Supplementary Figs.  3 – 15 . We recovered three competing hypotheses regarding the position of Ichneumonoidea, in combination with three alternative positions of Ceraphronoidea. We summarized these by establishing mutually exclusive topology groupings based on the superfamily-level relationships (summarized in Supplementary Table  1 ). We used the best supported (based on ufBS and Four-cluster Likelihood Mapping (FcLM) 44 ) superfamily-level topology C-1 (=topC-1: Ichneumonoidea sister to the rest of Apocrita; Ceraphronoidea sister to Evanioidea + Stephanoidea) and the most frequently recovered topology A-0 (=topA-0: Ichneumonoidea sister to Ceraphronoidea, together sister to Proctotrupomorpha) as input for divergence dating and macroevolutionary analyses. However, the alternate positions of these two superfamilies had little impact on our results investigating the evolution of key innovations and the diversification history of Hymenoptera.

Phylogeny shows relationships between hymenopteran families as estimated from 446 UCE loci in the nuc-70%-SWSC analysis. This data set was partitioned using the Sliding-Window Site Characteristics Entropy (SWSC-EN) algorithm 106 and PartitionFinder2 107 in combination with the r cluster algorithm 108 and analyzed using Maximum Likelihood (ML) best-tree and ultrafast bootstrap searches in IQ-TREE v1.6.10 109 . This result is referred to as topology C-1 (topC-1) throughout the text and the remaining figures and tables. Various nodes have been collapsed for clarity of display, with numbers of species subtended by the respective branches included in brackets. Ultrafast bootstrap (ufBS) support values are indicated by colored squares on respective nodes: dark blue = 100%, light blue = 91–99%, and yellow = 70–90%. Support values lower than ufBS = 70 are not shown. Scalebar represents substitutions/bp. Source data for this figure can be found in the Dryad repository at https://doi.org/10.5061/dryad.08kprr54m (folder 2.1.7).

Timescale of hymenopteran evolution and their life histories

We estimated divergence times within Hymenoptera, employing 12 fossil calibrations as minima plus a minimum-maximum bound on the root node. Time-calibrated phylogenies were then used to reconstruct the evolution of life history strategies (parasitoidism, phytophagy, predation, and secondary phytophagy) in Hymenoptera. A model with a rate matrix of equal transition frequencies (ER) provided the best fit for all ancestral state reconstructions. Analyses using topC-1 and topA-0 generated similar results (Fig.  2 ; Supplementary Figs  16 – 19 ; and Supplementary Data  6 – 8 ); thus, mainly results based on topC-1 are discussed unless specified.

Chronogram estimated using approximate likelihood in MCMCTREE with the PAMLv4.9 package 113 from the nuc-70% matrix and topology C-1. All outgroups were pruned from the tree and alignment prior to divergence time estimation. Twelve calibration points were used, which are indicated by a white star. Terminals have been collapsed down to family or clade-level post analysis. Families of Platygastroidea are shown in the chronogram, but were lumped at the superfamily level for ancestral state reconstructions; therefore, pies are absent in this clade. Ancestral state reconstructions with corHMM were estimated from topology C-1 and mapped in pie format onto the chronogram; states for terminals are indicated beside terminal branches. Divergence estimates and ancestral state probabilities can be accessed in Supplementary Data  6 and 8 , referring to numbers beside nodes. Pie states are red = parasitoid, blue = predatory, brown = phytophagous, yellow = secondarily phytophagous, and white = behavior unknown. For detailed information on methodology and results, also refer to the main text and the Supplementary Methods. Source data for this figure can be found in the Dryad repository at https://doi.org/10.5061/dryad.08kprr54m (folder 2.2.2).

Hymenoptera was estimated with a median crown-group origin of 280 million years ago (Ma) in the Permian (Fig.  2 and Table  1 ). The sawfly and woodwasp lineages diverged from the remaining Hymenoptera, respectively, over the course of the next 50 million years, throughout the Permian and Early to Mid-Triassic (Fig.  2 ). Unicalcarida and Vespina (Orussoidea + Apocrita) originated in the Early Triassic (node 4 and 6, Fig.  2 ) around 248 Ma and 234 Ma, respectively. Crown-group apocritan wasps with their constricted wasp waist appeared just a few million years later, around 226 Ma (node 7, Fig.  2 ). Our ancestral state reconstructions confirmed that the earliest hymenopterans were plant-feeding insects similar to the extant members of the sawfly and woodwasp grade. The first transition between phytophagy and parasitoidism is estimated in the Late Triassic in the most recent common ancestor (MRCA) of Vespina (node 6, Fig.  2 ; Supplementary Figs.  18 – 19 ). Parasitoidism thus evolved once and remained the dominant life strategy in Hymenoptera, with no subsequent major innovations in life history evolving until the Early Cretaceous around 140 Ma (Fig.  2 ). Proctotrupomorpha and the Aculeata + “Evaniomorpha” grade both originated in the Late Triassic, around 210 Ma and 206 Ma, respectively (node 9 and node 24, Fig.  2 and Table  1 ). The origin of crown-group Ichneumonoidea is also estimated within a similar timeframe in the Late Triassic (206 Ma, node 64; Fig.  2 ), while most of the other non-aculeate superfamilies originated throughout the Jurassic and Early Cretaceous (Table  1 ). Aculeata was estimated with an age of 142 Ma (node 26; Fig.  2 ), placing the origin of the modified ovipositor-stinger at the edge of the Jurassic-Cretaceous boundary. Within Aculeata, three transitions to a non-parasitoid strategy are reconstructed in the Late Cretaceous (starting ca. 80 Ma, Fig.  2 ), two to a mixed predatory and secondarily phytophagous lifestyle in vespid wasps and ants, and one to secondary phytophagy (pollen-collecting) in bees (node 36, Fig.  2 ). Another major transition to secondary phytophagy, in this case plant-galling, occurred within Cynipoidea in the Early Cretaceous (around 105 Ma, node 20, Fig.  2 ). Several additional adaptations to plant-galling in Chalcidoidea are not captured by these results, as all chalcidoid families were collapsed into one clade and coded as polymorphic in our analyses. Independent transitions from a parasitoid to a predatory lifestyle occurred in Evaniidae and Gasteruptiidae.

Diversification history of Hymenoptera

Net diversification rates and potential rate shifts in Hymenoptera were estimated using Bayesian Analysis of Mixture Models (BAMM 45 , 46 ) and stepwise AIC (MEDUSA 47 ). In BAMM, we estimated and plotted the best shift configuration with maximum a posteriori probability (MAP), mean phylorate, and cumulative shift probabilities. Six (topC-1) and eight (topA-0) rate shifts were estimated in the best shift configurations (Fig.  3a , Table  2 , and Supplementary Fig.  20a ). To quantify all rate shifts, we calculated the mean net diversification rates (estimated with BAMM) for all clades for which rate shifts were indicated by either of the above analyses (Table  2 ; Supplementary Data  9 ). Shifts in net diversification rate that are supported by BAMM across both topologies occur in Aculeata minus Chrysidoidea (non-chrysidoid aculeates hereafter), within the bees minus family Melittidae (non-melittid bees hereafter), on the branch leading to Cynipidae s.s. (sensu stricto, or in the narrow sense), on the branch leading to (or within) Eurytomidae (Eurytominae), and within Ichneumonidae (shifts 3–6, and 1, respectively; Fig.  3 and Supplementary Fig.  20 ). All of these shifts represent increases in net diversification compared to the background rate across the phylogeny (Table  2 ). Additionally, for topC-1 a rate increase (i.e. positive shift) is estimated on the branch leading to Apocrita minus Ichneumonoidea (shift 2, Fig.  3a , Table  2 ), while for topA-0 rate decreases (i.e. negative shifts) are estimated on the branches leading to Tenthredinoidea and Ichneumonoidea (shifts 7, 8), and a rate increase on the branch leading to Apocrita (shift 2) (Supplementary Fig.  20a , Table  2 ). These MAP shift configurations only had a probability of 0.019 (topC-1) and 0.009 (topA-0) among 9910 and 12,354 distinct shift configurations in the 95% credibility set, respectively; however, the next-best shift configurations were very similar (Supplementary Fig.  21 ).

We assessed potential shifts in diversification rates over time in Hymenoptera using a sampling fraction approach and clade-specific sampling probabilities in BAMM v2.5 45 , 46 and the associated R package BAMMtools v2.1.7 115 . All analyses shown are based on topology C-1. a Plot of best shift configuration with maximum a posteriori probability (MAP), indicating rate shifts on respective branches. Rates are shown as net diversification rates. Since this rate shift configuration only has a 0.019 probability among 9910 distinct shift configurations in the 95% credibility set, we summarized the cumulative probabilities for each branch that a shift occurred somewhere between the focal branch and the root of the tree. b Cumulative shift probability tree, indicating in dark cyan the branches with a shift probability ≥0.97 ≤ 0.99 and in gold the branches with a cumulative shift probability = 1. Golden branches thus occur in every distinct shift configuration and cyan branches in 97–99% of all distinct shift configurations. Numbered clades/shifts are (1) Ichneumonidae (internal); (2) Apocrita minus Ichneumonoidea; (3) Aculeata minus Chrysidoidea; (4) Bees minus Melittidae; (5) Cynipidae s.s.; (6) Eurytomidae (Eurytominae). For detailed information on methodology and results, refer to the main text, Supplementary Methods, and Supplementary Data  9 and 16 . Images of wasps courtesy of Matt Bertone. Source data for this figure can be found in the Dryad repository at https://doi.org/10.5061/dryad.08kprr54m (folder 3.2).

Due to the low probability for MAP shift configurations, we summarized support for shifts also by plotting cumulative shift probability (CSP) trees, which show the cumulative probability for each branch that a shift occurred somewhere between the root of the tree and the branch under scrutiny. For topC-1, the CSP tree clearly shows that only the positive shifts leading to the non-melittid bees, Cynipidae s.s., Eurytomidae (Eurytominae), and Ichneumonidae (internal) are present across most of the credible shift set (0.97–1.0 probability) (Fig.  3b ). Additionally, a shift is indicated leading up to Chalcidoidea that is not present in the best shift configuration. For topA-0, all shifts in the best shift configuration are present in the CSP tree with high probability, except the negative rate shifts on the branches leading to Tenthredinoidea and Apocrita (Supplementary Fig.  20b ).

MEDUSA analyses estimated six (topC-1) and five (topA-0) distinct rate regimes within Hymenoptera, which are summarized in Table  3 (see also Supplementary Fig.  22 ). Both analyses agreed on a positive shift in diversification rate within non-chrysidoid aculeates and within non-melittid bees (shifts ii and iii, Table  3 , respectively). The analysis using topC-1 additionally estimated shifts to a faster rate regime along the branch leading to Cynipoidea and further to a core cynipoid clade comprising Cynipidae s.s. and Figitidae s.l. (sensu lato, in the wider sense) (shifts vi and v, Supplementary Fig.  22a ; Table  3 ). Both analyses estimated a negative shift in diversification rate ancestral to Xiphydriidae + Siricidae (shift iv, Table  3 , and Supplementary Fig.  22 ), and the topA-0 analysis further estimated a decreased rate regime for Pamphiliidae + Xyelidae (shift v, Table  3 , and Supplementary Fig.  22b ).

Thus, two positive rate shifts are supported by the BAMM MAP configuration and MEDUSA: one on the branch to non-aculeate chrysidoids and one for non-melittid bees. Both analyses also place positive shifts within Cynipoidea but at varying locations. Rates for all clades that had a shift indicated in either analysis show a corresponding deviation from the background rate. Interestingly, net diversification rates for Vespina and Aculeata were elevated compared to the background rates, although no shifts were indicated in these clades. The highest net diversification rates are estimated within Ichneumonoidea, Cynipoidea (Cynipidae s.s.), Chalcidoidea (Eurytomidae: Eurytominae), and for non-melittid bees (Table  2 ); these rate patterns are also visible in mean phylorate plots (Supplementary Fig.  23 ).

The association of four key innovations (parasitoidism, wasp waist, stinger, secondary phytophagy) with diversification in Hymenoptera was assessed using 30 models in the HiSSE (Hidden State Speciation and Extinction) framework (Supplementary Data  10 ), which can incorporate unobserved traits (hidden states) that may influence diversification rate together with the observed traits 48 . We also analyzed carnivory and phytophagy (i.e., primary + secondary) as the fundamental feeding strategies, including parasitoidism and secondary phytophagy, respectively. HiSSE identifies the most probable combination of the observed trait as being absent (0) or present (1) with the estimated hidden state in state A or B, resulting in combined state inferences such as 0A, 0B, 1A, and 1B, which are reported below. We further estimated tip diversification rates across Hymenoptera using MiSSE (Missing State Speciation and Extinction), a trait-free extension of the HiSSE framework. Parameter estimates from best-fitting models are summarized in Supplementary Data  10 – 13 , and visualized in Fig.  4 and Supplementary Figs.  24 – 28 .

We tested 30 models of trait-dependent and trait-independent diversification in the HiSSE framework 48 for the four putative key innovations and plotted net diversification rates and state reconstructions from the best-fitting model onto the Hymenoptera phylogeny. All results shown were based on topology C-1. Inner branch colors represent the presence/absence of the respective states (black = presence; gray = absence); outer branch colors represent the net diversification rate (highest = gold, lowest = dark cyan). Clades discussed in the context of the results are labeled. The histograms in the lower right of each panel represent the distribution of net diversification rates associated with the observed states (0 = absent, 1 = present). a Wasp waist, model: HiSSE—full, irreversible states; b Stinger, model: CID-4−9 distinct transition rates; c Parasitoidism, model: HiSSE—full, all free parameters; d Secondary phytophagy, model: HiSSE—full, irreversible states. Best-fitting models for the wasp waist ( a ), parasitoidism ( c ), and secondary phytophagy ( d ) are trait-dependent models that suggest the association of a hidden state with the analyzed traits influencing diversification in Hymenoptera. The best-fitting model for the stinger ( b ) is a trait-independent model. For detailed information on methodology, results, and model scores, refer to the main text, Supplementary Methods, and Supplementary Data  10 – 12 and 16 – 17 . Source data for this figure can be found in the Dryad repository at https://doi.org/10.5061/dryad.08kprr54m (folder 3.3).

Trait-dependent diversification in association with a hidden state was supported for the wasp waist, as a full HiSSE model with irreversible states was found to be the best-scoring model (Table  4 ). Trait-diversification plots show a higher estimated diversification rate in association with the wasp waist (Fig.  4a ; Supplementary Fig.  25a ), and net diversification rate showed >100× increase after the transition to the wasp waist in combination with hidden state A and 8.2–11.9× in combination with hidden state B (Supplementary Data  11 ). Most tip states are estimated with higher probabilities in state 1A (Supplementary Data  12 ); however, state 1B has a higher net diversification rate than state 1A. Taxa whose marginal probabilities for state 1B are on the higher side (up to 0.58) overlap to some extent with clades that showed elevated rates in BAMM and MEDUSA analyses (Supplementary Data  12 ), but elevated probabilities for state 1B are also seen in taxa outside these clades.

An association of parasitoidism with diversification in Hymenoptera in combination with a hidden state was suggested, as the full HiSSE model received the most support (Table  4 ). However, a 0.83–0.84× decrease in net diversification rate from the non-parasitoid to the parasitoid state in combination with hidden state A (i.e., 0A to 1A) was indicated (Supplementary Data  11 ; Fig.  4c ; Supplementary Fig.  25b ), and this state was estimated as the likeliest state for most parasitoid taxa (Supplementary Data  12 ). By contrast, the net diversification rate of the parasitoid state in combination with hidden state B (i.e., 1B) increased by 17.7–29.2×. The highest net diversification rate was estimated for state 0A, yet all non-parasitoid taxa were reconstructed with state 0B as the most probable. Interestingly, some taxa in clades with rate shifts recovered (BAMM shifts 1, 3, 6, 8 and MEDUSA shifts ii, v, vi; Table  4 ) were estimated with a significant proportional probability in state 1B (up to 0.71), and taxa with probabilities on the higher side (up to 0.38) for state 0A clustered predominantly (but not exclusively) in clades with rate shifts (4, 5, iii; Table  4 ) as well.

For secondary phytophagy, trait-dependent diversification was also supported by a full HiSSE model with irreversible states (Fig.  4d ; Supplementary Fig.  24b ); net diversification rates showed a 1.6–21.1× increase in the observed trait in combination with hidden state A and a 0.33–0.95× increase in association with hidden state B, although state 1B has the highest net diversification rate (Supplementary Table  11 ). Most secondarily phytophagous clades (largely congruent with the rate shifts 4–6 and iii recovered by BAMM and MEDUSA; Table  4 ) are estimated with state 1A as the most probable, but also have a high proportion of probabilities for state 1B (up to 0.58; Supplementary Table  12 ). For non-secondarily phytophagous taxa the pattern is reversed, with state 0A being the most probable, but 0B having the faster net diversification rate. Trait-dependent diversification for the fundamental feeding strategy phytophagy was only supported by topC-1, whereas a character-independent model scored best for topA-0 (Table  4 , Supplementary Fig.  27 ). Character-independent models had the highest support for the stinger and carnivory (Table  4 ; Fig.  4b ; Supplementary Figs.  24a and 26 ), indicating that these traits were not associated with diversification rate.

The best-scoring models tested with MiSSE incorporated five (topC-1) and six (topA-0) hidden states. Tip net diversification rates ranged from 0.048 to 0.072 (topC-1) and 0.051 to 0.079 (topA-0). Several clades are highlighted with higher diversification rates (Supplementary Data  13 ; clades 1–4, Supplementary Fig.  28 ). These are congruent with BAMM/MEDUSA rate shifts 1, 4/iii, 5, and 6, three of which are associated with secondary phytophagy. The same hidden states (topC-1: 0D; topA-0: 0A) are estimated as most probable in these four clades (but not across most other taxa), suggesting that these states may be associated with secondary phytophagy and a higher diversification rate.

Our work targeted an increased sampling of parasitoid lineages to create an improved phylogenomic framework to evaluate the association of key traits with the diversification of Hymenoptera. A detailed discussion of the phylogenetic insights of our work can be found in the supplementary information accompanying this article. Below we focus on the implications of our time-calibrated hymenopteran phylogeny on the evolution of putative key innovations and the diversification of the order.

The pervasiveness of parasitoidism across Vespina (Orussidae and Apocrita) suggests the ancestry and antiquity of this behavior in Hymenoptera. Several attempts to trace the evolution of the parasitoid lifestyle have been made 24 , 26 , 49 , but our chronogram has enabled us to reconstruct the origin of parasitoidism with a maximum likelihood-based approach. We find a single evolution of parasitoidism most probable in the MRCA of Vespina, dating to the late Triassic (around 234 Ma). Branstetter et al. 38 , Peters et al. 36 , and Ronquist et al. 39 estimated the age of Vespina at ∼ 200 Ma, ∼ 247 Ma, and ∼ 270 Ma, respectively. Likely reasons for incongruities are differences in taxon composition between ours and the previous analyses, and different fossils and calibration strategies (i.e., node-dating in the present, and Branstetter et al. 38 and Peters et al. 36 analyses, whereas Ronquist et al. 39 also used a tip-dating and total-evidence approach). Naturally, the discovery of new fossil evidence may have unforeseeable consequences on our current estimates and the derived diversification patterns. Different sizes and types of sequence data (UCEs vs. transcriptomes in Peters et al. 36 and seven mitochondrial and nuclear markers in Ronquist et al. 39 ) and methods of divergence dating (MCMCTREE used by us and Peters et al. 36 , BEAST 50 in Branstetter et al. 38 , and MrBayes 3.2 51 in Ronquist et al. 39 ) most certainly also have had an impact on divergence estimates. We compare the above aspects in more detail in the supplementary discussion; overall, our results are closest to Peters et al.’s 36 estimates (Supplementary Data  6 ).

Our data suggest that parasitoidism has been the dominant life strategy in Hymenoptera since the late Triassic. Several subsequent switches from a parasitoid strategy to a secondarily phytophagous or predatory habit occurred, yet markedly only from the mid-Cretaceous onward. Diversification of angiosperms may have facilitated the evolution and diversification of gall-inducing and pollen-collecting hymenopterans, as evidenced by the known cases of codiversification between plants and their pollinating hymenopterans 52 , 53 , 54 , 55 . For angiosperm diversification to be a facilitator of secondary phytophagy, we would expect the onset of the former first, though not necessarily closely followed by the origins of the latter. The timing of the origin of the angiosperms is contentious, but most estimates (130–180 Ma 56 ) indeed predate the evolution of secondarily phytophagous clades (from ca. 105 Ma onwards based on our estimations) in Hymenoptera. Interestingly, the ages of secondarily phytophagous clades correspond with the beginning of the “Angiosperm Terrestrial Revolution” ca. 100 Ma 57 . Angiosperm diversification has also been suggested as a major element in the diversification of tenthredinoid sawflies, which are presently largely angiosperm-feeding ( ∼ 85% 58 ) and estimated in our analyses with a crown-group origin around 170 Ma.

Despite the more recent transitions to secondary phytophagy (and predation), the single appearance and long dominance of parasitoidism in Hymenoptera is striking. Parasitoidism does occur in other holometabolous insects, namely in the orders Diptera, Coleoptera, Lepidoptera, Trichoptera, and Neuroptera 24 , 59 , but Hymenoptera harbors the majority of parasitoid diversity (75–80% 24 ). In Diptera and Coleoptera, the other orders with substantial parasitoid members, parasitoidism has evolved repeatedly 24 , 60 but probably much more recently, at least in Diptera 61 . Our timeline of parasitoid diversification in Hymenoptera is generally older than scenarios suggested by the fossil record alone, which depict an expansion of insect parasitoid clades only starting in the late Early Jurassic and Early Cretaceous (“Mid-Mesozoic Parasitoid Revolution” 59 ). Yet our data imply a parasitoid regime already in full swing in the late Triassic. According to our current understanding of insect diversification 62 , parasitoidism was likely already dominant in Hymenoptera when many of their current primarily holometabolous host groups (e.g., Diptera and Lepidoptera) began to diversify. Thus, diversification in parasitoid Hymenoptera may have been mediated by both the long history and antiquity of the behavior, as a consequence of the longer timeframe for speciation events to take place and the opening of new niche space and resources by tracking host lineage diversification.

What roles have key innovations played in the diversification of Hymenoptera? Several clades are highlighted with a history of increased diversification rates across our analyses, related to some degree to the evolution of the four traits under investigation. An association of the wasp waist with diversification in Hymenoptera in combination with a hidden state was supported by HiSSE analyses, and both combinations of observed and hidden states show a significant increase in net diversification rates in apocritan waisted wasps. This confirms that the diversification patterns are largely driven by the hidden state(s), as there was no support from MEDUSA or BAMM analyses for a rate shift at the origin of Apocrita. Interestingly, BAMM and MEDUSA analyses recovered negative shifts or net diversification rate decreases in non-apocritan clades (i.e. Tenthredinoidea, Siricidae + Xiphydriidae, Pamphiliidae + Xyelidae), suggesting that the disparity in species richness we see between apocritan and non-apocritan lineages today may be due at least in part to a slowdown of diversification and higher extinction in the latter. It is possible that the rate increase associated with hidden states in the presence of the wasp waist is driven by the fact that lineages with this trait contain all the inferred rate shifts, while the absence of the wasp waist is restricted to lineages with low present-day diversity. Despite the wasp waist being undeniably a major morphological innovation in Hymenoptera 29 , there is thus no evidence that this character immediately accelerated the diversification of Apocrita.

Trait-dependent diversification was not supported for the stinger, although a positive diversification rate shift in non-chrysidoid Aculeata received support across BAMM and MEDUSA analyses, and net diversification in this clade is elevated compared to the background rate and the general rate in Aculeata. This observed rate shift is only imperfectly aligned with the origin of the stinger, excluding chrysidoids, which form a grade respective to all other aculeates. While the stinging apparatus evolved in the ancestor of Aculeata, this complex unit is composed of structures that have undergone several modifications. For example, all aculeates except chrysidoids have the third valvula of the sting shaft subdivided, a putative synapomorphy for this group, potentially leading to greater stinging precision 63 . Moreover, the loss of the egg delivery function in favor of exclusive delivery of venom by the ovipositor may not have been simultaneously fixed in the MRCA of aculeates, as there are several members of Chrysidoidea that still use the ovipositor to deliver eggs (e.g., Dryinidae, Embolemidae, Sclerogibbidae, and Chrysididae 64 , 65 ). Some of the innovative functions of the stinger may, therefore, only have had an impact on accelerating diversification once a series of modifications were completed. In any case, support for trait-independent diversification does not necessarily indicate that the aculeate stinger had no effect on the diversification rate at all, as it could be that elevated rates in other non-aculeate clades dilute the signal recovered for the stinger in statistical analyses.

We found indication for an association of parasitoidism with net diversification rate in combination with hidden states, but HiSSE results show an initial decrease in net diversification rate in Vespina rather than an increase. Parasitoidism can be viewed as a highly specialized strategy of carnivory (including predation), for which a character-independent model was supported. This may suggest the evolutionary significance of parasitoidism per se (as opposed to any form of carnivory) for the diversification in Hymenoptera. However, support for a hidden state model for parasitoidism over a character-independent model was not overwhelming, and there was no support by MEDUSA or BAMM analyses for a rate shift coinciding with the evolution of parasitoidism in Vespina. Wiegmann et al. 66 investigated parasitism (this broader definition also includes groups that do not kill their host) as a driver of diversification across insects using sister-clade comparisons but found no consistent association with increased diversification. They suggested that the specialization necessary for parasites to adapt to their host may actually decrease their evolutionary potential and negatively influence diversification 66 . In the case of Hymenoptera, our results indicate that parasitoidism evolved circa 230 Ma. This may have been too early for the group to have immediately benefited from the dramatic diversification of other holometabolous orders, which seems to have taken place in the last 150 million years 61 , 67 , 68 , 69 . Hence, the evolution of parasitoidism did not necessarily coincide with an abundance of available niches (in the form of host species) that would spur rapid adaptive radiation and lead to an increase in diversification rate. Instead, there may have been a lag time between the origin of parasitoidism and the diversification of parasitoid hymenopterans until their primarily holometabolous hosts became abundant. Such a delay in response to diversification has been observed in other systems as well 10 , for example, in the evolution of C 4 photosynthesis in grasses 70 .

Several rate shifts were suggested ancestral to or closely related to secondarily phytophagous clades. For instance, one of the rate shifts supported by both BAMM and MEDUSA highlights non-melittid bees as a clade with an exceptionally high net diversification rate. This rate shift is loosely associated with what could be seen as a key behavioral transition, a shift from parasitoidism to pollen collecting in bees, and confirms a result from an earlier analysis focusing on bee diversification dynamics 31 . Melittid bees are oligolectic, narrowly adapted to pollen-feeding from a few host plants, while many other bees are adapted to a wider host breadth (polylecty) and, thus, a broader ecological niche. Given the sister-group relationship of Melittidae to the rest of the bees, oligolecty is suggested to be the ancestral condition 71 . Murray et al. 31 suggested that this expansion of the host plant niche led to increased diversification in non-melittid bees. Similar to a hypothesized association of an advanced stinger with diversification in non-chrysidoid aculeates, polylecty could be seen as an advanced form of pollen-collecting in bees, the secondary innovation hereby driving diversification. Most analyses indicated some support for a positive diversification rate shift within Cynipoidea and particularly Cynipidae s.s., a group of gall-forming wasps, while another rate shift is suggested by BAMM within Eurytominae (Eurytomidae, Chalcidoidea), a group with multiple independent origins of phytophagy, commonly as gall associates (inducers or secondary feeders) or seed feeders. Both Cynipidae s.s. and Eurytominae were also estimated with very high net diversification rates (0.112–0.115 and 0.173–0.185, respectively). However, while our taxon sampling for cynipoids and bees was balanced and representative of species diversity, we cannot discount that a possible bias drives the accelerated diversification rates seen in Eurytominae within the more sparsely sampled Chalcidoidea, and the validity of the eurytomine rate shift, therefore, remains unclear.

The location of several diversification rates shifts ancestral to or within secondarily phytophagous clades, as well as their congruence with trait-dependent and tip diversification patterns (i.e. HiSSE and MiSSE), suggest that this life strategy played a major role in diversification. Phytophagy has repeatedly been suggested to promote diversification in insects 12 , 15 , 16 , 68 , 72 , 73 . Yet, the comparatively lower diversity of phytophagous clades appeared to be a major argument against applying this hypothesis to Hymenoptera. The ancestrally phytophagous “symphytan” lineages have not diversified exceptionally (7882 vs. 144,809 described species in Apocrita at the time of writing 18 ), and support for trait-dependent diversification for (fundamental) phytophagy also remained equivocal. Taken together, these results highlight the role of secondary phytophagy, but not phytophagy per se, in the more recent diversification history of Hymenoptera. This leads to the question: what adaptations do a parasitoid-converted-to-phytophage possess that may confer an evolutionary advantage over a primary phytophage?

We hypothesize that secondary phytophagy enabled further success because it is derived from the common parasitoid behavior of provisioning for their offspring by laying their eggs directly on or into a food resource 21 . This basic form of parental care increases the survival of offspring and could lead to a decreased extinction rate in the provisioning lineages compared to non-provisioning groups, although in our reconstructions, only one of the two hidden state combinations was associated with a lower extinction rate. While some forms of parental care, such as egg guarding, also occur in extant “symphytan” lineages (e.g., Argidae and Pergidae 74 ), parasitoids are particular in that they generally provide all resources (i.e., food and shelter or a nest) needed for the development of their offspring. Secondarily phytophagous hymenopterans such as bees, cynipids, or eurytomines may have retained this strategy from their parasitoid ancestors. This may have allowed secondarily phytophagous groups to more efficiently explore new plant-based food resources (such as pollen or concealed feeding in plant galls), providing a level of larval provisioning not previously possible. Diversification of these clades may have accelerated via a combination of decreased extinction and increased speciation rate as a response to the adaptation to these new trophic niches and the escape from the competition with parasitoid groups. In this context, parasitoid behavior potentially could be considered a pre-innovation or precursor (sensu Donoghue and Sanderson 10 ) to the evolution of specialized phytophagous strategies such as gall-inducing or pollen-collecting, and possibly represent the unobserved, hidden states influencing diversification rate. Of course, this scenario is not supported for the aforementioned melittid bees and other rather species-poor secondarily phytophagous clades, such as pollen wasps (Vespidae: Maserinae) or Krombeinictus , a monotypic genus of apoid wasps; conversely, it does not fit with the absence of rate shifts in the species-rich ants and the remaining vespid wasps.

Our study applied a broad definition of the key innovation concept widely used in the macroevolutionary literature; however, this interpretation has been criticized recently as being too simplistic 7 , 9 , 10 . We argue that an operational, simple definition is still desirable as a first step for macroevolutionary tests of key innovations: if a trait shows a strong correlation with a diversification rate increase in a clade, this may highlight its relevance in the evolutionary history of that lineage. However, in case of an absence of trait-dependent diversification, the reverse conclusion may be less straightforward. For example, there may be a time lag between the innovation and the increase in diversification rate, in which case statistical tests will fail to detect a correlation 10 . Confirming key innovation hypotheses when the trait under scrutiny has evolved only once or early on in the evolution of a group, as the case for several of our innovations, may also be particularly challenging from a statistical and conceptual point of view 7 , 48 , 75 . Extinction may have a considerable influence on net diversification in some clades as well, despite high speciation rates connected to innovation. Extinction rates are notoriously difficult to estimate from phylogenies 76 , and some authors have cautioned against inferring diversification dynamics from timetrees altogether 77 . Recent work further suggests that the location of diversification rate shifts may be influenced by sampling completeness and the number of alignment sites analyzed 78 . We show that our results are largely congruent and robust across data sets of different sizes and types of analyses, and careful measures were taken to minimize the effect of taxon sampling. Our diversification analyses corrected for unsampled diversity on the clade level by using a backbone sampling frequency for accounting for the few lineages (i.e. families) missing from our analyses. As these were mainly lineages with low diversity, we believe their exclusion likely had little effect on the overall diversity estimates. On the species level, we employed clade-specific taxon and trait sampling frequencies, which represent the only possible strategy for such a diverse group as Hymenoptera, for which complete phylogenetic sampling is out of reach with the current methodology. Sampling frequencies, of course, can only be as accurate as the underlying estimates of described species and family-level diversity, and it is, therefore, possible that future improvements in these estimates will warrant updated analyses.

In conclusion, our results indicate that the evolution of secondary phytophagous strategies has played a prominent role in the diversification of Hymenoptera, while the impact of the wasp waist, the aculeate stinger, and parasitoidism as direct accelerators of diversification or key innovations in the traditional sense remains unclear. We suggest that modifications or specializations of these latter three prominent characters and behaviors, rather than their first appearance, may relate to the diversification of Hymenoptera. This may be a common scenario, as many traits perceived as important innovations seemingly only had major evolutionary impacts when combined with additional adaptations. For instance, while winged flight has been traditionally considered a key innovation in insects, it was only after the evolution of wing flexing (in Neoptera) that insect diversity rapidly expanded in terms of both species diversity and niche occupation 14 . Therefore, we propose parasitoidism, the wasp waist and the stinger may be part of more complex character synergies akin to “synnovations” (sensu Donoghue et al. 10 ), i.e., characters that interact synergistically with other traits to open new evolutionary pathways. Future research should focus on dissecting these traits into their functional subcomponents to relate them with biological implications. A more nuanced analysis of these subcomponents may reveal innovations that can more directly be associated with diversification events. Searching for fossils that improve knowledge of divergence timing could also prove essential to better link innovations with diversification events. That parasitoidism was a superbly successful strategy in Hymenoptera cannot be disputed, given its long dominance in Vespina. Yet, among the characters assessed as potential key innovations by our study, only secondary phytophagy in the parasitoid lineages has left a discernible, direct imprint on diversification dynamics within the evolutionary history of the order.

Taxon sampling

We assembled a taxon set of 771 species across 94 out of 109 recognized extant families (sensu Huber 18 , with modifications by Chen et al. 79 , Pilgrim et al. 80 , and Sann et al. 81 ), belonging to all 22 recognized superfamilies within the Hymenoptera 18 , 80 , and six non-hymenopteran outgroups. Our taxon sampling aimed for the representation of major lineages within families while sampling across the respective root nodes on the family level, covering between 0.06–50% (=1–150 representatives) of the described species diversity. While we generated UCE sequence data de novo for most taxa, some sequences have already been published in other studies by some of us: 126 aculeate wasps 38 , 82 , 83 , 25 chalcidoids 84 , 85 , 86 , 76 cynipoids 87 , 26 Ichneumonidae 88 , 89 , 90 and 142 Braconidae 91 . We further included six representatives of other insect orders as outgroups by mining UCEs in silico from published genomes: Coleoptera ( Agrilus planipennis ), Diptera ( Aedes albopictus ), Lepidoptera ( Papilio glaucus ), Hemiptera ( Homalodisca vitripennis ), Psocodea ( Pediculus humanus corporis ), and Blattodea ( Blattella germanica ). Supplementary Data  1 list voucher information and NCBI accession numbers for all sequences, while more detailed specimen data is provided for sequences newly released in this article. All specimens were collected with the required permits and in accordance with local regulations at the time of their collection, and vouchers have been deposited in major collections.

UCE data collection and processing

We collected UCE data for this and allied studies using well-established library preparation and target enrichment protocols 92 , 93 , 94 , which we summarize in the following. Most UCE laboratory work was conducted in and with the support of the Laboratories of Analytical Biology (L.A.B.) facilities of the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA. Genomic DNA was extracted destructively or non-destructively (specimen retained after extraction) from whole specimens using the DNeasy Blood and Tissue Kit (Qiagen, Valencia, CA, USA), and quantified for each sample using a Qubit fluorometer (High sensitivity kit, Life Technologies, Inc., Carlsbad, CA). Between <5 ng and 1364 ng DNA was sheared for 0–60 s (amp = 25, pulse = 10) to a target size of approximately 250–600 bp by sonication (Q800, Qsonica Inc., Newtown, CT), depending on prior DNA degradation. A modified genomic DNA library preparation protocol (Kapa Hyper Prep Library Kit, Kapa Biosystems, Wilmington, MA) was applied to incorporate bead-based cleanup steps 95 and a generic SPRI substitute 96 as described by Faircloth et al. 82 , as well as TruSeq-style adapters during adapter ligation 97 . Libraries had post-PCR concentrations from 0.1 to 102 ng/µL. Library input statistics are provided in  Supplementary Data 1 . Groups of eight to ten libraries were combined at equimolar ratios, and each pool was enriched using a set of custom-designed probes (MYcroarray, Inc., now ArborBiosciences, Ann Arbor, MI) targeting 2590 UCE loci in Hymenoptera 98 (now sold as predesigned panel myBaits UCE Hymenoptera 2.5Kv2P). The pooled libraries were sequenced using several lanes of 125 bp paired-end sequencing on an Illumina HiSeq 2500 instrument. For 36 chalcidoid taxa, UCE data were generated at Center de Biologie et de Gestion des Populations (CBGP), Montpellier, France using the myBaits UCE Hymenoptera 1.5Kv1 panel 82 with similar protocols 84 , and sequenced on an Illumina MiSeq instrument. UCE sequences for the six non-Hymenoptera outgroup taxa were captured from genome assemblies published on NCBI ( www.ncbi.nlm.nih.gov , see Supplementary Data  1 for accession numbers), using scripts provided within the PHYLUCE package v.1.5.0 99 . We followed the tutorial “harvesting UCEs from genomes” ( https://phyluce.readthedocs.io/en/latest/tutorials/tutorial-3.html ), except we reduced the stringency of the minimum coverage parameter of the phyluce_probe_run_multiple_lastzs_sqlite script (-minCov = 50) and used the myBaits UCE Hymenoptera 2.5Kv2P panel as input. We captured between 235 and 634 UCE loci for outgroup taxa.

All UCE data were processed using scripts within the PHYLUCE package 99 . We first trimmed the demultiplexed FASTQ data output for adapter contamination and low-quality bases using Illumiprocessor v2.0.7 100 , based on the package Trimmomatic v0.32-1 101 . We assembled the cleaned reads using the program Trinity (version trinityrnaseq_r20140717) 102 and a wrapper script (phyluce_assembly_assemblo_trinity.py). At this step, we combined assemblies from previously published sequences (including outgroups) with these newly generated assemblies and aligned these to enrichment baits using phyluce_assembly_match_contigs_to_probes.py (min_coverage=50, min_identity=80), thereby creating a relational sqlite database containing the matched probes. Sequence quality statistics were calculated for Trinity contigs and UCE contigs using phyluce_assembly_get_fastq_lengths and are summarized in Supplementary Data  1 . We aligned the sequence data for individual UCE loci using MAFFT v7.130b 103 through phyluce_assembly_seqcap_align.py (settings: max_divergence=0.2, min-length=100, -no-trim). We performed internal trimming using Gblocks v0.91b 104 and a phyluce wrapper script (phyluce_assembly_get_gblocks_trimmed _alignment_from_untrimmed.py), with the relaxed trimming settings b1 = 0.5, b2 = 0.5, b3 = 12, b4 = 7. From 2590 trimmed UCE alignments, we prepared three different matrices using sets of loci recovered for at least 50%, 60%, and 70% of taxa for further analyses using the script phyluce_align_get_only_loci_with_min_taxa.py. 1118, 767 and 447 UCE loci were retained for analysis in a 50%, 60%, and 70% matrix, respectively. We have outlined the above deviations from default parameters for UCE processing only; for detailed documentation of this bioinformatics pipeline refer to https://phyluce.readthedocs.io/en/latest/tutorials/ . Supplementary Fig.  1 provides a flowchart-style overview of our data treatments and data sets.

To extract protein-coding loci from our captured loci, we followed a published pipeline 105 and used the required script available at https://github.com/marekborowiec/uce-to-protein , which uses BLASTX ( https://blast.ncbi.nlm.nih.gov/Blast.cgi ) to match unaligned UCE sequences to a reference protein database. Filtering of the protein-coding UCE loci retained 324 loci present in at least 50% of taxa, which were used for subsequent phylogenetic analyses.

Phylogenetic inference

We partitioned the nucleotide data matrices using the Sliding-Window Site Characteristics Entropy (SWSC-EN) algorithm 106 and PartitionFinder2 v2.1.1 107 employing the r cluster algorithm 108 . The resulting partitioned nucleotide data matrices (nuc-50%-SWSC, nuc-60%-SWSC, and nuc-70%-SWSC), as well as unpartitioned versions (nuc-50%-unpart, nuc-60%-unpart, and nuc-70%-unpart) were analyzed with Maximum Likelihood (ML) best-tree and ultrafast bootstrap searches in IQ-TREE v1.6.10 109 employing model selection for unpartitioned matrices while implementing a GTR + G model for data subsets in partitioned matrices. The 324 protein-coding loci were analyzed both as nucleotide matrix (prot-nuc-unpart) and translated to amino acids (prot-AA-unpart), employing model selection in IQ-TREE but no data partitioning. We kept third codon positions in the nucleotide data set to ensure comparability with the full data sets. All analyses were rooted using the outer, non-holometabolous outgroup (Blattodea: Blattella germanica ). We calculated several alignment statistics (e.g., alignment length, amount of missing data, number of parsimony-informative sites) with AMAS 110 , summarized in Supplementary Data  2 .

We used the Four-cluster Likelihood Mapping (FcLM) approach 44 to test four topological hypotheses regarding the position of Ichneumonoidea and Ceraphronoidea on each of our four main data sets (nuc-50%, nuc-60%, nuc-70% and prot-AA). The four hypotheses were investigated by defining four taxon groups (specified in Supplementary Data  14 and Supplementary Methods), and FcLM analyses were performed in IQ-TREE v1.6.12 using 100,000 randomly drawn quartets. Further information on additional phylogenetic sensitivity tests and exploratory analyses, such as GC content analyses, can be found in the Supplementary Methods. Coalescence-based phylogenetic inference was not pursued after the preliminary analysis stage because some characteristics of our data set (short alignment size and high levels of missing data for individual loci) suggested a high propensity for gene estimation errors.

Divergence dating

Divergence times were estimated using the information on twelve fossils within Hymenoptera, chosen following best practices for fossil calibrations 111 , and representing the oldest and most reliable available calibration points for superfamily and family-level nodes, except for one fossil calibrating a subfamily-level node. We restricted our calibrations to these 12 fossils as they covered all major lineages and deep divergences within Hymenoptera for which confidently placed fossils were available. Such deep calibrations have been shown to increase the accuracy of divergence estimates 112 ; calibrations at more shallow nodes were unlikely to improve the age estimates, yet increase the number of parameters in the computationally challenging analyses. Supplementary Data  15 detail the implemented calibrations, the characters used for their placement, and corresponding references. In addition to the fossil calibrations, we set a soft maximum bound of 283.7 Ma for crown Hymenoptera, which represents the upper 95% CI for the age of the order estimated by Misof et al. 62 (see Table  S25 in that paper). We employed approximate likelihood to estimate divergence times in mcmctree and codeml as included in PAMLv4.9 113 , using the two data sets and trees that were best-supported (by ufBS and FcLM) and most frequently recovered across our phylogenetic analyses (see Supplementary Methods and Discussion for further details): 1) the nuc-50% matrix and the best ML tree resulting from SWSC-EN partitioning of this matrix (topA-0), and 2) the nuc-70% matrix and the best ML tree resulting from SWSC-EN partitioning of this matrix (topC-1). Outgroups were pruned from the tree and alignment prior to divergence time estimation. All fossil calibrations were implemented as soft minima, except for the calibration on the root node on which we placed soft minimum and maximum bounds, and using default settings (heavy-tailed density based on a truncated Cauchy distribution with an offset p  = 0.1, a scale parameter c = 1, and a left tail probability of α=0.025 creating the soft bound). We set samplefreq=10 and n samples = 2,000,000, resulting in a potential chain length of 20,000,000 states (sample freq ×  n samples). We set up four separate runs for each data set and periodically checked progress and convergence parameters by visualizing mcmc convergence and effective sample sizes (ESS) using TracerV1.7.1 114 . Runs were stopped at 1,732,010–4,632,960 states once most parameters reached ESS values above 200 (excluding burnin). Most parameters well exceeded the ESS threshold in individual runs, but due to the large number of parameters to estimate (>750), a few only reached the threshold after combining results from the four runs. 731,251 samples for topC-1 and 1,180,127 for topA-0 were summarized across four runs each, after discarding 25–50% of samples as burnin. To evaluate the impact of our calibrations, we also performed analyses without sequence data using only the prior.

Diversification and comparative analyses

To assess potential shifts in diversification rates over time in Hymenoptera we used a sampling fraction approach in BAMM v2.5 45 , 46 and the associated R package BAMMtools v2.1.7 115 , as well as a taxonomic approach implemented in the MEDUSA function in the R package Geiger v2.0.7 116 in R v4.0.3. For BAMM, we created clade-specific sampling probabilities by assembling a richness matrix with the number of described species for all families of Hymenoptera included in our analyses. We predominantly used species estimates published in Huber 18 , except for the following groups for which the classification in that volume was outdated or did not correspond with natural monophyletic groupings. Within Aculeata we used the family-level classifications established by Pilgrim et al. 80 and Sann et al. 81 and species diversity estimates from Branstetter et al. 117 and Pulawski’s catalog 118 . Within Cynipoidea, we distinguished six monophyletic clades as identified in Blaimer et al. 87 and assigned species richness based on Buffington et al. 119 . Due to several non-monophyletic families in the Chalcidoidea (e.g., Pteromalidae) and the associated uncertainty about lineage-specific species richness, we treated this entire superfamily as one clade for the purpose of this analysis. Platygastroidea were treated similarly; the recently updated taxonomy of this group 79 could not yet be incorporated at the time of analysis. This merging and splitting of families into recognized monophyletic lineages resulted in 68 clades defined for analyses (with 13 missing). Clade-specific sampling probabilities were then calculated as proportions of sampled diversity divided by the total described species diversity in these clades and are listed in Supplementary Data  16 . We also applied a backbone sampling fraction (68 sampled/81 recognized clades = 0.8395) to account for the unsampled clades in the analyses. Sampling and parameter choice are further discussed in the Supplementary Discussion. We used the two chronograms (topA-0 and topC-1) generated in the dating analyses for two sets of BAMM analyses. Our analyses were configured using the function “setBAMMpriors” within BAMMtools to obtain appropriate priors for speciation-extinction analyses as outlined in the guidelines in the BAMM documentation ( http://bamm-project.org/ ); see Supplementary Methods for details on priors used. Our runs included four mcmc chains with a length of 200 million generations, sampling every 10,000 generations, and discarding a burnin of 10%. We confirmed that ESS values were appropriate (>200) and used “computeBayesFactors” to identify the best-supported model of rate shifts in our data. Results were analyzed and plotted with various functions in BAMMtools to infer mean phylorate plots (plot.bammdata), best shift configurations (getBestShiftConfiguration), credible shift sets (credibleShiftSet), and cumulative shift probability trees (cumulativeShiftProbsTree). Cumulative shift probabilities were displayed within a range of 0.97–1.00, as less stringent cutoff values support shifts leading to almost every major clade. We calculated mean speciation and extinction rates for specific clades for which rate shifts were indicated (both by BAMM and MEDUSA, see below).

A taxonomic method, in which clades are simply collapsed to terminal lineages of equal rank, may be more appropriate for incomplete sampling in diversification analyses 120 and circumvent some of the problems raised in the debates about BAMM 121 , 122 . The MEDUSA algorithm first fits a single diversification model (the background rate) to the entire data set, and then adds single breakpoints (i.e. shifts) in the diversification process in a step-wise fashion, so that different parts of the tree are allowed to evolve with different parameter values and have different rate regimes 47 . We implemented this analysis using clade-level chronograms for both topA-0 and topC-1, which we created by dropping all tips except one representative for each of the 68 clades also designated in BAMM analyses. The species richness matrix assembled for BAMM, composed of the described species diversity for each clade (Supplementary Data  16 ), was further designated to assign diversity estimates to the clade-level tree for this analysis. We used an AICc threshold of 3.760758 (computed automatically by MEDUSA) as a stopping criterion for the algorithm, at which further breakpoints in the diversification process are not added.

To investigate the evolution of parasitoidism in Hymenoptera, we used the same clade-level approach to integrate the entire character diversity within each group, thus also accounting for taxa missing from our phylogeny and avoiding bias by over- or underrepresentation of particular states. We inferred (1) the presence or absence of parasitoidism as a binary trait, and (2) the evolution of hymenopteran life strategies on a more detailed level, assigning the four categories parasitoidism, primary phytophagy (including xylophagy and mycophagy), secondary phytophagy (i.e. gall-inducing, pollen collecting), and predation. We also (3) contrasted carnivory (parasitoidism and predation combined) with phytophagy and secondary phytophagy in an analysis comprising three trait categories, though the results shown here focus on parasitoidism-centered analyses. Clades were assigned to one of these categories based on the life strategy exhibited by most (>80–95%) members. We allowed polymorphism for Chalcidoidea, Formicidae, and Vespidae, for which such a fully binary choice would not be representative of the group. Our rationale is further described in the Supplementary Discussion; see also Supplementary Data  16 for trait coding and the references used to score the biology of each group. We used the rayDISC function in the R package corHMM v2.1 in R v4.0.3 to reconstruct ancestral states using the reduced clade-level phylogenies created for MEDUSA analyses. We performed reconstructions under the “equal rates” model (ER) and the “all rates different” model (ARD) and compared the fit of these models with a likelihood ratio test.

We tested for state-dependent diversification associated with the four putative key innovations (wasp waist, stinger, parasitoidism, secondary phytophagy) and related traits (carnivory, phytophagy) in Hymenoptera using the HiSSE (Hidden State Speciation and Extinction) framework and associated R package 48 in R v4.0.3. The HiSSE framework has been developed to overcome some of the shortcomings of the SSE models (e.g., BiSSE 123 or MuSSE 124 ), which is a benefit of the more complex null models applied in this approach 48 , 125 , 126 . HiSSE models incorporate hidden states, which are unobserved traits that influence the diversification rate together with the observed traits 48 . In a HiSSE model where the focal states are, for example, 1 = parasitoidism present and 0 = parasitoidism absent, the diversification parameters of the hidden character will be modeled as a second character with states A and B. We decided on this approach rather than employing the Multistate Hidden Speciation and Extinction model (MuHiSSE 127 ), for example, as we specifically wanted to evaluate support for each trait as a key innovation independently. This framework addresses concerns about the false positive inference of state-dependent diversification when applying less complex models such as BiSSE 126 , 128 . We compared 30 models in the HiSSE framework, using the full set of models tested by Beaulieu and O’Meara 48 plus six models suggested in the HiSSE documentation and a study assessing the diversification of Squamates 129 . We tested the fit of our data to a full HiSSE model with unconstrained parameters and 17 subsets with various constraints on transition and diversification rates. Four models excluded hidden states and modeled trait-dependent diversification in a BiSSE-like fashion. Eight character-independent null models, or CID-2 and CID-4 models (sensu Beaulieu & O’Meara 48 ), were further tested, also including an extension of the currently implemented CID-4 models in the HiSSE package allowing for nine transitions rates 129 . We performed all analyses using the full phylogenies for topA-0 and topC-1 (same species-level trees as used for BAMM) but assigned character states based on the clade level. Calculating the occurrence of these traits across our sampling of Hymenoptera, we employed sampling fractions for character states as proportions of sampled vs. unsampled trait occurrences; these are specified in Supplementary Data  17 and the Supplementary Methods. Specifications of models and parameters are given in Supplementary Data  10 . Since the HiSSE function does not allow for polymorphic states or missing data, we resolved all polymorphisms to a present state, and taxa with uncertain states were coded with the state present if that was the case for their closest relatives. This follows the logic that a state can generally be counted as present in a clade even if not displayed in every taxon, while closely related taxa have a higher probability of sharing the same traits. The diversification rate and trait reconstruction results were plotted and summarized for the best-scoring models.

We also employed a recent direct extension of the HiSSE framework in the same R package (but using v2.1.9 in R v4.2.2), the MiSSE (Missing State Speciation and Extinction) model. MiSSE executes a trait-free version of the HiSSE model and thereby focuses only on the impact of unobserved, hidden states on the diversification dynamics of a clade 130 . We automated the process of fitting MiSSE models by using the “MiSSEGreedy” function and possible.combos=generateMiSSEGreedyCombinations(), using the default for stop.deltaAICc=10 and a sampling fraction of 0.005 based on our sampling of Hymenoptera (765 of 152,691 described species). We performed analyses using both topA-0 and C-1 topologies and summarized and plotted tip rates for the best-scoring model across both phylogenies.

Reporting summary

Further information on research design is available in the  Nature Portfolio Reporting Summary linked to this article.

Data availability

The raw sequence reads newly generated in this study have been deposited in the NCBI Sequence Read Archive under BioProject accession code PRJNA811764 and PRJNA632862 . The UCE sequence data from prior publications used in this study are available under BioProject accession codes PRJNA379583 , PRJNA248919 , PRJNA495844 , PRJNA814466 , PRJNA606284 , PRJNA647791 , PRJNA625490 , and PRJNA473845 ; the genome assemblies used in this study are available under accession codes GCA_001444175 , GCA_000699045 , GCA_000762945 , GCA_000696855 , GCA_000006295 , and GCA_000931545 . The raw sequence data can also be accessed using individual accession numbers given in Supplementary Data  1 . Source data for this study (assembled contig files, data matrices, tree and log files for phylogenetic analyses; input and output files for FcLM; R code and input and results files for comparative analyses) are available in the Dryad repository at https://doi.org/10.5061/dryad.08kprr54m 131 . Information on the location of voucher specimens is provided in Supplementary Data  1 .

Code availability

No new custom code is published with this article. We direct the reader to the Dryad repository accompanying this article ( https://doi.org/10.5061/dryad.08kprr54m ) which includes details for all code implemented in our analyses.

Heard, S. B. & Hauser, D. L. Key evolutionary innovations and their ecological mechanisms. Hist. Biol. 10 , 151–173 (1995).

Article   Google Scholar  

Hunter, J. P. Key innovations and the ecology of macroevolution. Trends Ecol. Evol. 13 , 31–36 (1998).

Article   CAS   PubMed   Google Scholar  

Schweizer, M., Seehausen, O. & Hertwig, S. T. Macroevolutionary patterns in the diversification of parrots: effects of climate change, geological events and key innovations. J. Biogeogr. 38 , 2176–2194 (2011).

Salzburger, W., Mack, T., Verheyen, E. & Meyer, A. Out of Tanganyika: genesis, explosive speciation, key-innovations and phylogeography of the haplochromine cichlid fishes. BMC Evol. Biol. 5 , 17 (2005).

Article   PubMed   PubMed Central   Google Scholar  

Chouvenc, T., Šobotník, J., Engel, M. S. & Bourguignon, T. Termite evolution: mutualistic associations, key innovations, and the rise of Termitidae. Cell. Mol. Life Sci. 78 , 2749–2769 (2021).

Nicholson, D. B., Ross, A. J. & Mayhew, P. J. Fossil evidence for key innovations in the evolution of insect diversity. Proc. R. Soc. B Biol. Sci. 281 , 20141823 (2014).

Rabosky, D. L. Phylogenetic tests for evolutionary innovation: the problematic link between key innovations and exceptional diversification. Philos. Trans. R. Soc. Lond. B Biol. Sci. 372 , 20160417 (2017).

Rainford, J. L., Hofreiter, M., Nicholson, D. B. & Mayhew, P. J. Phylogenetic distribution of extant richness suggests metamorphosis is a key innovation driving diversification in insects. PLoS ONE 9 , e109085 (2014).

Article   ADS   PubMed   PubMed Central   Google Scholar  

Donoghue, M. J. Key innovations, convergence, and success: macroevolutionary lessons from plant phylogeny. Paleobiology 31 , 77–93 (2005).

Donoghue, M. J. & Sanderson, M. J. Confluence, synnovation, and depauperons in plant diversification. N. Phytol. 207 , 260–274 (2015).

Bouchenak‐Khelladi, Y., Onstein, R. E., Xing, Y., Schwery, O. & Linder, H. P. On the complexity of triggering evolutionary radiations. N. Phytol. 207 , 313–326 (2015).

Mayhew, P. J. Why are there so many insect species? Perspectives from fossils and phylogenies. Biol. Rev. 82 , 425–454 (2007).

Article   PubMed   Google Scholar  

Carpenter, F. The geological history and evolution of insects. Am. Sci. 41 , 256–270 (1953).

Google Scholar  

Grimaldi, D. & Engel M.S. The Evolution of the Insects (Cambridge University Press, Cambridge, 2005).

Wiens, J. J., Lapoint, R. T. & Whiteman, N. K. Herbivory increases diversification across insect clades. Nat. Commun. 6 , 8370 (2015).

Article   ADS   CAS   PubMed   Google Scholar  

Mitter, C., Farrell, B. & Wiegmann, B. The phylogenetic study of adaptive zones: has phytophagy promoted insect diversification? Am. Sci. 132 , 107–128 (1988).

Forbes, A. A., Bagley, R. K., Beer, M. A., Hippee, A. C. & Widmayer, H. A. Quantifying the unquantifiable: why Hymenoptera, not Coleoptera, is the most speciose animal order. BMC Ecol. 18 , 21 (2018).

Huber, J.T. Biodiversity of Hymenoptera. In: Insect Biodiversity: Science and Society (Wiley-Blackwell, Oxford, 2017).

Huber, J. & Noyes, J. A new genus and species of fairyfly, Tinkerbella nana (Hymenoptera, Mymaridae), with comments on its sister genus Kikiki , and discussion on small size limits in arthropods. J. Hymenopt. Res. 32 , 17 (2013).

Nénon, J.-P., Kacem, N. & Le Lannic, J. Structure, sensory equipment, and secretions of the ovipositor in a giant species of Hymenoptera: Megarhyssa atrata F.(Ichneumonidae, Pimplinae). Can. Entomol. 129 , 789–799 (1997).

Godfray, H. Parasitoids: Behavioral and Evolutionary Ecology (Princeton University Press, 1994).

Mockford, E. L. A new species of Dicopomorpha (Hymenoptera: Mymaridae) with diminutive, apterous males. Ann. Entomol. Soc. 90 , 115–120 (1997).

Pennacchio, F. & Strand, M. R. Evolution of developmental strategies in parasitic Hymenoptera. Annu. Rev. Entomol. 51 , 233–258 (2006).

Eggleton, P. & Belshaw, R. Insect parasitoids: an evolutionary overview. Philos. Trans. R. Soc. Lond. B Biol. Sci. 337 , 1–20 (1992).

Article   ADS   Google Scholar  

Heraty, J. Parasitoid biodiversity and insect pest management. Insect Biodiversity: Science and Society (Wiley-Blackwell, Oxford, 2017).

Whitfield, J. B. Phylogenetic insights into the evolution of parasitism in Hymenoptera. Adv. Parasitol. 54 , 69–101 (2003).

Vilhelmsen, L. Before the wasp-waist: comparative anatomy and phylogenetic implications of the skeleto-musculature of the thoraco-abdominal boundary region in basal Hymenoptera (Insecta). Zoomorphology 119 , 185–221 (2000).

Vilhelmsen, L. Phylogeny and classification of the extant basal lineages of the Hymenoptera (Insecta). Zool. J. Linn. Soc. 131 , 393–442 (2001).

Vilhelmsen, L., Miko, I. & Krogmann, L. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zool. J. Linn. Soc. 159 , 22–194 (2010).

Brothers, D. Aculeate Hymenoptera: phylogeny and classification. In: Encyclopedia of Social Insects (Springer Nature, Switzerland, 2019).

Murray, E. A., Bossert, S. & Danforth, B. N. Pollinivory and the diversification dynamics of bees. Biol. Lett. 14 , 20180530 (2018).

Kristensen, N. P. Phylogeny of endopteryygote insects, the most successful lineage of living organisms. Eur. J. Entomol. 96 , 237–254 (1999).

Heraty, J. et al. Evolution of the hymenopteran megaradiation. Mol. Phylogenet. Evol. 60 , 73–88 (2011).

Sharkey, M. J. et al. Phylogenetic relationships among superfamilies of Hymenoptera. Cladistics 28 , 80–112 (2012).

Klopfstein, S., Vilhelmsen, L., Heraty, J. M., Sharkey, M. & Ronquist, F. The hymenopteran tree of life: evidence from protein-coding genes and objectively aligned ribosomal data. PLoS ONE 8 , e69344 (2013).

Article   ADS   CAS   PubMed   PubMed Central   Google Scholar  

Peters, R. S. et al. Evolutionary history of the Hymenoptera. Curr. Biol. 27 , 1013–1018 (2017).

Peters, R. S. et al. The taming of an impossible child: a standardized all-in approach to the phylogeny of Hymenoptera using public database sequences. BMC Biol. 9 , 1–14 (2011).

Branstetter, M. G. et al. Phylogenomic insights into the evolution of stinging wasps and the origins of ants and bees. Curr. Biol. 27 , 1019–1025 (2017).

Ronquist, F. et al. A total-evidence approach to dating with fossils, applied to the early radiation of the Hymenoptera. Syst. Biol. 61 , 973–999 (2012).

Chesters, D. The phylogeny of insects in the data‐driven era. Syst. Entomol. 45 , 540–551 (2020).

Baca, S. M., Alexander, A., Gustafson, G. T. & Short, A. E. Ultraconserved elements show utility in phylogenetic inference of Adephaga (Coleoptera) and suggest paraphyly of ‘Hydradephaga’. Syst. Entomol. 42 , 786–795 (2017).

Gustafson, G. T., Baca, S. M., Alexander, A. M. & Short, A. E. Z. Phylogenomic analysis of the beetle suborder Adephaga with comparison of tailored and generalized ultraconserved element probe performance. Syst. Entomol. 45 , 552–570 (2019).

Rasnitsyn, A. An outline of evolution of the hymenopterous insects (order Vespida). Orient. Insects 22 , 115–145 (1988).

Strimmer, K. & Von Haeseler, A. Likelihood-mapping: a simple method to visualize phylogenetic content of a sequence alignment. Proc. Natl Acad. Sci. USA 94 , 6815–6819 (1997).

Article   ADS   CAS   PubMed   PubMed Central   MATH   Google Scholar  

Rabosky, D. L. et al. Rates of speciation and morphological evolution are correlated across the largest vertebrate radiation. Nat. Commun. 4 , 1–8 (2013).

Rabosky, D. L. Automatic detection of key innovations, rate shifts, and diversity-dependence on phylogenetic trees. PLoS ONE 9 , e89543 (2014).

Alfaro, M. E. et al. Nine exceptional radiations plus high turnover explain species diversity in jawed vertebrates. Proc. Natl Acad. Sci. USA 106 , 13410–13414 (2009).

Beaulieu, J. M. & O’Meara, B. C. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65 , 583–601 (2016).

Dowton, M. & Austin, A. D. Simultaneous analysis of 16S, 28S, COI and morphology in the Hymenoptera: Apocrita—evolutionary transitions among parasitic wasps. Zool. J. Linn. Soc. 74 , 87–111 (2001).

Drummond, A. J., Suchard, M. A., Xie, D. & Rambaut, A. Bayesian phylogenetics with BEAUti and the BEAST 1.7. Mol. Biol. Evol. 29 , 1969–1973 (2012).

Article   CAS   PubMed   PubMed Central   Google Scholar  

Ronquist, F. et al. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst. Biol. 61 , 539–542 (2012).

Weiblen, G. D. & Bush, G. L. Speciation in fig pollinators and parasites. Mol. Ecol. 11 , 1573–1578 (2002).

Cruaud, A. et al. An extreme case of plant–insect codiversification: figs and fig-pollinating wasps. Syst. Biol. 61 , 1029–1047 (2012).

Cardinal, S. & Danforth, B. N. Bees diversified in the age of eudicots. Proc. R. Soc. B. 280 , 20122686 (2013).

Michez, D., Vanderplanck M. & Engel M.S. Fossil bees and their plant associates. In: Evolution of Plant-Pollinator Relationships (Cambridge University Press, Cambridge, 2012).

Soltis, D. et al. The age and diversification of early angiosperms: Integration of the fossil record and molecular dates. In: Phylogeny and evolution of the angiosperms: revised and updated edition (University of Chicago Press, Chicago, 2018).

Benton, M. J., Wilf, P. & Sauquet, H. The Angiosperm Terrestrial Revolution and the origins of modern biodiversity. N. Phytol. 233 , 2017–2035 (2022).

Isaka, Y. & Sato, T. Was species diversification in Tenthredinoidea (Hymenoptera: Symphyta) related to the origin and diversification of angiosperms? Can. Entomol. 147 , 443–458 (2015).

Labandeira, C.C. & Li L. The history of insect parasitism and the Mid-Mesozoic Parasitoid Revolution. In: The Evolution and Fossil Record of Parasitism: Identification and Macroevolution of Parasites (Springer, New York, 2021).

Eggleton, P. & Belshaw, R. Comparisons of dipteran, hymenopteran and coleopteran parasitoids: provisional phylogenetic explanations. Biol. J. Linn. Soc. 48 , 213–226 (1993).

Wiegmann, B. M. et al. Episodic radiations in the fly tree of life. Proc. Natl Acad. Sci. USA 108 , 5690–5695 (2011).

Misof, B. et al. Phylogenomics resolves the timing and pattern of insect evolution. Science 346 , 763–767 (2014).

Kumpanenko, A., Gladun, D. & Vilhelmsen, L. Functional morphology and evolution of the sting sheaths in Aculeata (Hymenoptera). Arthropod Syst. Phylogeny 77 , 325–338 (2019).

Yamada, Y. Charactiristics of the Oviposition of a Parasitoid, Chrysis shanghaiensis : Hymenoptera: Chrysididae. Appl. Entomol. Zool. 22 , 456–464 (1987).

Oeser, R. Vergleichend-morphologische Untersuchungen über den Ovipositor der Hymenopteren. Mitt. Museum Naturkunde Berl. Zool. Museum Inst. Spez. Zool. (Berl.) 37 , 3–119 (1961).

Wiegmann, B. M., Mitter, C. & Farrell, B. Diversification of carnivorous parasitic insects: extraordinary radiation or specialized dead end? Am. Nat. 142 , 737–754 (1993).

McKenna, D. D. et al. The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Syst. Ent. 40 , 835–880 (2015).

Zhang, S.-Q. et al. Evolutionary history of Coleoptera revealed by extensive sampling of genes and species. Nat. Commun. 9 , 1–11 (2018).

ADS   Google Scholar  

Kawahara, A. Y. et al. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proc. Natl Acad. Sci. USA 116 , 22657–22663 (2019).

Spriggs, E. L., Christin, P.-A. & Edwards, E. J. C4 Photosynthesis promoted species diversification during the Miocene grassland expansion. PLoS ONE 9 , e97722 (2014).

Dötterl, S. & Vereecken, N. The chemical ecology and evolution of bee–flower interactions: a review and perspectives. Can. J. Zool. 88 , 668–697 (2010).

Farrell, B. D. “Inordinate fondness” explained: why are there so many beetles? Science 281 , 555–559 (1998).

Futuyma, D. J. & Agrawal, A. A. Macroevolution and the biological diversity of plants and herbivores. Proc. Natl Acad. Sci. USA 106 , 18054–18061 (2009).

Malagón-Aldana, L. A., Jensen, A. R., Smith, D. R., Shinohara, A. & Vilhelmsen, L. Sawflies out of Gondwana: phylogenetics and biogeography of Argidae (Hymenoptera). Syst. Ent. 47 , 231–258 (2021).

Maddison, W. P. & FitzJohn, R. G. The unsolved challenge to phylogenetic correlation tests for categorical characters. Syst. Biol. 64 , 127–136 (2015).

Rabosky, D. L. Challenges in the estimation of extinction from molecular phylogenies: a response to Beaulieu and O’Meara. Evolution 70 , 218–228 (2016).

Louca, S. & Pennell, M. W. Extant timetrees are consistent with a myriad of diversification histories. Nature 580 , 502–505 (2020).

Craig, J. M., Kumar, S. & Hedges, S. B. Limitations of phylogenomic data can drive inferred speciation rate shifts. Mol. Biol. Evol. 39 , msac038 (2022).

Chen, H. et al. An integrated phylogenetic reassessment of the parasitoid superfamily Platygastroidea (Hymenoptera: Proctotrupomorpha) results in a revised familial classification. Syst. Entomol. 46 , 1088–1113 (2021).

Pilgrim, E. M., von Dohlen, C. D. & Pitts, J. P. Molecular phylogenetics of Vespoidea indicate paraphyly of the superfamily and novel relationships of its component families and subfamilies. Zool. Scr. 37 , 539–560 (2008).

Sann, M. et al. Phylogenomic analysis of Apoidea sheds new light on the sister group of bees. BMC Evol. Biol. 18 , 71 (2018).

Faircloth, B. C., Branstetter, M. G., White, N. D. & Brady, S. G. Target enrichment of ultraconserved elements from arthropods provides a genomic perspective on relationships among Hymenoptera. Mol. Ecol. Res. 15 , 489–501 (2015).

Article   CAS   Google Scholar  

Blaimer, B. B., Ward, P. S., Schultz, T. R., Fisher, B. L. & Brady, S. G. Paleotropical diversification dominates the evolution of the hyperdiverse ant tribe Crematogastrini (Hymenoptera: Formicidae). Insect Syst. Div. 2 , 3 (2018).

Cruaud, A. et al. Optimized DNA extraction and library preparation for minute arthropods: application to target enrichment in chalcid wasps used for biocontrol. Methods Ecol. Evol. 19 , 702–710 (2019).

CAS   Google Scholar  

Cruaud, A. et al. Ultra-Conserved Elements and morphology reciprocally illuminate conflicting phylogenetic hypotheses in Chalcididae (Hymenoptera, Chalcidoidea). Cladistics 37 , 1–35 (2021).

Rasplus, J.-Y. et al. A first phylogenomic hypothesis for Eulophidae (Hymenoptera, Chalcidoidea). J. Nat. Hist. 54 , 597–609 (2020).

Blaimer, B. B., Gotzek, D., Brady, S. G. & Buffington, M. L. Comprehensive phylogenomic analyses re-write the evolution of parasitism within cynipoid wasps. BMC Evol. Biol. 20 , 155 (2020).

Santos, B. F., Perrard, A. & Brady, S. G. Running in circles in phylomorphospace: host environment constrains morphological diversification in parasitic wasps. Proc. R. Soc. B Biol. Sci. 286 , 20182352 (2019).

Santos, B. F. et al. Phylogenomics of Ichneumoninae (Hymenoptera, Ichneumonidae) reveals pervasive morphological convergence and the shortcomings of previous classifications. Syst. Ent. 46 , 704–724 (2021).

Supeleto, F. A., Santos, B. F., Brady, S. G. & Aguiar, A. P. Phylogenomic analyses reveal a rare new genus of wasp (Hymenoptera, Ichneumonidae, Cryptinae) from the Brazilian Atlantic Forest. Syst. Biodivers. 18 , 646–661 (2020).

Jasso-Martinez, J. M. et al. Phylogenomics of braconid wasps (Hymenoptera, Braconidae) sheds light on classification and the evolution of parasitoid life history traits. Mol. Phylogen. Evol. 173 , 107452 (2022).

Longino, J. T. & Branstetter, M. G. The truncated bell: an enigmatic but pervasive elevational diversity pattern in Middle American ants. Ecography 42 , 272–283 (2019).

Blaimer, B. B., LaPolla, J. S., Branstetter, M. G., Lloyd, M. W. & Brady, S. G. Phylogenomics, biogeography and diversification of obligate mealybug-tending ants in the genus Acropyga . Mol. Phylogen. Evol. 102 , 20–29 (2016).

Blaimer, B. B., Lloyd, M. W., Guillory, W. X. & Brady, S. G. Sequence capture and phylogenetic utility of genomic ultraconserved elements obtained from pinned insect specimens. PLoS ONE 11 , e0161531 (2016).

Fisher, S. et al. A scalable, fully automated process for construction of sequence-ready human exome targeted capture libraries. Genome Biol. 12 , R1 (2011).

Rohland, N. & Reich, D. Cost-effective, high-throughput DNA sequencing libraries for multiplexed target capture. Genome Res. 22 , 939–946 (2012).

Faircloth, B. C. & Glenn, T. C. Not all sequence tags are created equal: designing and validating sequence identification tags robust to indels. PLoS ONE 7 , e42543 (2012).

Branstetter, M. G., Longino, J. T., Ward, P. S. & Faircloth, B. C. Enriching the ant tree of life: enhanced UCE bait set for genome-scale phylogenetics of ants and other Hymenoptera. Methods Ecol. Evol. 8 , 768–776 (2017).

Faircloth, B. C. PHYLUCE is a software package for the analysis of conserved genomic loci. Bioinformatics 32 , 786–788 (2016).

Faircloth, B. Illumiprocessor: a trimmomatic wrapper for parallel adapter and quality trimming. https://doi.org/10.6079/J6079ILL (2013).

Bolger, A. M., Lohse, M. & Usadel, B. Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics 30 , 2114–2120 (2014).

Grabherr, M. G. et al. Full-length transcriptome assembly from RNA-Seq data without a reference genome. Nat. Biotechnol. 29 , 644–652 (2011).

Katoh, K., Asimenos G. & Toh H. Multiple alignment of DNA sequences with MAFFT. In: Bioinformatics for DNA sequence analysis ). (Springer, 2009).

Castresana, J. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Mol. Biol. Evol. 17 , 540–552 (2000).

Borowiec, M. L. Convergent evolution of the army ant syndrome and congruence in big-data phylogenetics. Syst. Biol. 68 , 642–656 (2019).

Tagliacollo, V. A. & Lanfear, R. Estimating improved partitioning schemes for UltraConserved Elements (UCEs). Mol. Biol. Evol. 37 , 1798–1811 (2018).

Lanfear, R., Frandsen, P. B., Wright, A. M., Senfeld, T. & Calcott, B. PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Mol. Biol. Evol. 34 , 772–773 (2017).

CAS   PubMed   Google Scholar  

Lanfear, R., Calcott, B., Kainer, D., Mayer, C. & Stamatakis, A. Selecting optimal partitioning schemes for phylogenomic datasets. BMC Evol. Biol. 14 , 82 (2014).

Nguyen, L.-T., Schmidt, H. A., von Haeseler, A. & Minh, B. Q. IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Mol. Biol. Evol. 32 , 268–274 (2015).

Borowiec, M. L. AMAS: a fast tool for alignment manipulation and computing of summary statistics. PeerJ 4 , e1660 (2016).

Parham, J. F. et al. Best practices for justifying fossil calibrations. Syst. Biol. 61 , 346–359 (2011).

Mello, B. & Schrago, C. G. Assignment of calibration information to deeper phylogenetic nodes is more effective in obtaining precise and accurate divergence time estimates. Evol. Bioinform. Online 10 , 79–85 (2014).

Yang, Z. PAML 4: Phylogenetic analysis by maximum likelihood. Mol. Biol. Evol. 24 , 1586–1591 (2007).

Rambaut, A., Drummond, A. J., Xie, D., Baele, G. & Suchard, M. A. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Syst. Biol. 67 , 901–904 (2018).

Rabosky, D. L. et al. BAMMtools: an R package for the analysis of evolutionary dynamics on phylogenetic trees. Methods Ecol. Evol. 5 , 701–707 (2014).

Pennell, M. W. et al. geiger v2.0: an expanded suite of methods for fitting macroevolutionary models to phylogenetic trees. Bioinformatics 30 , 216–2218 (2014).

Branstetter, M. G. et al. Genomes of the Hymenoptera. Curr. Opin. Insect Sci. 25 , 65–75 (2018).

Pulawski, W. Catalogue of Sphecidae sensu lato. https://www.calacademy.org/scientists/projects/catalog-of-sphecidae . Accessed February, 2021.

Buffington, M. L., Forshage, M., Liljeblad, J., Tang, C.-T. & van Noort, S. World Cynipoidea (Hymenoptera): a key to higher-level groups. Insect Syst. Div. 4 , 1 (2020).

Chang, J., Rabosky, D. L. & Alfaro, M. E. Estimating diversification rates on incompletely sampled phylogenies: theoretical concerns and practical solutions. Syst. Biol. 69 , 602–611 (2019).

Moore, B. R., Höhna, S., May, M. R., Rannala, B. & Huelsenbeck, J. P. Critically evaluating the theory and performance of Bayesian analysis of macroevolutionary mixtures. Proc. Natl Acad. Sci. USA 113 , 9569–9574 (2016).

Meyer, A. L. S. & Wiens, J. J. Estimating diversification rates for higher taxa: BAMM can give problematic estimates of rates and rate shifts. Evolution 72 , 39–53 (2018).

Maddison, W. P., Midford, P. E. & Otto, S. P. Estimating a binary character’s effect on speciation and extinction. Syst. Biol. 56 , 701–710 (2007).

FitzJohn, R. G. Diversitree: comparative phylogenetic analyses of diversification in R. Methods Ecol. Evol. 3 , 1084–1092 (2012).

O’Meara, B. C. & Beaulieu, J. M. Past, future, and present of state-dependent models of diversification. Am. J. Bot. 103 , 792–795 (2016).

Uyeda, J. C., Zenil-Ferguson, R. & Pennell, M. W. Rethinking phylogenetic comparative methods. Syst. Biol. 67 , 1091–1109 (2018).

Nakov, T., Beaulieu, J. M. & Alverson, A. J. Diatoms diversify and turn over faster in freshwater than marine environments. Evolution 73 , 2497–2511 (2019).

Rabosky, D. L. & Goldberg, E. E. Model inadequacy and mistaken inferences of trait-dependent speciation. Syst. Biol. 64 , 340–355 (2015).

Harrington, S. & Reeder, T. W. Rate heterogeneity across Squamata, misleading ancestral state reconstruction, and the importance of proper null model specification. J. Evol. Biol. 30 , 313–325 (2017).

Vasconcelos, T., O’Meara, B. C. & Beaulieu, J. M. A flexible method for estimating tip diversification rates across a range of speciation and extinction scenarios. Evolution 76 , 1420–1433 (2022).

Blaimer, B. B. et al. Data from: Key innovations and the diversification of Hymenoptera, Dryad, Dataset. https://doi.org/10.5061/dryad.08kprr54m (2023).

Download references

Acknowledgements

Laboratory work was conducted in and with the support of the L.A.B. facilities of the National Museum of Natural History. We acknowledge general support for molecular work from L.A.B. staff and thank Michael Lloyd for assistance in laboratory methods. Many thanks to Taina Litwak for the illustrations and Matt Bertone for the images of wasps. Phylogenomic analyses utilized the Smithsonian Institution High-Performance Cluster (SI/HPC). Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the USDA. USDA is an equal opportunity provider and employer. This study was supported by the National Science Foundation (DEB-1555905, S.G.B.) and a Smithsonian Institute for Biodiversity Genomics and Global Genome Initiative grant (S.G.B., M.L.B., and B.B.B.), as well as a grant from the GGI Peer-Review Awards Program (B.F.S.). R.R.K., M.W.G., and M.L.B., as well as part of the sequencing effort, were funded by SEL-ARS. B.B.B. was supported by the Museum für Naturkunde, Berlin, during the analyses and writing stages of this project. J.Y.R. and AC were supported by the INRAe SPE department. B.F.S. was funded by a GGI Peter Buck Postdoctoral Fellowship (Smithsonian Institution) during much of this work. E.J.T. was supported by the Florida Department of Agriculture and Consumer Services, Division of Plant Industry.

Open Access funding enabled and organized by Projekt DEAL.

Author information

Authors and affiliations.

Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity Science, Center for Integrative Biodiversity Discovery, Invalidenstraße 43, Berlin, 10115, Germany

Bonnie B. Blaimer & Bernardo F. Santos

National Museum of Natural History, Smithsonian Institution, 10th & Constitution Ave. NW, Washington, DC, USA

Bonnie B. Blaimer, Bernardo F. Santos & Seán G. Brady

CBGP, INRAe, CIRAD, IRD, Montpellier SupAgro, Université de Montpellier, Montpellier, France

Astrid Cruaud & Jean-Yves Rasplus

Systematic Entomology Laboratory, USDA-ARS, c/o NMNH, Smithsonian Institution, 10th & Constitution Ave. NW, Washington, DC, USA

Michael W. Gates, Robert R. Kula, David R. Smith & Matthew L. Buffington

Department of Biological Sciences, University of New Hampshire, Durham, NH, USA

István Mikó

Florida State Collection of Arthropods, Division of Plant Industry, Florida Department of Agriculture and Consumer Services, 1911 SW 34th St, Gainesville, FL, 32608, USA

Elijah J. Talamas

You can also search for this author in PubMed   Google Scholar

Contributions

B.B.B., S.G.B., and M.L.B. conceived the study, with significant input from all other authors. R.R.K., B.F.S., M.W.G., M.L.B., E.J.T., I.M., D.R.S., J.Y.R., and A.C. provided samples. S.G.B., M.L.B., and M.W.G. provided funding or resources. B.B.B., B.F.S., J.Y.R., and A.C. generated ultraconserved element data. B.B.B., B.F.S., and A.C. processed data, and B.B.B. performed phylogenomic and macroevolutionary analyses. B.B.B. wrote the initial paper draft, and all other authors contributed to revising subsequent drafts of the paper and approved the final version.

Corresponding author

Correspondence to Bonnie B. Blaimer .

Ethics declarations

Competing interests.

The authors declare no competing interests.

Peer review

Peer review information.

Nature Communications thanks Heather Hines and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.  Peer reviewer reports are available.

Additional information

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Supplementary information

Supplementary information, peer review file, description of additional supplementary files, supplementary data 1-17, reporting summary, rights and permissions.

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/ .

Reprints and permissions

About this article

Cite this article.

Blaimer, B.B., Santos, B.F., Cruaud, A. et al. Key innovations and the diversification of Hymenoptera. Nat Commun 14 , 1212 (2023). https://doi.org/10.1038/s41467-023-36868-4

Download citation

Received : 25 March 2022

Accepted : 21 February 2023

Published : 03 March 2023

DOI : https://doi.org/10.1038/s41467-023-36868-4

Share this article

Anyone you share the following link with will be able to read this content:

Sorry, a shareable link is not currently available for this article.

Provided by the Springer Nature SharedIt content-sharing initiative

This article is cited by

• David Peris
• Fabien L. Condamine

Nature Communications (2024)

Terebra steering in chalcidoid wasps

• Benjamin Eggs
• Stefan Fischer
• Oliver Betz

Frontiers in Zoology (2023)

Elevated rates of dietary generalization in eusocial lineages of the secondarily herbivorous bees

BMC Ecology and Evolution (2023)

By submitting a comment you agree to abide by our Terms and Community Guidelines . If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.

Quick links

• Explore articles by subject
• Guide to authors
• Editorial policies

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Journal of Hymenoptera Research

Subject Area and Category

• Animal Science and Zoology
• Ecology, Evolution, Behavior and Systematics
• Insect Science

Pensoft Publishers

Publication type

10709428, 13142607

Information

How to publish in this journal

[email protected]

The set of journals have been ranked according to their SJR and divided into four equal groups, four quartiles. Q1 (green) comprises the quarter of the journals with the highest values, Q2 (yellow) the second highest values, Q3 (orange) the third highest values and Q4 (red) the lowest values.

The SJR is a size-independent prestige indicator that ranks journals by their 'average prestige per article'. It is based on the idea that 'all citations are not created equal'. SJR is a measure of scientific influence of journals that accounts for both the number of citations received by a journal and the importance or prestige of the journals where such citations come from It measures the scientific influence of the average article in a journal, it expresses how central to the global scientific discussion an average article of the journal is.

Evolution of the number of published documents. All types of documents are considered, including citable and non citable documents.

This indicator counts the number of citations received by documents from a journal and divides them by the total number of documents published in that journal. The chart shows the evolution of the average number of times documents published in a journal in the past two, three and four years have been cited in the current year. The two years line is equivalent to journal impact factor ™ (Thomson Reuters) metric.

Evolution of the total number of citations and journal's self-citations received by a journal's published documents during the three previous years. Journal Self-citation is defined as the number of citation from a journal citing article to articles published by the same journal.

Evolution of the number of total citation per document and external citation per document (i.e. journal self-citations removed) received by a journal's published documents during the three previous years. External citations are calculated by subtracting the number of self-citations from the total number of citations received by the journal’s documents.

International Collaboration accounts for the articles that have been produced by researchers from several countries. The chart shows the ratio of a journal's documents signed by researchers from more than one country; that is including more than one country address.

Not every article in a journal is considered primary research and therefore "citable", this chart shows the ratio of a journal's articles including substantial research (research articles, conference papers and reviews) in three year windows vs. those documents other than research articles, reviews and conference papers.

Ratio of a journal's items, grouped in three years windows, that have been cited at least once vs. those not cited during the following year.

Evolution of the percentage of female authors.

Evolution of the number of documents cited by public policy documents according to Overton database.

Evoution of the number of documents related to Sustainable Development Goals defined by United Nations. Available from 2018 onwards.

Leave a comment

Name * Required

Email (will not be published) * Required

* Required Cancel

The users of Scimago Journal & Country Rank have the possibility to dialogue through comments linked to a specific journal. The purpose is to have a forum in which general doubts about the processes of publication in the journal, experiences and other issues derived from the publication of papers are resolved. For topics on particular articles, maintain the dialogue through the usual channels with your editor.

Follow us on @ScimagoJR Scimago Lab , Copyright 2007-2024. Data Source: Scopus®

Cookie settings

Cookie Policy

Legal Notice

Privacy Policy

Journal of Hymenoptera Research

( API-Link )

Impact Factor : 1.300 (based on Web of Science 2022)

• # 42 / 84 (Q2) in Entomology

Partner: • University Press Alert

The Journal of Hymenoptera Research is a peer-reviewed, open-access, rapid online journal launched to accelerate research on all aspects of Hymenoptera, including biology, behavior, ecology, systematics, taxonomy, genetics, and morphology.

Some content from Wikipedia , licensed under CC BY-SA

Journal of Hymenoptera Research

• Date 6 hours 12 hours 1 day 3 days all
• Rank Last day 1 week 1 month all
• LiveRank Last day 1 week 1 month all
• Popular Last day 1 week 1 month all

Oldest family of jewel wasps discovered in Cretaceous amber from Lebanon

Jewel wasps (Chalcidoidea) are one of the most diverse groups of insects, with more than 120,000 species described and an estimated true diversity of nearly 1 million. The chalcids are parasitoid wasps, which attack other ...

Oct 25, 2023

New species of parasitic wasp associated with soybean pest in Minnesota

Researchers at the University of Minnesota have discovered a new species of tiny parasitic wasp that might prove beneficial to managing soybean gall midge, a recently emerged pest in Midwest soybean fields that can have devastating ...

May 11, 2023

Tracking an invasion: A single Asian hornet may have sparked the ongoing spread across Europe

In Europe, the Asian (or "Yellow-legged") Hornet (Vespa velutina) is a predator of insects such as honeybees, hoverflies, and other wasps, and poses serious risks to apiculture, biodiversity and pollination services. This ...

Plants & Animals

Dec 6, 2022

New unusual bee species discovered with dog-like snout

A new native bee species with a dog-like "snout" has been discovered in Perth bushland though Curtin-led research that sheds new light on our most important pollinators.

Oct 31, 2022

New 'menehune' wasp species discovered on UH Manoa campus

A new species of wasp was discovered on the University of Hawaiʻi at Mānoa campus. Mymarommatoidea—a group of miniscule but very beautiful wasps, around 0.5 mm in length—had been emerging from branches of a banyan tree ...

Mar 15, 2022

Accidental tree wound reveals novel symbiotic behavior by ants

One afternoon, during the early days of the COVID-19 pandemic in Panama, a bored teenager with a slingshot and a clay ball accidentally shot entry and exit holes in a Cecropia tree trunk. These are "ant-plant" trees, which ...

Jan 3, 2022

The wild cocoons of a new species of Japanese parasitoid wasp

A unique "star" was discovered from the Ryukyu Islands, a biodiversity hot spot in subtropical Japan: a star-shaped structure that turned out to be the cocoon mass of a new species of parasitoid wasp. Researchers Shunpei ...

Nov 30, 2021

The ants, bees and wasps of Canada, Alaska and Greenland: A checklist of 9250 species

Knowing what species live in which parts of the world is critical to many fields of study, such as conservation biology and environmental monitoring. This is also how we can identify present or potential invasive and non-native ...

May 5, 2021

Bees thrive where it's hot and dry: A unique biodiversity hotspot located in North America

The United States-Mexico border traverses through large expanses of unspoiled land in North America, including a newly discovered worldwide hotspot of bee diversity. Concentrated in 16 km2 of protected Chihuahuan Desert are ...

Rare bee found after 100 years

A widespread field search for a rare Australian native bee not recorded for almost a century has found it's been there all along—but is probably under increasing pressure to survive.

Feb 25, 2021

E-mail newsletter

International Society of Hymenopterists

Publications

ISH regularly releases a society newsletter,  Hamuli , and also publishes a peer-reviewed research journal,  Journal of Hymenoptera Research .

ISH Publications

• Journal of Hymenoptera Research
• Genera of New World Braconidae. In English or Spanish. Out of Print.
• Taxapad database of world Ichneumonidae (digital)
• Taxapad database of Chalcidoidea (digital)
• Papers of A. A. Girault. (digital)

Hamuli – The Newsletter of the International Society of Hymenopterists

Hamuli ISSN 2224-2791

 Other hymenopterist newsletters

An archive of important, relevant hymenopterist newsletters.

• Chalcid Forum
• volume 1, issue 1
• volume 1, issue 2
• volume 2, issue 1
• volume 2, issue 2
• volume 3, issue 1

• Special Issues
• Topical collections

Author Guidelines

• Editorial Team

Focus and Scope

Journal of Hymenoptera Research publishes papers of high scientific quality reporting comprehensive research on all aspects of Hymenoptera, including biology, behavior, ecology, systematics, taxonomy, genetics, and morphology. Taxonomic papers describing single species are acceptable if the species has economic importance or provides new data on the biology or evolution of the genus or higher taxon. Manuscript length generally should not exceed 50 typed pages; however, no upper limit on length has been set for papers of exceptional quality and importance, including taxonomic monographs at generic or higher level.

Journal of Hymenoptera Research considers the following categories of papers for publication:

Original research articles,

Reviews as longer articles, offering a comprehensive overview, historical analysis or/and future perspectives of a topic,

Monographs and collection of papers with no limit in size, published as 'special issues',

Short communications,

Letters and Discussion papers,

Book reviews.

The journal will be published online (standard) and as a print version (on request).

The Journal of Hymenoptera Research is published on behalf the International Society of Hymenopterists ( www.hymenopterists.org ).

Open Access

This journal provides immediate open access to its content on the principle that making research freely available to the public supports a greater global exchange of knowledge.

Copyright Notice

License and copyright agreement.

In submitting the manuscript to any of Pensoft’s journals, authors certify that: 

• They are authorized by their co-authors to enter into these arrangements. 
• The work described has not been published before (except in the form of an abstract or as part of a published lecture, review or thesis); it is not under consideration for publication elsewhere; its publication has been approved by all author(s) and responsible authorities – tacitly or explicitly – of the institutes where the work has been carried out. 
• They secure the right to reproduce any material that has already been published or where copyright is owned by someone else. 
• They agree to the following license and copyright agreement:
• Copyright on any article is retained by the author(s) or the author's employer. Regarding copyright transfers please see below. 
• Authors grant Pensoft Publishers a license to publish the article and identify itself as the original publisher. 
• Authors grant any third party the right to use the article freely as long as its original authors and citation details are identified. 
• The article and published supplementary material are distributed under the Creative Commons Attribution License (CC BY 4.0) :

Creative Commons Attribution License (CC BY 4.0)

Anyone is free:

to Share — to copy, distribute and transmit the work  to Remix — to adapt the work

Under the following conditions:

Attribution. The original authors must be given credit. 

• For any reuse or distribution, it must be made clear to others what the license terms of this work are. 
• Any of these conditions can be waived if the copyright holders give permission. 
• Nothing in this license impairs or restricts the author's moral rights.

The full legal code of this license.

Copyright Transfers

Any usage rights are regulated through the Creative Commons License. Since Pensoft Publishers is using the Creative Commons Attribution License (CC BY 4.0), anyone (the author, their institution/company, the publisher and the public) is free to copy, distribute, transmit and adapt the work as long as the original author is credited (see above). Therefore, specific usage rights cannot be reserved by the author or their institution/company and the publisher cannot include a statement "all rights reserved" in any published paper.

Website design and publishing framework: Copyright © Pensoft Publishers.

CLOCKSS system has permission to ingest, preserve, and serve this Archival Unit.

Privacy Statement

The names and email addresses entered in this journal site will be used exclusively for the stated purposes of this journal and will not be made available for any other purpose or to any other party.

Authorship/Contributorship

Some journals are integrated with  Contributor Role Taxonomy  (CRediT), in order to recognise individual author input within a publication, thereby ensuring professional and ethical conduct, while avoiding authorship disputes, gift / ghost authorship and similar pressing issues in academic publishing.

During manuscript submission, the submitting author is strongly recommended to specify a contributor role for each of co-author, i.e. Conceptualization, Methodology, Software, Validation, Formal analysis, Investigation, Resources, Data Curation, Writing - Original draft, Writing - Review and Editing, Visualization, Supervision, Project administration, Funding Acquisition (see  more ). For the journals that are not integrated with CRediT, the submitting author is encouraged to specify the roles as a free text. Once published the article will include the contributor role for all authors in the article metadata.

Authors must be transparent about the use of Artificial Intelligence tools such as ChatGPT and other large language models in the manuscript preparation, and disclose details of how the AI tool was used within the "Materials and Methods" section.

Desk Rejection

During the pre-review evaluation, Editors-in-Chief or Subject editors check the manuscript for compliance with the journal's guidelines, focus, and scope. At this point, they may reject a manuscript prior to sending it out for peer review, specifying the reasons. The most common ones are non-conformity with the journal's focus, scope and policies and/or low scientific or linguistic quality. In such cases, authors are encouraged to considerably improve their manuscript and resubmit it for a review. We encourage authors whose manuscripts have been desk rejected due to being out of the scope of this journal to consider another potentially suitable title from the  Pensoft portfolio .

In case the manuscript is suitable for the journal but has to be corrected technically or linguistically, it will be returned to the authors for improvement. The authors will not need to re-submit the manuscript but only to upload the corrected file(s) to their existing submission.

Peer Review

This journal uses a single-blind peer review process. This means that the names of reviewers are hidden from the authors (the author does not know the identity of the reviewer, but the reviewer knows the identity of the author). Notwithstanding that, the reviewers are encouraged to disclose their identities, if they wish to do so. Each article is reviewed by at least two independent experts, with a final decision on acceptance being made by the Subject Editor / Editor-in-Chief. Front-matter articles, such as editorials, correspondence, biographies, and similar articles, can be published after editorial evaluation only.

Please consider the Editor and Reviewer Guidelines in the About webpage of this journal for more details and stepwise instructions on the editorial and peer review process.

Indexing and Archiving

Publication ethics and malpractice statement.

The publishing ethics and malpractice policies follow the  Principles of Transparency and Best Practice in Scholarly Publishing (joint statement by COPE, DOAJ, WAME, and OASPA), the  NISO Recommended Practices for the Presentation and Identification of E-Journals (PIE-J), and, where relevant, the  Recommendations for the Conduct, Reporting, Editing, and Publication of Scholarly Work in Medical Journals  from ICMJE.

Privacy statement

The personal information used on this website is to be used exclusively for the stated purposes of each particular journal. It will not be made available for any other purpose or to any other party. 

Open access

Pensoft and ARPHA-hosted journals adhere strictly to gold open access to accelerate the barrier-free dissemination of scientific knowledge. All published articles are made freely available to read, download, and distribute immediately upon publication, given that the original source and authors are cited ( Creative Commons Attribution License (CC BY 4.0) ).

Open data publishing and sharing

Pensoft and ARPHA encourage open data publication and sharing, in accordance with  Panton’s Principles  and  FAIR Data Principles . For the domain of biodiversity-related publications Pensoft has specially developed extended  Data Publishing Policies and Guidelines for Biodiversity Data . Specific data publishing guidelines are available on the journal website. 

Data can be published in various ways, such as preservation in data repositories linked to the respective article or as data files or packages supplementary to the article. Datasets should be deposited in an appropriate, trusted repository and the associated identifier (URL or DOI) of the dataset(s) must be included in the data resources section of the article. Reference(s) to datasets should also be included in the reference list of the article with DOIs (where available). Where no discipline-specific data repository exists authors should deposit their datasets in a general repository such as, for example  Zenodo  or others. 

Submission, peer review and editorial process

The peer review and editorial processes are facilitated through an online editorial system and a set of email notifications. Pensoft journals’ websites display stepwise description of the editorial process and list all necessary instructions and links. These links are also included in the respective email notification.

General: Publication and authorship

• All submitted papers are subject to a rigorous peer review process by at least two international reviewers who are experts in the scientific field of the particular paper. 
• The factors that are taken into account in review are relevance, soundness, significance, originality, readability and language. 
• A declaration of potential Conflicts of Interest is a mandatory step in the submission process. The declaration becomes part of the article metadata and is displayed in both the PDF and HTML versions of the article.
• The journals allow several rounds of review of a manuscript. The ultimate responsibility for editorial decisions lies with the respective Subject Editor and, in some cases, with the Editor-in-Chief. All appeals should be directed to the Editor-in-Chief, who may decide to seek advice among the Subject Editors and Reviewers.
• The possible decisions include: (1) Accept, (2) Minor revisions, (3) Major revisions, (4) Reject, but re-submission encouraged and (5) Reject. 
• If Authors are encouraged to revise and re-submit a submission, there is no guarantee that the revised submission will be accepted. 
• The paper acceptance is constrained by such legal requirements as shall then be in force regarding libel, copyright infringement and plagiarism. 
• No research can be included in more than one publication.
• Editors-in-Chief, managing editors and their deputies are strongly recommended to limit the amount of papers co-authored by them. As a rule of thumb, research papers (co-)authored by Editors-in-Chief, managing editors and their deputies must not exceed 20% of the publications a year, with a clear task to drop this proportion below 15%. By adopting this practice, the journal is taking extra precaution to avoid endogeny and conflicts of interest, while ensuring the editorial decision-making process remains transparent and fair.
• Editors-in-Chief, managing editors and handling editors are not allowed to handle manuscripts co-authored by them.

Responsibility of Authors

• Authors are required to agree that their paper will be published in open access under the  Creative Commons Attribution License (CC BY 4.0 ) license.
• Authors must certify that their manuscripts are their original work. 
• Authors must certify that the manuscript has not previously been published elsewhere. 
• Authors must certify that the manuscript is not currently being considered for publication elsewhere. 
• Authors should submit the manuscript in linguistically and grammatically correct English and formatted in accordance with the journal’s Author Guidelines.
• Authors must participate in the peer review process. 
• Authors are obliged to provide retractions or corrections of mistakes. 
• All Authors mentioned are expected to have significantly contributed to the research. 
• Authors must notify the Editors of any conflicts of interest. 
• Authors must identify all sources used in the creation of their manuscript. 
• Authors must report any errors they discover in their published paper to the Editors.
• Authors should acknowledge all significant funders of the research pertaining to their article and list all relevant competing interests.   
• Other sources of support for publications should also be clearly identified in the manuscript, usually in an acknowledgement (e.g. funding for the article processing charge; language editing or editorial assistance).
• The Corresponding author should provide the declaration of any conflicts of interest on behalf of all Authors. Conflicts of interest may be associated with employment, sources of funding, personal financial interests, membership of relevant organisations or others.

Responsibility of Reviewers

• The manuscripts will be reviewed by two or three experts in order to reach first decision as soon as possible. Reviewers do not need to sign their reports but are welcome to do so. They are also asked to declare any conflicts of interests.
• Reviewers are not expected to provide a thorough linguistic editing or copyediting of a manuscript, but to focus on its scientific quality, as well as for the overall style, which should correspond to the good practices in clear and concise academic writing. If Reviewers recognize that a manuscript requires linguistic edits, they should inform both Authors and Editor in the report.
• Reviewers are asked to check whether the manuscript is scientifically sound and coherent, how interesting it is and whether the quality of the writing is acceptable.
• In cases of strong disagreement between the reviews or between the Authors and Reviewers, the Editors can judge these according to their expertise or seek advice from a member of the journal's Editorial Board.

Reviewers are also asked to indicate which articles they consider to be especially interesting or significant. These articles may be given greater prominence and greater external publicity, including press releases addressed to science journalists and mass media.

• During a second review round, the Reviewer may be asked by the Subject Editor to evaluate the revised version of the manuscript with regards to Reviewer’s recommendations submitted during the first review round.
• Reviewers are asked to be polite and constructive in their reports. Reports that may be insulting or uninformative will be rescinded.
• Reviewers are asked to start their report with a very brief summary of the reviewed paper. This will help the Editors and Authors see whether the reviewer correctly understood the paper or whether a report might be based on misunderstanding.
• Further, Reviewers are asked to comment on originality, structure and previous research: (1) Is the paper sufficiently novel and does it contribute to a better understanding of the topic under scrutiny? Is the work rather confirmatory and repetitive? (2) Is the introduction clear and concise? Does it place the work into the context that is necessary for a reader to comprehend the aims, hypotheses tested, experimental design or methods? Are Material and Methods clearly described and sufficiently explained? Are reasons given when choosing one method over another one from a set of comparable methods? Are the results clearly but concisely described? Do they relate to the topic outlined in the introduction? Do they follow a logical sequence? Does the discussion place the paper in scientific context and go a step beyond the current scientific knowledge on the basis of the results? Are competing hypotheses or theories reasonably related to each other and properly discussed? Do conclusions seem reasonable?  Is previous research adequately incorporated into the paper? Are references complete, necessary and accurate? Is there any sign that substantial parts of the paper were copies of other works?
• Reviewers should not review manuscripts in which they have conflicts of interest resulting from competitive, collaborative, or other relationships or connections with any of the authors, companies, or institutions connected to the papers.
• Reviewers should keep all information regarding papers confidential and treat them as privileged information. 
• Reviewers should express their views clearly with supporting arguments. 
• Reviewers should identify relevant published work that has not been cited by the authors.
• Reviewers should also call to the Editors’ attention any substantial similarity or overlap between the manuscript under consideration and any other published paper of which they have personal knowledge.

Responsibility of Editors

• Editors in Pensoft’s journals carry the main responsibility for the scientific quality of the published papers and base their decisions solely on the papers' importance, originality, clarity and relevance to publication's scope.
• The Subject Editor takes the final decision on a manuscript’s acceptance or rejection and his/her name is listed as "Academic Editor" in the header of each article.
• The Subject Editors are not expected to provide a thorough linguistic editing or copyediting of a manuscript, but to focus on its scientific quality, as well as the overall style, which should correspond to the good practices in clear and concise academic writing. 
• Editors are expected to spot small errors in orthography or stylistic during the editing process and correct them.
• Editors should always consider the needs of the Authors and the Readers when attempting to improve the publication. 
• Editors should guarantee the quality of the papers and the integrity of the academic record. 
• Editors should preserve the anonymity of Reviewers, unless the latter decide to disclose their identities. 
• Editors should ensure that all research material they publish conforms to internationally accepted ethical guidelines. 
• Editors should act if they suspect misconduct and make all reasonable attempts to obtain a resolution to the problem. 
• Editors should not reject papers based on suspicions, they should have proof of misconduct.
• Editors should not allow any conflicts of interest between Authors, Reviewers and Board Members.
• Editors are allowed to publish a limited proportion of papers per year co-authored by them, after considering some extra precautions to avoid an impression of impropriety, endogeny, conflicts of interest and ensure that the editorial decision-making process is transparent and fair.
• Editors-in-Chief, managing editors and handling editors are not allowed to handle manuscripts co-authored by them.              

Neutrality to geopolitical disputes

The strict policy of Pensoft and its journals is to stay neutral to any political or territorial dispute. Authors should depoliticize their studies by avoiding provoking remarks, disputable geopolitical statements and controversial map designations; disputable territories should be referred to as well-recognised and non-controversial geographical areas. Тhe journal reserves the right to mark such areas at least as disputable at or after publication, to publish editor's notes, or to reject/retract the paper.

Authors' affiliations

Pensoft does not take decisions regarding the actual affiliations of institutions. Authors are advised to provide their affiliation as indicated on the official internet site of their institution.

Editors 

Editorial decisions should not be affected by the origins of the manuscript, including the nationality, ethnicity, political beliefs, race, or religion of the authors. Decisions to edit and publish should not be determined by the policies of governments or other agencies outside of the journal itself.

Human and animal rights

The ethical standards in medical and pharmacological studies are based on the Helsinki declaration (1964, amended in 1975, 1983, 1989, 1996, 2000 and 2013) of the World Medical Association and the  Publication Ethics Policies for Medical Journals  of the World Association of Medical Journals (WAME).

Authors of studies including experiments on humans or human tissues should declare in their cover letter a compliance with the ethical standards of the respective institutional or regional committee on human experimentation and attach committee’s statement and informed consent; for those researchers who do not have access to formal ethics review committees, the principles outlined in the Declaration of Helsinki should be followed and declared in the cover letter. Patients’ names, initials, or hospital numbers should not be used, not in the text nor in any illustrative material, tables of databases, unless the author presents a written permission from each patient to use his or her personal data. Photos or videos of patients should be taken after a warning and agreement of the patient or of a legal authority acting on his or her behalf.

Animal experiments require full compliance with local, national, ethical, and regulatory principles, and local licensing arrangements and respective statements of compliance (or approvals of institutional ethical committees where such exists) should be included in the article text.

Informed consent

Individual participants in studies have the right to decide what happens to the identifiable personal data gathered, to what they have said during a study or an interview, as well as to any photograph that was taken. Hence it is important that all participants gave their informed consent in writing prior to inclusion in the study. Identifying details (names, dates of birth, identity numbers and other information) of the participants that were studied should not be published in written descriptions, photographs, and genetic profiles unless the information is essential for scientific purposes and the participant (or parent or guardian if the participant is incapable) gave written informed consent for publication. Complete anonymity is difficult to achieve in some cases, and informed consent should be obtained if there is any doubt. If identifying characteristics are altered to protect anonymity, such as in genetic profiles, authors should provide assurance that alterations do not distort scientific meaning.

The following statement should be included in the article t ext in one of the following ways:

• "Informed consent was obtained from all individual participants included in the study."
• "Informed consent was obtained from all individuals for whom identifying information is included in this article." (In case some patients’ data have been published in the article or supplementary materials to it).

Gender issues

We encourage the use of gender-neutral language, such as 'chairperson' instead of 'chairman' or 'chairwomen', as well as 'they' instead of 'she/he' and 'their' instead of  'him/her' (or consider restructuring the sentence).

Conflict of interest

During the editorial process, the following relationships between editors and authors are considered conflicts of interest: Colleagues currently working in the same research group or department, recent co-authors, and doctoral students for which the editor served as committee chair. During the submission process, the authors are kindly advised to identify possible conflicts of interest with the journal editors. After manuscripts are assigned to the handling editor, individual editors are required to inform the managing editor of any possible conflicts of interest with the authors. Journal submissions are also assigned to referees to minimize conflicts of interest. After manuscripts are assigned for review, referees are asked to inform the editor of any conflicts that may exist.

Appeals and open debate

We encourage academic debate and constructive criticism. Authors are always invited to respond to any editorial correspondence before publication. Authors are not allowed to neglect unfavorable comments about their work and choose not to respond to criticisms. 

No Reviewer’s comment or published correspondence may contain a personal attack on any of the Authors. Criticism of the work is encouraged. Editors should edit (or reject) personal or offensive statements. Authors should submit their appeal on editorial decisions to the Editorial Office, addressed to the Editor-in-Chief or to the Managing Editor. Authors are discouraged from directly contacting Editorial Board Members and Editors with appeals.

Editors will mediate all discussions between Authors and Reviewers during the peer review process prior to publication. If agreement cannot be reached, Editors may consider inviting additional reviewers if appropriate. 

The Editor-in-Chief will mediate all discussions between Authors and Subject Editors.

The journals encourage publication of open opinions, forum papers, corrigenda, critical comments on a published paper and Author’s response to criticism.

Research misconduct may include: (a)  manipulating research materials, equipment or processes; (b) changing or omitting data or results such that the research is not accurately represented in the article; c) plagiarism. Research misconduct does not include honest error or differences of opinion. If misconduct is suspected, journal Editors will act in accordance with the relevant  COPE guidelines .

Plagiarism and duplicate publication policy A special case of misconduct is plagiarism, which is the appropriation of another person's ideas, processes, results or words without giving appropriate credit. Plagiarism is considered theft of intellectual property and manuscripts submitted to this journal which contain substantial unattributed textual copying from other papers will be immediately rejected. Editors are advised to check manuscripts for plagiarism via the iThenticate service by clicking on the "ïThenticate report" button. Journal providing a peer review in languages other than English (for example, Russian) may use other plagiarism checking services (for example, Antiplagiat).  Instances, when authors re-use large parts of their publications without providing a clear reference to the original source, are considered duplication of work. Slightly changed published works submitted in multiple journals is not acceptable practice either. In cases of plagiarism in an already published paper or duplicate publication, an announcement will be made on the journal publication page and a procedure of retraction will be triggered.

Responses to possible misconduct

All allegations of misconduct must be referred to the Editor-In-Chief. Upon the thorough examination, the Editor-In-Chief and deputy editors should conclude if the case concerns a possibility of misconduct. All allegations should be kept confidential and references to the matter in writing should be kept anonymous, whenever possible.

Should a comment on potential misconduct be submitted by the Reviewers or Editors, an explanation will be sought from the Authors. If it is satisfactory and the issue is the result of either a mistake or misunderstanding, the matter can be easily resolved. If not, the manuscript will be rejected or retracted and the Editors may impose a ban on that individual's publication in the journals for a certain period of time. In cases of published plagiarism or dual publication, an announcement will be made in both journals explaining the situation.

When allegations concern authors, the peer review and publication process for their submission will be halted until completion of the aforementioned process. The investigation will be carried out even if the authors withdraw the manuscript, and implementation of the responses below will be considered.

When allegations concern reviewers or editors, they will be replaced in the review process during the ongoing investigation of the matter. Editors or reviewers who are found to have engaged in scientific misconduct should be removed from further association with the journal, and this fact reported to their institution.

Retraction policies

Article retraction

According to the  COPE Retraction Guidelines  followed by this Journal, an article can be retracted because of the following reasons:

• Unreliable findings based on clear evidence of a misconduct (e.g. fraudulent use of the data) or honest error (e.g. miscalculation or experimental error).
• Redundant publication, e.g., findings that have previously been published elsewhere without proper cross-referencing, permission or justification.
• Plagiarism or other kind of unethical research.

Retraction procedure

• Retraction should happen after a careful consideration by the Journal editors of allegations coming from the editors, authors, or readers.
• The HTML version of the retracted article is removed (except for the article metadata) and on its place a retraction note is issued.
• The PDF of the retracted article is left on the website but clearly watermarked with the note "Retracted" on each page.
• In some rare cases (e.g., for legal reasons or health risk) the retracted article can be replaced with a new corrected version containing apparent link to the retracted original version and a retraction note with a history of the document.

Expression of concern

In other cases, the Journal editors should consider issuing an expression of concern, if evidence is available for:

• Inconclusive evidence of research or publication misconduct by the authors.
• Unreliable findings that are unreliable but the authors’ institution will not investigate the case.
• A belief that an investigation into alleged misconduct related to the publication either has not been, or would not be, fair and impartial or conclusive.
• An investigation is underway but a judgement will not be available for a considerable time.

Errata and Corrigenda

Pensoft journals largely follow the ICMJE guidelines for corrections and errata.

Admissible and insignificant errors in a published article that do not affect the article content or scientific integrity (e.g. typographic errors, broken links, wrong page numbers in the article headers etc.) can be corrected through publishing of an erratum. This happens through replacing the original PDF with the corrected one together with a correction notice on the Erratum Tab of the HTML version of the paper, detailing the errors and the changes implemented in the original PDF. The original PDF will be marked with a correction note and an indication to the corrected version of the erratum article. The original PDF will also be archived and made accessible via a link in the same Erratum Tab.

Authors are also encouraged to post comments and indicate typographical errors on their articles to the Comments tab of the HTML version of the article.

Corrigenda should be published in cases when significant errors are discovered in a published article. Usually, such errors affect the scientific integrity of the paper and could vary in scale. Reasons for publishing corrigenda may include changes in authorship, unintentional mistakes in published research findings and protocols, errors in labelling of tables and figures or others. In taxonomic journals, corrigenda are often needed in cases where the errors affect nomenclatural acts. Corrigenda are published as a separate publication and bear their own DOI. Examples of published corrigenda are available here .

The decision for issuing errata or corrigenda is with the editors after discussion with the authors.

COPE Compliance

This journal endorses the COPE (Committee on Publication Ethics) guidelines and will pursue cases of suspected research and publication misconduct (e.g. falsification, unethical experimentation, plagiarism, inappropriate image manipulation, redundant publication). For further information about COPE please see the website for COPE at http://www.publicationethics.org and journal's Publication Ethics and Malpractice Statement .

Terms of Use

This document describes the Terms of Use of the services provided by the Journal of Hymenoptera Research journal, hereinafter referred to as "the Journal" or "this Journal". All Users agree to these Terms of Use when signing up to this Journal. Signed Journal Users will be hereinafter referred to as "User" or "Users".

The publication services to the Journal are provided by  Pensoft Publishers Ltd., through its publishing platform ARPHA , hereinafter referred to as "the Provider".

The Provider reserves the right to update the Terms of Use occasionally. Users will be notified via posting on the site and/or by email. If using the services of the Journal after such notice, the User will be deemed to have accepted the proposed modifications. If the User disagrees with the modifications, he/she should stop using the Journal services. Users are advised to periodically check the Terms of Use for updates or revisions. Violation of any of the terms will result in the termination of the User's account. The Provider is not responsible for any content posted by the User in the Journal.

Account Terms

• For registration in this Journal or any of the services or tools hosted on it, Users must provide their full legal name, a valid email address, postal address, affiliation (if any),  and any other information requested.
• Accounts created via this journal automatically sign in the User to the ARPHA Platform.
• Users are responsible for maintaining the security of their account and password. The Journal cannot and will not be liable for any loss or damage from failure to comply with this security obligation.
• Users are solely responsible for the content posted via the Journal services (including, but not limited to data, text, files, information, usernames, images, graphics, photos, profiles, audio and video clips, sounds, applications, links and other content) and all activities that occur under their account.
• Users may not use the service for any illegal or unauthorised purpose. Users must not, in the use of the service, violate any laws within their jurisdiction (including but not limited to copyright or trademark laws).
• Users can change or pseudonomyse their personal data, or deactivate their accounts at any time through the functionality available in the User’s personal profile. Deactivation or pseudonomysation will not affect the appearance of personal data in association with an already published work of which the User is author, co-author, editor, or reviewer.
• Users can report to the Journal uses of their personal data, that they might consider not corresponding to the current Terms of Use.
• The User’s personal data is processed by the Journal on the legal basis corresponding to Article 6, paragraph 1, letters a, b, c and f. of the General Data Protection Regulation (hereinafter referred to as GDPR) and will be used for the purpose of Journal’s services in accordance with the present Terms and Use, as well as in those cases expressly stated by the legislation.
• User’s consent to use the information the Journal has collected about the User corresponds to Article 6(1)(a) of the GDPR.
• The ‘legitimate interest’ of the Journal to engage with the User and enable him/her to participate in Journal’s activities and use Journal’s services correspond to Article 6(1)(f) of the GDPR.

Services and Prices

The Provider reserves the right to modify or discontinue, temporarily or permanently, the services provided by the Journal. Plans and prices are subject to change upon 30 days notice from the Provider. Such notice may be provided at any time by posting the changes to the relevant service website.

The Authors retain full ownership to their content published in the Journal. We claim no intellectual property rights over the material provided by any User in this Journal. However, by setting pages to be viewed publicly (Open Access), the User agrees to allow others to view and download the relevant content. In addition, Open Access articles might be used by the Provider, or any other third party, for data mining purposes. Authors are solely responsible for the content submitted to the journal and must confirm [during the submission process] that the content does not contain any materials subject to copyright violation including, but not limited to, text, data, multimedia, images, graphics, photos, audio and video clips. This requirement holds for both the article text and any supplementary material associated with the article.

The Provider reserves the rights in its sole discretion to refuse or remove any content that is available via the Website.

Copyrighted Materials

Unless stated otherwise, the Journal website may contain some copyrighted material (for example, logos and other proprietary information, including, without limitation, text, software, photos, video, graphics, music and sound - "Copyrighted Material"). The User may not copy, modify, alter, publish, transmit, distribute, display, participate in the transfer or sale, create derivative works or, in any way, exploit any of the Copyrighted Material, in whole or in part, without written permission from the copyright owner. Users will be solely liable for any damage resulting from any infringement of copyrights, proprietary rights or any other harm resulting from such a submission.

Exceptions from this rule are e-chapters or e-articles published under Open Access (see below), which are normally published under  Creative Commons Attribution 3.0 license (CC-BY), or Creative Commons Attribution 4.0 license (CC-BY), or Creative Commons Public Domain license (CC0).

Open Access Materials

This Journal is a supporter of open science. Open access to content is clearly marked, with text and/or the open access logo, on all materials published under this model. Unless otherwise stated, open access content is published in accordance with the Creative Commons Attribution 4.0 licence (CC-BY). This particular licence allows the copying, displaying and distribution of the content at no charge, provided that the author and source are credited.

• Users agree to submit their personal data to this Journal, hosted on the ARPHA Platform provided by Pensoft.
• The Journal collects personal information from Users (e.g., name, postal and email addresses, affiliation) only for the purpose of its services.
• All personal data will be used exclusively for the stated purposes of the website and will not be made available for any other purpose or to third parties.
• In the case of co-authorship of a work published through the Journal services, each of the co-authors states that they agree that their personal data be collected, stored and used by the Journal.
• In the case of co-authorship, each of the co-authors agrees that their personal data publicly available in the form of a co-authorship of a published work, can be distributed to external indexing services and aggregators for the purpose of the widest possible distribution of the work they co-author.
• When one of the co-authors is not registered in the Journal, it is presumed that the corresponding author who is registered has requested and obtained his/her consent that his/her personal data will be collected, stored and used by the Journal.
• The registered co-author undertakes to provide an e-mail address of the unregistered author, to whom the Journal will send a message in order to give the unregistered co-author’s explicit consent for the processing of his/her personal data by the Journal.
• The Journal is not responsible if the provided e-mail of the unregistered co-author is inaccurate or invalid. In such cases, it is assumed that the processing of the personal data of the unregistered co-author is done on a legal basis and with a given consent.
• The Journal undertakes to collect, store and use the provided personal data of third parties (including but not limited to unregistered co-authors) solely for the purposes of the website, as well as in those cases expressly stated by the legislation.
• Email notifications to authors, reviewers and editors who are engaged with authoring, reviewing or editing a manuscript submitted to the Journal.
• Email alerts sent via email subscription service, which can happen only if the User has willingly subscribed for such a service. Unsubscription from the service can happen through a one-click link provided in each email alert notification.
• Information emails on important changes in the system or in its Terms of Use which are sent via Mailchimp are provided with "Unsubscribe" function.
• Registered users can be invited to provide a peer review on manuscripts submitted to the Journal. In such cases, the users can decline the review invitation through a link available on the journal’s website.
• Each provided peer review can be registered with external services (such as Web of Science Reviewer Recognition Service, formerly Publons). The reviewer will be notified if such registration is going to occur and can decline the registration process.
• In case the Journal starts using personal data for purposes other than those specified in the Terms of Use, the Journal undertakes to immediately inform the person and request his/her consent.
• If the person does not give his/her consent to the processing of his or her personal data pursuant to the preceding paragraph, the Journal shall cease the processing of the personal data for the purposes for which there is no consent, unless there is another legal basis for the processing.
• Users can change/correct their personal data anytime via the functionality available in the User’s profile. Users can request the Journal to correct their personal data if the data is inaccurate or outdated and the Journal is obliged to correct the inaccurate or outdated personal data in a timely manner.
• Users may request the Journal to restrict the use of their personal data insofar as this limitation is not contrary to the law or the Terms of Use.
• Users may request their personal data to be deleted (the right to be forgotten) by the Journal, provided that the deletion does not conflict with the law or the Terms of Use.
• whether his or her personal data have been processed;
• for which purposes the Journal processes the personal data;
• the ways in which his/her personal data are processed;
• the types of personal data that Journal processes.
• The user undertakes not to interfere with and impede the Journal’s activities in the exercise of the provided rights.
• In case of non-fulfillment under the previous paragraph, the Journal reserves the right to delete the user's profile.

Disclaimer of Warranty and Limitation of Liability

Neither Pensoft and its affiliates nor any of their respective employees, agents, third party content providers or licensors warrant that the Journal service will be uninterrupted or error-free; nor do they give any warranty as to the results that may be obtained from use of the journal, or as to the accuracy or reliability of any information, service or merchandise provided through Journal.

Legal, medical, and health-related information located, identified or obtained through the use of the Service, is provided for informational purposes only and is not a substitute for qualified advice from a professional.

In no event will the Provider, or any person or entity involved in creating, producing or distributing Journal or the contents included therein, be liable in contract, in tort (including for its own negligence) or under any other legal theory (including strict liability) for any damages, including, but without limitation to, direct, indirect, incidental, special, punitive, consequential or similar damages, including, but without limitation to, lost profits or revenues, loss of use or similar economic loss, arising from the use of or inability to use the journal platform. The User hereby acknowledges that the provisions of this section will apply to all use of the content on Journal. Applicable law may not allow the limitation or exclusion of liability or incidental or consequential damages, so the above limitation or exclusion may not apply to the User. In no event will Pensoft’s total liability to the User for all damages, losses or causes of action, whether in contract, tort (including own negligence) or under any other legal theory (including strict liability), exceed the amount paid by the User, if any, for accessing Journal.

Third Party Content

The Provider is solely a distributor (and not a publisher) of SOME of the content supplied by third parties and Users of the Journal. Any opinions, advice, statements, services, offers, or other information or content expressed or made available by third parties, including information providers and Users, are those of the respective author(s) or distributor(s) and not of the Provider.

Cookies Policy

a) Session cookies

We use cookies on our website. Cookies are small text files or other storage technologies stored on your computer by your browser. These cookies process certain specific information about you, such as your browser, location data, or IP address.  

This processing makes our website more user-friendly, efficient, and secure, allowing us, for example, to allow the "Remember me" function.

The legal basis for such processing is Art. 6 Para. 1 lit. b) GDPR, insofar as these cookies are used to collect data to initiate or process contractual relationships.

If the processing does not serve to initiate or process a contract, our legitimate interest lies in improving the functionality of our website. The legal basis is then Art. 6 Para. 1 lit. f) GDPR.

When you close your browser, these session cookies are deleted.

b) Disabling cookies

You can refuse the use of cookies by changing the settings on your browser. Likewise, you can use the browser to delete cookies that have already been stored. However, the steps and measures required vary, depending on the browser you use. If you have any questions, please use the help function or consult the documentation for your browser or contact its maker for support. Browser settings cannot prevent so-called flash cookies from being set. Instead, you will need to change the setting of your Flash player. The steps and measures required for this also depend on the Flash player you are using. If you have any questions, please use the help function or consult the documentation for your Flash player or contact its maker for support.

If you prevent or restrict the installation of cookies, not all of the functions on our site may be fully usable.

What Can I Publish

Book review.

Published on agreement

Peer-reviewed and indexed; large-scale catalogues will be treated as monographs and will bear both ISBN and ISSN numbers

Peer-reviewed and indexed; large-scale checklists will be treated as monographs and will bear both ISBN and ISSN numbers

Published upon editorial decision

Peer-reviewed and indexed

Forum Paper

Published on editorial decision; indexed

In Memoriam

Letter to the editor.

Peer-reviewed and indexed in both journal and book registers; bears ISSN and ISBN numbers

Research Article

Review article, short communication.

Title: The title should be in a sentence case (only scientific, geographic or person names should be with a first capital letter, i.e. Elater ferrugineus L., Germany, etc.), and should include an accurate, clear and concise description of the reported work, avoiding abbreviations. The titles of taxonomic papers should always include the upper rank taxa separated with comma in the following order, in brackets: (Order, Family).

Authors and Affiliations: Provide the complete names of all authors, and their addresses for correspondence, including e.g., institutional affiliation (e.g. university, institute), location (street, boulevard), city, state/province (if applicable), and country. One of the authors should be designated as the corresponding author. It is the corresponding author's responsibility to ensure that the author list, and the individual contributions to the study are accurate and complete. If the article has been submitted on behalf of a consortium, all consortium members and their affiliations should be listed after the Acknowledgements section.

Abstract and Keywords: Please have your abstract and keywords ready for input into the submission module.

Body Text: Manuscripts must be submitted in English. Authors should confirm the English language quality of their texts or alternatively request thorough linguistic editing prior to peer-review at a price. Manuscripts written in poor English are a subject of rejection prior to peer-review. Use either British/Commonwealth or American English provided that the language is consistent within the paper. Each text must be written with precision, clarity, and economy, whenever appropriate in active voice and first person. Avoid the use of parenthetical comments and italics or bold for emphasis. This journal discourages the use of quotation marks except for direct quotations, words defined by the author, and words used in unusual contexts. Short quotations should be embedded in the text and enclosed in double quotation marks (''). Long quotations should be on a separate line, italicized, but without quotation marks. Single quotation marks are to be used only for a quotation that occurs within another quotation.

Spacing, Fonts, Line and Page Numbering: 1.5 or double space all text and quotations, single space figure legends, tables, references, etc. Separate paragraphs with a blank line. Use a 12-point font (preferably Times New Roman or Arial). Please provide line numbers as well as page numbers.

Capitals: First capital letters should be used only in the beginning of a sentence, in proper names and in headings and subheadings, as well as to indicate tables, graphs and figure/s within the text. Software programmes should be written with capital letters (e.g., ANOVA, MANOVA, PAUP).

Italicization/Underlining: Scientific names of species and genera, long direct quotations and symbols for variables and constants (except for Greek letters), such as p, F, U, T, N, r, but not for SD (standard deviation), SE (standard error), DF (degrees of freedom) and NS (non significant) should be italicized. These symbols in illustrations and equations should be in italics to match the text. Italics should not be used for emphasis, and not in abbreviations such as e.g., i.e., et al., etc., cf. Underlining of any text is not acceptable.

Abbreviations: Abbreviations should be followed by ‘.' (full stop or period; for instance: i.e., e.g., cf., etc.). Note that you shouldn't add a full stop at the end of abbreviated words if the last letter of the abbreviation is the same as the last letter of the full word. For example, you should abbreviate "Eds", "Dr", "Mr" without full stop at the end. All measures, for instance mm, cm, m, s, L, should be written without full stop.

On the use of dashes: (1) Hyphens are used to link words such as personal names, some prefixes and compound adjectives (the last of which vary depending on the style manual in use). (2) En-dash or en-rule (the length of an 'n') is used to link spans. In the context of our journal en-dash should be used to link numerals, sizes, dates and page numbers (e.g., 1977–1981; figs 5–7; pp. 237–258); geographic or name associations (Murray–Darling River; a Federal–State agreement); and character states combinations such as long–pubescent or red–purple. (3) Em-dash or em-rule (the length of an 'm') should be used rarely, only for introducing a subordinate clause in the text that is often used much as we use parentheses. In contrast to parentheses an em-dash can be used alone. En-dashes and em-dashes should not be spaced.

Footnotes: Avoid footnotes in the body text of the manuscript. It is always possible to incorporate the footnote into the main text by rewording the sentences, which greatly facilitates reading. Additionally, footnotes are not always handled well by the journal software, and their usage may cause a failure of submission. Footnotes are acceptable only below tables; instead of numbers, please use (in order): †, ‡, §, |, ¶, #, ††, ‡‡, §§, ||, ¶¶, ##.

Geographical coordinates: It is strongly recommended to list geographical coordinates as taken from GPS or online gazetteer, or georeferencer (

http://wwold.gbif.org/prog/digit/Georeferencing ). Geographical coordinates must be listed in one of the following formats:

Definition: The locality consists of a point represented by coordinate information in the form of latitude and longitude. Information may be in the form of

Degrees, Minutes and Seconds (DMS),

Degrees and Decimal Minutes (DDM), or

Decimal Degrees (DD).

Records should also contain a hemisphere (E or W and N or S) or, with Decimal Degrees, minus (–) signs to indicate western and/or southern hemispheres.

Example 1: 36°31'21"N; 114°09'50"W (DMS)

Example 2: 36°31.46'N; 114°09.84'W (DDM)

Example 3: 36.5243°S; 114.1641°W (DD)

Example 4: −36.5243; −114.1641 (DD using minus signs to indicate southern and western hemispheres)

Note on accuracy: Because GPS units are very commonly used today to record latitude/longitude, many authors simply give the GPS readings for their localities. However, these readings are much too accurate. For example, a GPS unit might give the latitude in decimal seconds as 28°16'55.87"N. Since one second of latitude is about 30 m on the ground, the second figure after the decimal in 55.87 represents 30 cm, yet a typical handheld GPS unit is only accurate at best to a few metres.

We therefore recommend two ways to report GPS-based locations. If you give the GPS reading without rounding off, make sure you include an uncertainty figure as a context for the over-accurate GPS reading. We recommend the Darwin Core definition of uncertainty ( http://rs.tdwg.org/dwc/terms/index.htm#coordinateUncertaintyInMeters ):

"The horizontal distance (in meters) from the given decimalLatitude and decimalLongitude describing the smallest circle containing the whole of the Location."

If you only give the GPS reading, please round it off to an implied precision appropriate to the error in the measurement, or to the extent of the area sampled. We suggest rounding off

• to the nearest second in degree-minute-second format (28°16'56"N), which implies roughly ± 25-30 m at middle latitudes
• to four decimal places in decimal degree format (28.2822°N), which implies roughly ± 10-15 m at middle latitudes
• to two decimal places in decimal minute format (28°16.93'N), which implies roughly 15-20 m at middle latitudes

Altitude: Many GPS users simply record the elevation given by their GPS unit. However, GPS elevation is NOT the same as elevation above sea level. GPS units record the elevation above a mathematical model of the earth's surface. The difference between this elevation and elevation above sea level can be tens of metres. In any case, the accuracy of a GPS elevation is often the same as the usual accuracy in horizontal position, so a GPS elevation such as '753 m' is much too accurate and should be rounded off to 'ca 750 m'.

We strongly recommend the use of Example 2 (the DDM format). The other three are also possible but will be recalculated to DDM during the process of online mapping from the HTML version of the paper.

The only restriction on format is in creating a KML (Keyhole Markup Language) file. KML latitudes and longitudes must be in the DD format shown above in Example 4.

Please also consider submitting a table of localities with your manuscript, either as a spreadsheet or in CSV text format. By doing so you will make your specimen localities much more easily available for use in biodiversity databases and geospatial investigations. The geospatial table will be put online as supplementary material for your paper. A minimum table will have three fields: species (or subspecies) name, latitude and longitude. A full table will have the same data for each specimen lot as appears in the text of your paper. Please check latitude/longitude carefully for each entry.

Units: Use the International System of Units ( SI ) for measurements. Consult Standard Practice for Use of the International System of Units (ASTM Standard E−380−93) for guidance on unit conversions, style, and usage.

Statistics: Use leading zeroes with all numbers, including probability values (e.g., P < 0.001). For every significant F−statistic reported, provide two df values (numerator and denominator). Whenever possible, indicate the year and version of the statistical software used.

Web (HTML) links: Authors are encouraged to include links to other Internet resources in their article. This is especially encouraged in the reference section. When inserting a reference to a web-page, please include the http:// portion of the web address.

Supplementary files: Larger datasets can be uploaded separately as Supplementary Files. Tabular data provided as supplementary files can be uploaded as an Excel spreadsheet (.xls), as an OpenOffice spreadsheets (.ods) or comma separated values file (.csv). As with all uploaded files, please use the standard file extensions.

Headings and subheadings: Main headings: The body text should be subdivided into different sections with appropriate headings. Where possible, the following standard headings should be used: Introduction, Methods, Results, Discussion, Conclusions, Acknowledgements, References. These headings need to be in bold font on a separate line and start with a first capital letter. Please do not number headings or subheadings.

Introduction − The motivation or purpose of your research should appear in the Introduction, where you state the questions you sought to answer, and then provide some of the historical basis for those questions.

Methods − Provide sufficient information to allow someone to repeat your work. A clear description of your experimental design, sampling procedures, and statistical procedures is especially important in papers describing field studies, simulations, or experiments. If you list a product (e.g., animal food, analytical device), supply the name and location of the manufacturer. Give the model number for equipment used. Supply complete citations, including author (or editor), title, year, publisher, and version number, for computer software mentioned in your article.

Results − Results should be stated concisely and without interpretation.

Discussion − Focus on the rigorously supported aspects of your study. Carefully differentiate the results of your study from data obtained from other sources. Interpret your results, relate them to the results of previous research, and discuss the implications of your results or interpretations. Point out results that do not support speculations or the findings of previous research, or that are counter-intuitive. You may choose to include a Speculation subsection in which you pursue new ideas suggested by your research, compare and contrast your research with findings from other systems or other disciplines, pose new questions that are suggested by the results of your study, and suggest ways of answering these new questions.

Conclusion −This should state clearly the main conclusions of the research and give a clear explanation of their importance and relevance. Summary illustrations may be included.

References − The list of References should be included after the final section of the main article body. A blank line should be inserted between single-spaced entries in the list. Authors are requested to include links to online sources of articles, whenever possible!

Where possible, the standard headings should be used in the order given above. Additional headings and modifications are permissible.

Subordinate headings: Subordinate headings (e.g. Field study and Simulation model or Counts, Measurements and Molecular analysis ), should be left-justified, italicized, and in a regular sentence case. All subordinate headings should be on a separate line.

English Language Editing

This journal has well-defined policies for English language editing. 

Authors are required to have their manuscripts written in fluent English or edited by a professional English language editor  BEFORE submission. Authors have to confirm by checking a tick box in the submission process that they have followed the above requirement:

"The text is edited by a professional English language editor, duly acknowledged in the manuscript. I am aware that non-edited manuscripts could be rejected prior to peer-review".

The submission process includes an option to request a professional linguistic editing at a price of EURO 15 per 1800 characters :

The authors are NOT obliged to use Journal's linguistic services, but they must ensure that their manuscripts have passed a proper linguistic editing before submission.

Citations and References

Citations within the text:  Before submitting the manuscript, please check each citation in the text against the References and vice-versa to ensure that they match exactly.

Citations in the text should be formatted as follows:

One author:  Smith (1990) or (Smith 1990) 

Note:  The citations format depends on the way it is incorporated within the article’s text:

According to Smith (1990), these findings…

These findings have been first reported in the beginning of the nineties (Smith 1990).

Two authors:   Brock and Gunderson (2001) or (Brock and Gunderson 2001)

Note:  When choosing between formats refer back to examples above.

Three or more authors:   Smith et al. (1998) or (Smith et al. 1998)

When  citing more than one source , in-text citations should be ordered by the year of publication, starting with the earliest one:

(Smith et al. 1998, 2000, 2016; Brock and Gunderson 2001; Felt 2006).

Note:  When you have a few citations from the same author but from different years (such as the case with Smith et al. above), the first year is taken into consideration when ordering the sources (in this case 1998, which is why Smith et al. come first in the list).

When having  two or more fully identical citations  (this can happen when you have more than one reference with exactly the same authors and years for one or two authors, or the same first author and year for author teams of three or more), the references are distinguished by adding the letters 'a', 'b', 'c', etc. after the years and this marking is followed in the in-text citations, respectively:

(Reyes-Velasco et al. 2018a, Reyes-Velasco et al. 2018b)

Authorship  references for species  should include a "," between author and year:

Brianmyia stuckenbergi  Woodley, 2012.

References: It is important to format the references properly, because all references will be linked electronically as completely as possible to the papers cited. It is desirable to add a DOI (digital object identifier) number for either the full-text or title and abstract of the article as an addition to traditional volume and page numbers. If a DOI is lacking, it is recommended to add a link to any online source of an article. Please use the following style for the reference list (or download the Pensoft EndNote style ): here . It is   also available in Zotero, when searched by journal name.

Published Papers: Araújo MB, Cabeza M, Thuiller W, Hannah L, Williams PH (2004) Would climate change drive species out of reserves? An assessment of existing reserve-selection methods. Global Change Biology 10: 1618-1626.

Accepted Papers: Same as above, but ''in press'' appears instead the year in parentheses. 

Electronic Journal Articles: Thuiller W, Araújo MB, Pearson RG, Whittaker R J, Brotons L, Lavorel S (2004) Biodiversity conservation - Uncertainty in predictions of extinction risk. Nature 430.  https://doi.org/10.1038/nature02716 .

Paper within conference proceedings: Orr AG (2006) Odonata in Bornean tropical rain forest formations: Diversity, endemicity and applications for conservation management. In Cordero Rivera A (Ed.) Forests and Dragonflies: Fourth WDA International Symposium of Odonatology, Pontevedra (Spain), July 2005. Pensoft Publishers (Sofia-Moscow): 51-78.

Book chapter: May RM, Lawton JH, Stork NE (1995) Assessing extinction rates. In: Lawton JH, May RM (Eds) Extinction rates. Oxford University Press (New York): 1-24.

Book: Hanski I (2005) The shrinking world: Ecological consequences of habitat loss. International Ecological Institute (Oldendorf/Luhe): 1-307.

Book with institutional author: European Environment Agency (2005) Vulnerability and Adaptation to Climate Change in Europe. EEA Technical Report 7/2005: 1-84.

PhD thesis: Kawanishi K (2002) Population status of tigers ( Panthera tigris ) in a primary rainforest of Peninsular Malaysia. PHD Thesis. University of Florida (Gainesville).

Link/URL: BBC News: Plants 'can recognise themselves'.  http://news.bbc.co.uk/earth/hi/earth_news/newsid_8076000/8076875.stm

Citations of Public Resource Databases:

It is highly recommended all appropriate datasets, images, and information to be deposited in public resources. Please provide the relevant accession numbers (and version numbers, if appropriate). Accession numbers should be provided in parentheses after the entity on first use. Examples of such databases include, but are not limited to:

• IPNI ( www.ipni.org )
• Index Fungorum ( www.indexfungorum.org )
• LIAS ( www.liasnames.lias.net )
• ZooBank ( www.zoobank.org )
• Genbank ( www.ncbi.nlm.nih.gov/Genbank )
• BOLD ( www.barcodinglife.org )

Providing accession numbers to data records stored in global data aggregators allows us to link your article to established databases, thus integrating it with a broader collection of scientific information. Please hyperlink all accession numbers through the text or list them directly after the References in the online submission manuscript.

All journal titles should be spelled out completely and should not be italicized.

Provide the publisher's name and location when you cite symposia or conference proceedings; distinguish between the conference date and the publication date if both are given. Do not list abstracts or unpublished material in the References. They should be quoted in the text as personal observations, personal communications, or unpublished data, specifying the exact source, with date, if possible. When possible, include URLs for articles available online through library subscription or individual journal subscription, or through large international archives, indexes and aggregators, e.g., PubMedCentral, Scopus, CAB Abstracts, etc. URLs for pdf articles that are posted on personal websites only should be avoided.

Authors are encouraged to cite in the References list the publications of the original descriptions of the taxa treated in their manuscript.

Ordering references:  All references should be ordered alphabetically.

If the references have  the same first author and a varying number of co-authors,  the ordering should be based on the number of co-authors starting with the lowest as follows:

Smith J (2018) Article Title. Journal Name 1: 1-10.  https://doi.org/10.3897

Smith J, Gunderson A (2017) Article Title. Journal Name 1: 10-20.  https://doi.org/10.3897

Smith J, Gunderson A, Brock B (2015) Article Title. Journal Name 1: 20-30.  https://doi.org/10.3897

In the occasion of  more than one article from the same first author  within any of the categories above, the references should be ordered chronologically.

If both the  first author and year of publication match  within the categories above, the references are distinguished by adding the letters 'a', 'b', 'c', etc. after the year of publication and this marking is followed in the in-text citations, respectively.

Illustrations, Figures and Tables

Figures and illustrations are accepted in the following image file formats:

• EPS (preferred format for diagrams)
• TIFF (at least 300dpi resolution, with LZW compression)
• PNG (preferred format for photos or images)
• JPEG (preferred format for photos or images)

Vector files in any of the following formats EPS, SVG  or PDF are requested for phylogenetic trees and cladograms.

The journal is printed in A4 paper size with the maximum printing area of 167 mm × 242 mm. Whenever possible, individual figures should be prepared as composite figures.

Should you have any problems in providing the figures in one of the above formats, or in reducing the file below 20 MB , please contact the Editorial Office at [email protected]

Figure legends: All figures should be referenced consecutively in the manuscript; legends should be listed consecutively immediately after the References. For each figure, the following information should be provided: Figure number (in sequence, using Arabic numerals − i.e. Figure 1, 2, 3 etc.); short title of figure (maximum 15 words); detailed legend, up to 300 words.

Illustrations of measurable morphological traits should bear mute scale bars, whose real size is to be given in the figure captions.

Please note that it is the responsibility of the author(s) to obtain permission from the copyright holder to reproduce figures or tables that have previously been published elsewhere.

Figure citations in the text should always be with Capital "F" and En-dash for ranges. One figure with a full stop, figures without.

Example: Fig. 1, Figs 1–3, Fig. 2A–E.

Citations of figures from other publications should always be Lower Case (fig. / figs). When two subsequent figures or parts are cited (for instance figures 1 and 2 or A and B), a comma should be used.

Example:   Figs 1, 2 and Fig. 1A, B.

Parts belong to one figure.

Example: Fig. 1A, B and Fig. 2A-E.

On the use of Google Maps All uses of  Google Maps  and  Google   Earth Content must provide attribution to Google , according to Google Maps/Earth Additional Terms of Service  (see also  Permission Guidelines for Google Maps and Google Earth ). The attribution should be visible on each map in the form, for example: "Map data 2019 (C) Google".

Tables: Each table should be numbered in sequence using Arabic numerals (i.e. Table 1, 2, 3 etc.). Tables should also have a title that summarizes the whole table, maximum 15 words. Detailed legends may then follow, but should be concise.

Small tables can be embedded within the text, in portrait format (note that tables on a landscape page must be reformatted onto a portrait page or submitted as additional files). These will be typeset and displayed in the final published form of the article. Such tables should be formatted using the 'Table object' in a word processing program to ensure that columns of data are kept aligned when the file is sent electronically for review. Do not use tabs to format tables or separate text. All columns and rows should be visible, please make sure that borders of each cell display as black lines. Colour and shading should not be used; neither should commas be used to indicate decimal values. Please use a full stop to denote decimal values (i.e., 0.007 cm, 0.7 mm).

Larger datasets can be uploaded separately as Supplementary Files. Tabular data provided as supplementary files can be uploaded as an Excel spreadsheet (.xls), as an OpenOffice spreadsheets (.ods) or comma-separated values file (.csv). As with all uploaded files, please use the standard file extensions.

Material Citations Formatting Guidelines

Authors are strongly encouraged to adhere to the new fine-grained formatting of the material examined (species occurrence records) as shown below. Please note that this new fine-grained formatting is NOT COMPULSORY and is introduced for testing for an undefined period, during which we hope to receive your feedback. Nonetheless, it is recommended to follow the guidelines to ensure accurate conversion of your records to XML and consequent indexing in GBIF, Plazi, and other important resources. Indexing of each individual occurrence record in GBIF and linking back to your article will provide much higher visibility, data usability, dissemination, and citation probability of your work!  Adapted from: CETAF best practices in electronic publishing in taxonomy ( https://doi.org/10.5852/ejt.2018.475 ). 

Each material citation is composed of diverse data fields (material, locality, date, etc.) that are tagged, using Darwin Core (DWC) terms. To efficiently perform this, it is important to ensure that the different fields of a material citation are consistently presented in the same order throughout the article or, at the very least, within a taxon treatment.

The preferred order is as follows:

For botanical and mycological data, please use "Collection number" instead of "collector [followed by "leg."]". The collection number encompasses all gatherings from a single specimen (e.g., leaves, flowers, piece of wood) which may be preserved on different herbarium sheets and in different herbaria.

The preferred format for botanical/mycological data should be as follows:

Punctuation

A bullet point (• [unicode: 2022]) is used to signify the beginning of a material citation. Within each citation, the different fields are delimited by a semicolon. A single field can be composed of several elements, which are separated by commas (e.g. the details region, area, town and street for the ‘locality’ field).

Semicolons should not be used elsewhere in a material citation.

Repetitive data

Authors can indicate repetitive data with indications such as "same data as for holotype", "same data as for preceding", "same locality", "ibid", etc. as long as the same method and wording are used consistently throughout the paper.

If a material citation is identical to another with only one or two differences, the exceptions should be listed after the mention of repeated data e.g.:

‘Missing’ elements

It is not necessary to include information such as "no date" or "no locality data"; just list the elements that are available.

Label citations

We recommend including photos of labels as figures if they contain data that cannot be standardised. Double quote marks (" ") must be used to represent label citations that do not correspond to (or cannot be reliably interpreted as) specific DWC terms. This data will simply be parsed as a verbatim citation.

Only quote marks should be used to present verbatim label data and they should not appear elsewhere in a material citation.

Author interpretation

Use square brackets [ ] to distinguish data that has been interpreted by the author e.g., coordinates interpreted from a locality, or translations of label data:

Data fields

The different data fields that are tagged in a material citation are explained below, along with the formatting needed to achieve maximum output and precision.

Any specimen data presented in a separate table cannot be linked back to the citation and tagged for conversion.

Country ( country / waterBody )

The citations must be listed by either country or water body, using a separate paragraph for each new zone. Countries should be listed in alphabetical order. If another method is used (e.g. geographic groups) please state this in the Material and Methods section.

If the material is organised by region, please use the following format:

Material ( organismQuantity / organismQuantityType / sex / typeStatus )

This field comprises several indications about the specimens cited: number , type (e.g. specimen, juv., shell, excuviae), sex and type status . All subsequent elements of a citation will be applied to the specimens presented in this field.

Locality ( higherGeography )

The locality data is listed from least to most specific, using commas to divide each detail. It is recommended to employ the English name in current usage where possible.

If there is a particular reason to use a different system, e.g. spelling/transcription variations or archaic names, such details should preferably be identified using quotes, with their current names given in square brackets , but this is not mandatory.

Geo-coordinates ( decimalLatitude / decimalLongitude )

Diverse formats are accepted but it is important to include the degrees symbol (° [unicode: 00B0]), which distinguishes the data as a geo-coordinate. It is also preferable to include the direction (N/E/S/W):

Geo-coordinates should be presented to a maximum of 5 decimal places.

Altitude/elevation/depth ( verbatimAltitude / verbatimDepth )

This type of measurement should be explicit in the material citations, e.g.:

Collection date ( eventDate )

Format: d(d) Mmm. YYYY

Date ranges should be shown with an n-dash, e.g.:

Collector ( recordedBy )

The name(s) of the collector(s) should always be followed by "leg."; for institutions or collecting programmes, "exped." can be used, e.g. MNHN exped.

For botanical and mycological data, "Collection number" instead of "collector [followed by "leg."]". The collection number encompasses all gatherings from a single specimen (e.g., leaves, flowers, piece of wood) which may be preserved on different herbarium sheets and in different herbaria.

Additional data

Ideally, the data fields identified above should be listed before other collection data. If you choose to use a different order, it is important to be as consistent as possible throughout the paper, or at least within a single treatment. You may use a semicolon to separate the additional data into appropriate fields, e.g.:

Additional data can also be given in the appropriate field between brackets, e.g.:

Associated sequences

Accession numbers and barcodes should be identified as such, e.g.:

Repository data ( institutionCode / catalogNumber )

The repository data field should be composed of an institution acronym and a catalogue number (where available), using a colon to separate the two elements.

Institution acronym

All acronyms for repositories must feature in a distinct list in the Materials and Methods section, under the title Repositories, Institutional acronyms or Institutional abbreviations .

Specimen code

The specimen/catalogue code(s) should be listed after the institution code.

Where a specimen code is available, it should be explicit which specimen it refers to. This guarantees unambiguous interpretation, both by readers and upon XML conversion.

E.g., in the citation below, we cannot distinguish which specimens are catalogued under which code:

This citation should be presented as follows:

Exceptions:

1.   If several specimens share the same code, e.g.:

2.   If the specimens possess identical data, including the sex, their specimen codes can be given together, e.g.:

Ranges & multiple specimens

Use the word "to" in order to show a range of specimen numbers. E.g.:

Materials and Methods

In line with responsible and reproducible research, as well as FAIR ( Findability, Accessibility, Interoperability and Reusability)  data principles, we highly recommend that authors describe in detail and deposit their science methods and laboratory protocols in the open access repository protocols.io .

Once deposited on protocols.io , protocols and methods will be issued a unique digital object identifier (DOI), which could be then used to link a manuscript to the relevant deposited protocol. By doing this, authors could allow for editors and peers to access the protocol when reviewing the submission to significantly expedite the process.

Furthermore, an author could open up his/her protocol to the public at the click of a button as soon as their article is published.

Stepwise instructions:

• Prepare a detailed protocol via protocols.io .
• Click Get DOI to assign a persistent identifier to your protocol.
• Add the DOI link to the Methods section of your manuscript prior to submitting it for peer review.
• Click Publish to make your protocol openly accessible as soon as your article is published (optional).
• Update your protocols anytime.

Supplementary Files

Online publishing allows an author to provide datasets, tables, video files, or other information as supplementary information, greatly increasing the impact of the submission. Uploading of such files is possible in Step 9 of the submission process.

The maximum file size for each Supplementary File is 20 MB.

The Supplementary Files will not be displayed in the printed version of the article but will exist as linkable supplementary downloadable files in the online version.

While submitting a supplementary file the following information should be completed:

File format (including name and a URL of an appropriate viewer if format is unusual)

Title of data

Description of data

All supplementary files should be referenced explicitly by file name within the body of the article, e.g. 'See supplementary file 1: Movie 1" for the original data used to perform this analysis.

Ideally, the supplementary files should not be platform-specific, and should be viewable using free or widely available tools. Suitable file formats are:

For supplementary documentation:

PDF (Adobe Acrobat)

For animations:

SWF (Shockwave Flash)

For movies:

MOV (QuickTime)

For datasets:

XLS (Excel spreadsheet)

CSV (Comma separated values)

ODS (OpenOffice spreadsheets)

As for images, file names should be given in the standard file extensions. This is especially important for Macintosh users, since the Mac OS does not enforce the use of standard file extensions. Please also make sure that each additional file is a single table, figure or movie (please do not upload linked worksheets or PDF files larger than one sheet).

Author Contributions

The journal is integrated with  Contributor Role Taxonomy  (CRediT), in order to recognise individual author input within a publication, thereby ensuring professional and ethical conduct, while avoiding authorship disputes, gift / ghost authorship and similar pressing issues in academic publishing.

During manuscript submission, the submitting author is strongly recommended to select a contributor role for each of co-author, using a list of 14 predefined roles, i.e. Conceptualization, Methodology, Software, Validation, Formal analysis, Investigation, Resources, Data Curation, Writing - Original draft, Writing - Review and Editing, Visualization, Supervision, Project administration, Funding Acquisition (see  more ). Once published, the article will be including the contributor role for all authors in the article metadata.

Revising Your Article

Authors must submit the revised version of the manuscript using Track Changes/Comments tools of Word so that the Subject Editor can see the corrections and additions.

Authors must address all critiques of the referees in a response letter to the editor and submit it along with the revised manuscript through the online editorial system. In case a response letter is not submitted by the authors, the editor has the right to reject the manuscript without further evaluation. When resubmitting a manuscript that has been previously rejected with resubmission encouraged, authors must include the response letter to the article text file, and the pdf review version, so that it gets to the Subject Editor and the reviewers during the peer review.

When submitting corrections to proofs (during the layout stage), authors  must  upload the latest proof (in PDF format) containing their revisions as track changes.

Concise Copyediting Instructions

The copyediting instructions below represent a concise summary of the journal's formatting requirements. The instructions are intended for use by the authors during preparation of the final revised versions of their manuscripts, technical editors, copy editors and typesetters.  

Author names

• Omit titles, degrees, etc.
• Provide ORCID if available

Affiliation

(Department,) Institution, City, Country

Article title

Title of article: Subtitle of article

• Title: Sentence case
• Colon between title and subtitle (if any)
• No footnotes
• No bold (use when needed sub-/superscript, and/or italics only for the terms in Latin)
• Higher taxa within the title should be separated with commas and not with a semicolon

Running head

• A short version of title up to 50 characters (including spaces); normally the short title should have been suggested by the authors and checked for clarity by the copy editor
• No references to tables, figures, etc., no footnotes
• If citations unavoidable: Complete citations, allowing unambiguous identification of cited publication!
• Should be written consistently in either third or first person
• Note: The abstract has to be a stand-alone entity, to present a really well written and concise summary of the article! A special care for copy editors to check!
• Designations of nomenclatural novelties should be in bold and spelled in the way suggested ( sp. nov., gen. nov., comb. nov. )

Keywords (up to 8 words)

keyword a, keyword b, keyword n

• Do not repeat words from the title
• Listed in alphabetical order and separated by commas
• Lowercase letters, except proper names
• No bold font
• Without any punctuation marks after last keyword
• Table 2. Table caption text.
• Numbered consecutively with Arabic numerals
• Heading for every column (including the leftmost!)
• No shading of cells, rows, columns; no colored fonts
• No horizontal or vertical lines in table body
• Same number of decimal places for same statistics (usually within same column)
• Text formatting in the cell without paragraph and line break
• Table must be in an editable format (.docx, .xlsx, etc., not as images)
• Caption and footnotes as texts (not as part of a table)
• Figure 6. Figure caption text.
• Figure 1. Figure general caption text. A part caption text B part caption text N part caption text.
• If abbreviations are used, these are placed after the parts with a colon, i.e.: Abbreviations: xxxx
• If there are scale bars on the figure parts, reference to them is last and in the format: Scale bars: 20 μm ( D, N, O, Q ); 50 μm ( F, K ); 10 μm ( G, P ); 5 μm ( H ); 100 μm ( M ).
• High quality (at least 300 dpi)
• Text sharp and readable (e.g., no overlap of text and graphical elements like lines)
• White or transparent background
• No image border
• Caption as text (not as part of the image)

Capitalization

• Article title: Sentence case
• Running head: Sentence case
• For separated titles (usually H1-H3): Sentence case
• For paragraph titles (usually H4): Sentence case
• Table captions: Sentence case
• Sentence case or lower case (check for consistency only!)
• Figure captions: Sentence case
• Fig. 4; Figs 1, 2; Table 2; Appendix 1
• In text body: Titles of articles, book chapters, books, tests
• In references: Sentence case

Equations and statistical symbols

• standard typeface for Greek letters, sub-/superscripts, and abbreviations that are not variables
• italic typeface for all other statistical symbols
• Space before and after equal/inequality signs
• Same number of decimal places for decimal values
• Use leading zeros before a decimal fraction including for statistical values pertaining to probability
• Abbreviations e.g., i.e., et al., etc., cf., vs.
• Greek letter e.g., α, β, γ, δ, ε, σ, φ, χ, ω
• Scientific names of taxa of species and genera (authorities in regular font, not in italics)
• Long direct quotations
• Symbols for variables and constants, such as p , F , U , T , N , r , but not for SD (standard deviation), SE (standard error), DF (degrees of freedom), and NS (non significant). These symbols in illustrations and equations should be in italics to match the text.
• Do not use italics for emphasis
• No underlining
• Subheadings, sections and subsections
• Table 1. Table caption text.
• In systematic sections for specimen designation such us: holotype , paratype, syntype , lectotype , isotype , etc.
• NHML Natural History Museum, London
• MW Naturhistorisches Museum, Vienna
• EL length of elytra
• EW maximum width of elytra
• TL total length (PL+EL)
• In species descriptions – designation of main anatomical structures followed by a colon mark, i.e. Head: …, Thorax: …, Legs: …, Abdomen: … , etc., in this case these should be followed by a section describing other anatomical organs and structures attached to these.
• COUNTRY • specimens [e.g. 1 ♂, size ]; geographic/locality data [from largest to smallest]; coordinates; altitude/elevation/depth [using alt./m a.s.l. etc.]; date [format: 16 Jan. 1998]; collector [followed by "leg."]; other collecting data [e.g. micro habitat/host/method of collecting]; barcodes/identifiers [e.g. GenBank: MG779236]; institution code and specimen code [e.g. CBF 06023]. For Example: Holotype: CHINA • ♀; Sichuan, Kangding; 30.04°N, 101.57°E; 15.VI.2017; Yanzhou Zhang leg.; Hyp-2018-06, original number ZYZ-2017-28. Paratypes: CHINA • 1♀1♂; Sichuan, Kangding; 29.VI.2017; Yanzhou Zhang leg.; Hyp-2018-01, Hyp-2018-02, original number ZYZ-2017-08 • 1♀; Sichuan: Kangding; 2.VIII.2017; Yanzhou Zhang leg.; Hyp-2018-03, original number ZYZ-2017-20 • 1♂, Sichuan: Kangding; 29.VI.2017; Yanzhou Zhang leg.; Hyp-2018-08, original number ZYZ-2017-029.
• Punctuation: A bullet point "•" (unicode: 2022) is used to signify the beginning of a material citation. Within each citation, the different fields are delimited by a semicolon. A single field can be composed of several elements, which are separated by commas (e.g. the details region, area, town and street for the ‘locality’ field). Semicolons should not be used elsewhere in a material citation.
• Repetitive data: Authors can indicate repetitive data with indications such as " same data as for holotype ", "same data as for preceding", " same locality ", " ibid ", etc. as long as the same method and wording are used consistently throughout the paper.
• ‘Missing’ elements: It is not necessary to include information such as "no date" or "no locality data"; just list the elements that are available.
• see more details here
• Avoid quotation marks except for direct quotations, words defined by the author, and words used in unusual contexts.
• Short quotations should be embedded in the text and enclosed in double quotation marks ("). Long quotations should be on a separate line, italicized, but without quotation marks.
• Single quotation marks are to be used only for a quotation that occurs within another quotation.
• Consistent use of (-, –, —).
• In contrast to parentheses an em-dash can be used alone.
• link words such as personal names, some prefixes and compound adjectives (the last of which vary depending on the style manual in use)
• link spans.
• link numerals, sizes, dates and page numbers (e.g., 1977–1981; figs 5–7; pp. 237–258 )
• geographic or name associations (e.g., Murray–Darling River; a Federal–State agreement )
• character states combinations (e.g., long–pubescent or red–purple ).
• only for introducing a subordinate clause in the text that is often used much as we use parentheses.

Section hierarchy

• No more than 4 levels, from hierarchical level 1 (H1) to hierarchical level 4 (H4)
• Unambiguous hierarchy levels
• No numbering of hierarchical levels

Section titles

Mandatory statements.

• The author has no funding to report.
• The authors have no funding to report.
• The author has declared that no competing interests exist.
• The authors have declared that no competing interests exist.
• The author has no support to report.
• The authors have no support to report.
• Data Resources (mandatory for empirical articles)

Geographical coordinates

One of the following formats should be used:

• 36°31'21"N; 114°09'50"W
• 36°31.46'N; 114°09.84'W
• 36.5243°S; 114.1641°W
• −36.5243; −114.1641 (using minus to indicate southern and western hemispheres)

In-Text Citations

• Jackson and Miller (2012) found out that...
• A recent study (Jackson and Miller 2012) confirmed that...
• Jackson et al. (2012) found out that...
• A recent study (Jackson et al. 2012) confirmed that...
• Jackson and Miller (2012, 2015) found out that...
• Recent studies (Jackson et al. 2012, 2015) confirmed that...
• (Smith et al. 1998, 2000, 2016; Brock and Gunderson 2001; Felt 2006)
• Jackson 2008a, 2008b
• Jackson and Miller 2014a, 2014b
• Reyes-Velasco et al. 2018a, 2018b
• Jackson and Miller (2012: 120–121) found out that
• A recent study (Jackson and Miller 2012: 120) confirmed that
• Fig. 1A–D
• Figs 1–3
• Figs 1A, B, 3F, G, 7A
• Tables 1, 2
• Tables 1–3
• Appendices 1, 2
• Appendices 1–4
• All figures, tables, etc., from other sources should be written with small letters i.e.: see fig. 2 in Author (Year) ...
• de Albuquerque PRA
• Different authors separated by comma
• Year in brackets; no comma or full stop after it
• No italics (except for Latin terms)

Published papers:

Polaszek A, Alonso-Zarazaga M, Bouchet P, Brothers DJ, Evenhuis NL, Krell FT, Lyal CHC, Minelli A, Pyle RL, Robinson N, Thompson FC, van Tol J (2005) ZooBank: The open-access register for zoological taxonomy: Technical Discussion Paper. Bulletin of Zoological Nomenclature 62: 210–220.

Accepted papers:

Same as above, but ''in press'' appears instead of the year in parentheses.

Electronic journal articles:

Mallet J, Willmott K (2002) Taxonomy: Renaissance or Tower of Babel? Trends in Ecology and Evolution 18(2): 57–59. https://doi.org/10.1016/S0169-5347(02)00061-7

Paper within conference proceedings:

Orr AG (2006) Odonata in Bornean tropical rain forest formations: Diversity, endemicity and applications for conservation management. In: Cordero Rivera A (Ed.) Forest and Dragonflies. Fourth WDA International Symposium of Odonatology, Pontevedra (Spain), July 2005. Pensoft Publishers, Sofia-Moscow, 51–78.

Book chapters:

Mayr E (2000) The biological species concept. In: Wheeler QD, Meier R (Eds) Species concepts and phylogenetic theory: A debate. Columbia University Press, New York, 17–29.

Goix N, Klimaszewski J (2007) Catalogue of Aleocharine Rove Beetles of Canada and Alaska. Pensoft Publishers, Sofia-Moscow, 166 pp.

Book with institutional author:

ICZN [International Commission on Zoological Nomenclature] (1999) International code of zoological nomenclature. Fourth Edition. The International Trust for Zoological Nomenclature,  London.

PhD thesis:

Dalebout ML (2002) Species identity, genetic diversity and molecular systematic relationships among the Ziphiidae (beaked whales). PhD Thesis, University of Auckland, Auckland, ## pp.

BBC News (2012) Island leopard deemed new species http://news.bbc.co.uk/ [Accessed on dd.mm.yyyy]

Submission Guidelines

Submission procedure.

Already have a Username/Password?

Go to Login

Need a Username/Password?

Go to Registration

Registration and login are required to submit items online and to check the status of current submissions.

Submission of manuscripts to this journal is possible only through the online submission module. We kindly request authors to consult the Focus and Scope section prior to submission. In order to submit a manuscript to the journal, authors are required to register with the journal and/or to login. Once logged in, you will find the online submission system either by clicking the " Submit manuscript " button.

The manuscript submission process is separated into the following steps:

• Step 1:  Specifying the manuscript type and completing the submission checklist
• Step 2:  Choosing the payment option and requesting optional services
• Step 3:  Typing in the author(s) names and affiliation, title, abstract, keywords, and other metadata
• Step 4:  Assigning classifications categories for your manuscript using hierarchical classification trees
• Step 5:  Completing the submission metadata by adding details about any supporting agencies, conflict of interest, ethical statement, comments to the editors
• Step 6:  Agreeing with the journal's Data Publishing Policy and specifying the availability of the data underpinning your article
• Step 7:  Uploading the submission file and the additional files (see below for details on how to prepare it)
• Step 8:  Confirming the automatically generated pdf review version of the article, and the metadata (or revising them, if needed)
• Step 9:  Uploading supplementary files (see below for details) and associated metadata
• Step 10:  Suggesting reviewers, final verification of the submitted files and confirmation

Stepwise guidance on new manuscript submission, with screenshots of the interface embedded, is available online in  this section of the User Manual .

Organizing Your Submission

Before starting your submission please make sure that your manuscript is formatted in accordance with the Author Guidelines.

Before attempting an online submission, please consider preparing the following file types:

1. Submission file

Review  version of the manuscript: a TEXT (MS WORD) file in either DOC, DOCX, RTF or ODT format. The total file size must be no larger than 80 MB . The system allows two options for the submission file upload:

it could contain all figures embedded at their respective places within the manuscript: 

Advantage: The review version of the manuscript will be more convenient for reading and understanding by the reviewers and editors. Likewise, if you opt to post your manuscript on ARPHA Preprints and this is allowed by the current journal’s policies, it will be better organised for the readers.

Drawback: Additional effort is needed to place and number the figures within the text.

it could contain the article text only, while the figures are added separately in the allowed formats (see below), so that the system can add them automatically to the PDF version that will be sent for review. The authors have the option to check and replace, if needed, the PDF review version generated at the first submission step:

Advantage: No additional effort is needed for placing and numbering the figures within the text.

Drawback: All figures will be placed at the end of the manuscript and the review version will be less convenient for reading and understanding by the reviewers and editors. The same concerns your preprint if you decide to post it on ARPHA Preprints and this is allowed by the current journal’s policies.

2. Additional files

High-resolution figures must be submitted during the same submission process as the additional files (Step 7) in one of the accepted file formats (see below). These may be compressed in order to reduce bandwidth during upload:

• Figures (each figure as an individual file in one of the following image file formats:  EPS, TIFF, JPEG, PNG, GIF, BMP, not larger than 20 MB each )
• Equations (each equation as an individual file in one of the above-mentioned image file formats)

Please note that the  maximum file size  that may be uploaded through our online submission system is  20 MB .

3. Supplementary files (appendices)

Large datasets or multimedia files, usually published as appendices in conventional print journals, should be uploaded as  supplementary files  complete with the associated metadata on the online submission form. Supplementary files should have their own legends.

Most file formats are accepted. Text-only appendices must be in DOC, DOCX, RTF, or ODF formats.

Should you have any technical problems in submitting a manuscript to this journal, please contact the  Editorial Office  at  [email protected] .

We encourage authors to send an inquiry to the respective Subject Editor prior to submitting a manuscript. The purpose of the presubmission inquiry is to solicit rapid initial feedback on the suitability of the manuscript for publication in this journal. Pre-submission inquiries may also be sent to the Editorial Office at  [email protected] .

Special issues & Topical collections

Special issues and Topical collections are collections of articles grouped together by a shared topic or interest group, such as an emerging area of research, proceedings of a conference, outcomes of a research project or a Festschrift volume.

Article collections aim to aid the dissemination and outreach of multiple research outcomes and also bring together research teams from around the globe working on similar topics, thus increasing the opportunities for collaboration, sharing and re-use of research. Article collections bring credit, increased discoverability, visibility and recognition to both their collection editors and participating authors.

Special issues are available only in journals published in consequent issues within an yearly volume and are subject to a submission deadline set at the time the call for papers is issued. The publication date of a Special issue is also pre-scheduled and all papers are published simultaneously on the same date as a separately numbered issue within the yearly journal's volume. This means that the Special issue will be published only when all articles are ready to be published. 

Topical collections can be opened in any journal hosted on ARPHA and can be permanent or made subject to a submission deadline. It is only up to the Collection editor(s) to decide whether and when the collection is to be closed for submission (given a timely public announcement is provided). The articles are published on a rolling basis, as soon as they are ready for publication, and can be part of different journal issues, published across many years.

Article collections are managed by a Collection editor and associated Guest editors. To pitch a Special issue or a Topical collection, either contact the Editor-in-Chief or submit an Open an article collection proposal form. Before pitching a Special issue or a Topical collection, please make yourself aware of the specificity of the focus, scope and policies of the journal and the associated responsibilities and benefits for you as a Collection editor.

How It Works

The following guidelines apply for both Special issues and Topical collections in the ARPHA journals.

Article collections can be opened in any of the ARPHA-hosted journals. It is subject to the journal's policy, however, to offer this feature or not. 

Special issues are available only in the journals that publish consecutively numbered issues within a yearly volume, while topical collections are available to all ARPHA-hosted journals, depending on their policies.

Collections may have subcollections, for example, topical subcollections. A subcollection cannot be managed separately from the parent collection, except in the case of conference proceedings submission workflows available at some ARPHA-hosted journals.

Opening and managing a collection

The article collections are managed by a Collection editor and Guest editors. The Collection editor is responsible for approving or declining manuscripts submitted to the article collection; assigning a Guest editor to each manuscript for handling the peer review process; and managing the collection on the journal’s website (e.g. change the collection’s description or the order of the papers). The Collection editor has the full rights of a Guest editor and can also handle manuscripts.

Before pitching a collection, assure that you are ready to appoint other Guest editors, if necessary. The Collection editor and the Guest editors are also expected to commission an initial set of manuscripts to be submitted soon after the opening of the collection.

Submit the Article collection proposal form or contact the Editor-in-Chief via email. The collections should fully comply with the journal’s focus, scope and editorial policies.

Online proposals are forwarded to the Editor-in-Chief and to the journal’s editorial office for approval. The editorial office checks and confirms the guest editors' credentials.

Upon approval of the proposal, the journal’s editorial office will set up the collection on the journal’s website.

Open collections will be promoted through the journal’s website and social media in collaboration with the Collection and Guest editors. 

Editors of Special issues need to assure that the minimum volume of articles is met within the set deadline and that, if necessary, a deadline extension is announced well in advance.

Editors of Topical collections with no set submission deadlines need to inform the journal’s editorial office if they wish to close the collection for submissions in a timely manner. 

Authors opt for assigning their manuscript to a collection during submission. In case the manuscript is declined from the collection, it undergoes the regular evaluation and peer review process at the journal.

Once the manuscript passes the initial pre-review screening by the Editor-in-Chief and the journal's editorial office, it is forwarded to the Collection editor to either approve or decline it for the collection. The Collection editor is notified about each new submission to the collection via email sent by the system.

After reading the paper, the Collection editor can:

accept it in the collection and assign it to a Guest editor.

decline it from the collection and send it back to the journal's editorial office.

Once a manuscript is assigned to a Guest editor, he or she takes on the responsibility to invite reviewers and provide an editorial decision for revision, rejection or acceptance of the manuscript, based on the reviews and personal evaluation. Papers submitted by the guest editor(s) must be handled under an independent review process and make up no more than 25% of the collection's total.

The editorial decisions are automatically forwarded to the authors by the system.

The guest editors are overseen by the journal's Editor-in-Chief and/or dedicated board members, and may intervene in the editorial process. Depending on the journal’s policy, the journal’s Editor-in-Chief might need to approve the Guest editor’s final decision before the manuscript is accepted for publication.

Benefits of Editing a Collection

The main advantages to open and edit an article collection can be summarised as follows:

Credit and recognition for the Collection and Guest editors who take care to organise and manage the article collection.

Facilitates discoverability and usability of topically related studies, which in turn benefits both authors and readers.

Increases the visibility of related papers, even when papers might otherwise lack in viewership. 

Prompts simultaneous citation of multiple articles related to a certain subject.

Facilitates citation and referencing of the whole issue as a complete entity.

Editor’s Responsibilities

By proposing an article collection (Special issue or Topical collection), you agree to act as a Collection editor, whose main responsibilities are:

Working with the editorial office to set up the article collection on the journal’s website.

Appoint Guest editors for the article collection.

Approve or decline each manuscript submitted to the article collection.

Assign a Guest editor for each manuscript submitted to the article collection.

Assure that the article collections complies with any relevant requirements, as set up by the journal and the agreement (if any).

Inform the journal’s editorial office about any changes or issues concerning the management of the collection in due time.

You will also be granted the user rights of a Guest editor necessary to handle manuscripts in the system (i.e. assign reviewers and provide an editorial decision on the acceptance/rejection of the manuscript). 

The responsibilities of a Guest editor are:

Handling the peer review of the manuscripts they have been assigned to.

Making an editorial decision for revision, acceptance or rejection of the manuscripts they have been assigned to, based on the reviews provided and personal evaluation.

Taking into consideration the recommendations of the journal’s Editor-in-Chief.

The journal adheres to the " Best practices for guest edited collections " by the Committee on Publication Ethics (COPE). The collection editors should first familiarise themselves with those guidelines before applying for or starting working on a guest edited article collection.

For more information about the editorial workflow, visit How it works ?

Article Processing Charges

Publisher's statement

A key policy and strategic aim of Pensoft is to provide high-quality and inclusive publishing services at highly competitive and affordable Article Processing Charges (APCs) or for free through its diamond open access journals. See Pensoft’s journal portfolio here .

In order to ensure long-term sustainability of the journals and cover the cost of the associated in-house publishing services, our journals require Article Processing Charges (APCs). These charges apply only after a submitted manuscript is accepted for publication, and may be partially or fully covered by institutional funds to reduce financial burdens on authors of research.

Pensoft strongly supports measures that ensure an inclusive and FAIR publishing environment, which in turn prompts quality, sustainability and reasonable pricing in scholarly publishing. You can find more about the publisher’s view on quality, transparency, openness and equity in scholarly publishing in Pensoft’s official statement , prompted by the publication of the European Union’s Conclusions on high-quality, transparent, open and equitable scholarly publishing . 

In compliance with the Plan S requirements , Pensoft provides a breakdown of the APC following the  guidelines by the Fair Open Access Alliance (FOAA) . The report on the journal’s APC is submitted on a yearly basis to the  Journal Comparison Service by Coalition S and the detailed breakdown is available to the participating funding institutions on the platform.

Authors who are unable to pay their APCs for several reasons, should consult the Journal’s Discounts and Waivers page, use the diamond open-access journals (free to publish and free to read) hosted on Pensoft’s ARPHA Publishing Platform , or contact the journal’s Editor-in-Chief directly. 

Core Charges

Core services included in our Article Processing Charges

• Online submission and editorial management system, professional peer review and editorial assistance.
• Personal attitude, technical support and fast reply to any inquiry coming from authors, editors or reviewers.
• Automated email notification and alert system to save you time from tracking the progress of your manuscript.
• Automated registration of peer reviews at Clarivate (formerly Publons).
• Copy-editing, technical editing, typesetting and proofreading services.
• Publication in 3 digital formats: semantically enhanced HTML, PDF and machine-readable JATS XML.
• Rapid publication process, normally within 1-2 weeks time after a manuscript is accepted for publication.
• Full-color (no extra-charges for color), high-resolution hardcopy of reprints or whole issues.
• Advanced data publishing workflows. 
• Semantic Web enhancements to the article text. 
• Markup and visualization of all biological taxon names and taxon treatments in your work, if present.
• Immediate free access to the article on the day of publication.
• Copyright retained by the authors, articles distributed under the Creative Commons Attribution (CC-BY) 4.0 license.
• Active dissemination and promotion through social bookmarking tools and social media.
• Automated email acknowledgements to editors and reviewers upon publication.
• Automated alert service through email and RSS on the day of publication. 
• Registration of all new taxa in ZooBank, IPNI, MycoBank or Index Fungorum (where relevant).
• Export and display of taxon treatments to Encyclopedia of Life (EOL), Plazi, Species-ID, Globalnames, and other aggregators (where relevant).
• Immediate distribution of your publication to scientific databases, indices and search engines (Web of Science, Scopus, Google Scholar, CAB Abstracts, DOAJ Content and others).
• Archiving in international repositories (PubMedCentral, CLOCKSS, Zenodo).
• Bibliography search and discovery tool. 
• Citation export in various formats.
• Cited-by records statistics and display.
• Article- and sub-article-level metrics (Altmetric, Dimensions, number of downloads separately for the PDF, XML and HTML, usage stats for figures, tables and supplementary files).

Please note that the charges below are applicable for all manuscripts submitted after  26th of January 2024 . 

*At least one of the authors must be a member of ISH in good standing for the reduced fee to be applicable.

The above prices do not include VAT (Value Added Tax). VAT is applicable only for VAT NON-registered customers based within the European union. To avoid charging VAT, the EU companies or persons should provide their VAT registration numbers validated with the EU taxation database ( https://ec.europa.eu/taxation_customs/vies/ ).

Please note that the charges below are applicable for all manuscripts submitted before  25th of January 2024 . 

Publication fees in open access journals cover article processing costs associated with editorial process, layout, publication and dissemination, and ensure a free distribution of your paper at no charge for the readers. The authors submitting manuscripts to this journal benefit from:

• Online submission and editorial management system, professional review and editorial assistance
• Typesetting, proofreading and publication
• No limit in manuscript length; large revisionary works, checklists, catalogues, etc. are published as separate monographs
• Rapid publication process, usually within 1-2 weeks time after a manuscript is accepted for publication
• Advanced publishing technologies, possibilities for data publication and various semantic Web enhancements
• Immediate free access for everyone to your work
• Immediate Alert Service through Email, RSS feeds and social networks (Twitter, Facebook, mendeley and others)
• Immediate distribution of your publication to scientific databases, indices and search engines (Web of Science, Google Scholar, CAB Abstracts, and others)
• Archiving of your publication, electronically and in print, in trusted (e-) archives and copyright libraries
• Author copyright and distribution under the Creative Commons Attribution 3.0 license
• No extra charges for colour illustrations
• Full-color, high-resolution print version

Offprints and printed issues of the journal can be purchased additionally (see Printed Version & Reprints).

Discounts and Waivers

Discounts and waivers for this journal can be provided only by the  International Society of Hymenopterists

Journal of Hymenoptera Research is published in identical print (high-resolution, full-color) and online (PDF) versions.

Printed versions of this journal may be ordered in parts or subscribed for. To subscribe please contact us by writing to  [email protected] .

Please include the full delivery address and indicate your preferred payment method. Please contact us if you need a quotation or proforma invoice.

Separate issues or reprints (high-resolution, full-color) can be ordered using the "Order now" button available under each issue or article on the journal's website.

Prices are given in EURO and are exclusive of postage and handling. Payment in USD is also possible according to the exchange rate on the day of payment.

IMPORTANT: Our prices do not include VAT. Orders from countries outside the European Union (EU) or from VAT-registered EU customers will be processed VAT-free. VAT (20%) will be added ONLY to NOT VAT-registered customers based in the European Union.

Prices of full-color, high-resolution printed version (separate article and complete issues)

Additional Services (Optional)

*This service can be discounted or waived for articles of outstanding importance for the science and society.

Data Publishing Guidelines

We strongly encourage and support various strategies and methods for data publication. The preferable way is to store data in internationally recognised data repositories and link back to the data set(s) in the respective article. Data can also be published as supplementary files to the articles, however this should be an exception rather than a rule (see How to publish data ). The key to discover, use and cite your data is to include the data references in the reference lists of the articles and  always  include the DOIs of the data sets, when available, in the citation record . You may read more about this in  How to cite data  section of the article below. A good example of concise data citation guidelines using DOIs is also available on the  GBIF website  and on other data repositories.

Darwin Core-structured species occurrence records and observations (primary biodiversity data) should be published with GBIF using either the Integrated Publishing Toolkit (IPT) (for which Pensoft maintains an instance, in case such is not available to the authors).  Alternatively, DwC data could also be published in trusted and community-recognised repositories (for example, Atlas of Living Australia, Symbiota,  Arctos or others), however deposition at GBIF should always have a priority over the alternatives. In case a dataset is deposited in more than one repository, the data paper should link to the dataset which is actually described, again with GBIF having a priority over the others.

Authors who want to publish species occurrence data as supplementary files only or through generic repositories (e.g. Zenodo, Dryad), instead of submitting these to GBIF, should justify their decision to do so in a letter to the editors.

For biodiversity and biodiversity-related data the reader may consult the  Strategies and guidelines for scholarly publishing of biodiversity data   (Penev et al. 2017,   Research Ideas and Outcomes 3: e12431.  https://doi.org/10.3897/rio.3.e12431 ).   For reader's convenience, we list here the hyperlinked table of contents of these extensive guidelines:

• Introduction
• What is a Dataset
• Why Publish Data
• How to Publish Data
• How to Cite Data
• General Policies for Biodiversity Data
• Data Publishing Licenses
• General Information
• Taxonomic Data  
• Species-by-Occurrence and Sample-Based Data
• Phylogenies
• Gene Sequence
• Protein Sequence
• Other Omics
• Various Data Types
• Biodiversity Literature
• Data Published within Supplementary Information Files
• Import of Darwin Core Specimen Records into Manuscripts
• Data Published in Data Papers
• Data Papers Describing Primary Biodiversity Data
• Data Papers Describing Ecological and Environmental Data
• Data Papers Describing Genomic Data
• Software Description Papers
• Quality of the Manuscript
• Quality of the Data
• Consistency between Manuscript and Data

The core of the data publishing project of Pensoft is the concept of "Data Paper" developed in a cooperation with the Global Biodiversity Information Facility (GBIF). Data Papers are peer-reviewed scholarly publications that describe the published datasets and provide an opportunity to data authors to receive the academic credit for their efforts. Currently, Pensoft offers the opportunity to publish Data Papers describing occurrence data and checklists, Barcode-of-Life genome data and biodiversity-related software tools, such as interactive keys and others.

Examples of data papers

ZooKeys: Antarctic, Sub-Antarctic and cold temperate echinoid database A dataset from bottom trawl survey around Taiwan Project Description: DNA Barcodes of Bird Species in the National Museum of Natural History, Smithsonian Institution, USA Literature based species occurrence data of birds of northeast India MOSCHweb — a matrix-based interactive key to the genera of the Palaearctic Tachinidae (Insecta, Diptera) Amundsen Sea Mollusca from the BIOPEARL II expedition Iberian Odonata distribution: data of the BOS Arthropod Collection (University of Oviedo, Spain FORMIDABEL: The Belgian Ants Database Circumpolar dataset of sequenced specimens of Promachocrinus kerguelensis (Echinodermata, Crinoidea)

PhytoKeys: Florabank1: a grid-based database on vascular plant distribution in the northern part of Belgium (Flanders and the Brussels Capital region) Database of Vascular Plants of Canada (VASCAN): a community contributed taxonomic checklist of all vascular plants of Canada, Saint Pierre and Miquelon, and Greenland Herbarium of Vascular Plants Collection of the University of Extremadura (Spain)

Nature Conservation: Antarctic macrobenthic communities: A compilation of circumpolar information

Press releases on data papers New incentive for biodiversity data publishing Dat a publishing policies and guidelines for biodiversity data by Pensoft First database-derived 'data paper' published in journal A new type of data papers designed to publish online interactive keys Data paper describes Antarctic biodiversity data gathered by 90 expeditions since 1956 Unique information on Belgian ants compiled and published through FORMIDABEL data paper Database simplifies finding Canadian plant names and distribution A synthesis of the 36451 specimens from the UNEX Herbarium in a new data paper

Data Quality Checklist and Recommendations

INTRODUCTION

An empowering aspect of digital data is that they can be merged, reformatted and reused for new, imaginative uses that are more than the sum of their parts. However, this is only possible if data are well curated. To help authors avoid some common mistakes we have created this document to highlight those aspects of data that should be checked before publication.

By "mistakes" we do not mean errors of fact, although these should also be avoided! It is possible to have entirely correct digital data that are low-quality because they are badly structured or formatted, and, therefore, hard or impossible to move from one digital application to another. The next reader of your digital data is likely to be a computer program, not a human. It is essential that your data are structured and formatted so that they are easily processed by that program, and by other programs in the pipeline between you and the next human user of your data.

The following list of recommendations will help you maximise the re-usability of your digital data. Each represents a test carried out by Pensoft when auditing a digital dataset at the request of an author. Following the list, we provide explanations and examples of each recommendation.

Authors are encouraged to perform these checks themselves prior to data publication. For text data, a good text editor ( https://en.wikipedia.org/wiki/List_of_text_editors ) can be used to find and correct most problems. Spreadsheets usually have some functions for text checking functions, e.g. the "TRIM" function that removes unneeded whitespace from a data item. The most powerful text-checking tools are on the command line, and the website "A Data Cleaner's Cookbook" ( https://www.datafix.com.au/cookbook/ ) is recommended for authors who can use a BASH shell.

When auditing datasets for authors, Pensoft does not check taxonomic or bibliographic details for correctness, but we will do basic geochecks upon request, e.g. test to see if the stated locality is actually at or near the stated latitude/longitude. We also recommend checking that fields do not show "domain schizophrenia", i.e. fields misused to containing data of more than one type.

Proofreading data takes at least as much time and skill as proofreading text. Just as with text, mistakes easily creep into data files unless the files are carefully checked. To avoid the embarrassment of publishing data with such mistakes, we strongly recommend that you take the time to run these basic tests on your data.

• The dataset is UTF-8 encoded
• The only characters used that are not numbers, letters or standard punctuation, are tabs and whitespaces
• Each character has only one encoding in the dataset
• No line breaks within data items
• No field-separating character within data items (tab-separated data preferred)
• No "?" or replacement characters in place of valid characters
• No Windows carriage returns
• No leading, trailing, duplicated or unnecessary whitespaces in individual data items
• No broken records, i.e. records with too few or too many fields
• No blank records
• No duplicate records (as defined by context)
• No empty fields
• No evident truncation of data items
• No unmatched braces within data items
• No data items with values that are evidently invalid or inappropriate for the given field
• Repeated data items are consistently formatted
• Standard data items such as dates and latitude/longitude are consistently formatted
• No evident disagreement between fields
• No unexpectedly missing data

RECOMMENDATIONS

Characters  

Computer programs do not "read" characters like "A" and "4". Instead, they read strings of 0's and 1's and interpret these strings as characters according to an encoding scheme. The most universal encoding scheme is called UTF-8 and is based on the character set called Unicode. Text data should always be shared with UTF-8 encoding, as errors can be generated when non-UTF-8 encodings (such as Windows-1252) are read by a program expecting UTF-8, and vice-versa. (See also below, on replacement characters).  

• The only characters used that are not numbers, letters or standard punctuation are tabs and whitespaces

Unusual characters sometimes appear in datasets, especially when databases have been merged. These "control" or "gremlin" characters are sometimes invisible when data are viewed within a particular application (such as a spreadsheet or a database browser) but can usually be revealed when the data are displayed in a text editor. Examples include vertical tab, soft hyphen, non-breaking space and various ASCII control characters ( https://en.wikipedia.org/wiki/Control_character ).

We have seen individual datasets in which the degree symbol (°) is represented in three different ways, and in which a single quotation mark (') is also represented as a prime symbol, a right single quotation mark and a grave accent. Always use one form of each character, and preferably the simplest form, e.g. plain quotes rather than curly quotes.

Spreadsheet and database programs often allow users to have more than one line of text within a data item, separated by linebreaks or carriage returns. When these records are processed, many computer programs understand the embedded linebreak as the end of a record, so that the record is processed as several incomplete records:

item A  itemB1          itemC

               itemB2

itemA           itemB1

itemB2          itemC

Data are most often compiled in table form, with a particular character used to separate one field ("column") from the next. Depending on the computer program used, the field-separating character might be a comma (CSV files), a tab (TSV files), a semicolon, a pipe (|) etc.

Well-structured data keeps the field-separating character out of data items, to avoid confusion in processing. Because commas are commonly present within data items, and because not all programs understand how to process CSVs, we recommend using tabs as field-separating characters (and avoiding tabs within data items!): https://en.wikipedia.org/wiki/Tab-separated_values .

When text data are moved between different character encodings, certain characters can be lost because the receiving program does not understand what the sending program is referring to. In most cases, the lost character is then represented by a question mark, as in "Duméril" becoming "Dum?ril", or by a replacement character, usually a dark polygon with a white question mark inside.

It is important to check for these replacements before publishing data, especially if you converted your data to UTF-8 encoding from another encoding.

On UNIX, Linux and Mac computers, a linebreak is built with just one character, the UNIX linefeed '\n' ('LF'). On Windows computers, a linebreak is created using two characters, one after the other: '\r\n' ('CRLF'), where '\r' is called a 'carriage return' ('CR'). Carriage returns are not necessary in digital data and can cause problems in data processing on non-Windows computers. Check the documentation of the program in which you are compiling data to learn how to remove Windows carriage returns.

Like "control" and "gremlin" characters, whitespaces are invisible and we pay little attention to them when reading a line of text. Computer programs, however, see whitespaces as characters with the same importance as "A" and "4". For this reason, the following four lines are different and should be edited to make them the same:

Aus bus (Smith, 1900)

   Aus bus (Smith, 1900)

Aus bus (Smith,   1900)

Aus  bus   (Smith, 1900  )

  Records

If a data table contains records with, for example, 25 fields, then every record in the table should have exactly 25 data items, even if those items are empty. Records with too few fields are often the result of a linebreak or field separator within a data item (see above). Records with too many fields also sometimes appear when part of a record has been moved in a spreadsheet past the end of the table.

Blank records contribute nothing to a data table because they contain no information, and a tidy data table has no blank lines. Note, however, that a computer program looking for blank lines may not find what looks to a human like a blank line, because the "blank" line actually contains invisible tabs or whitespaces.

It can be difficult to find duplicate records in some datasets, but our experience is that they are not uncommon. One cause of duplicates is database software assigning a unique ID number to the same line of data more than once. Context will determine whether one record is a duplicate of another, and data compilers are best qualified to look for them.

  Fields

  Fields containing no data items do not add anything to the information content of a dataset and should be omitted.

•   No evident truncation of data items

The end of a data item is sometimes cut off, for example when a data item with 55 characters is entered into a database field with a 50-character maximum limit. Truncated data items should be repaired when found, e.g.

Smith & Jones in Smith, Jones and Bro

repaired to :

Smith & Jones in Smith, Jones and Brown, 1974

These are surprisingly common in datasets and are either data entry errors or truncations, e.g.

Smith, A. (1900 A new species of Aus. Zool. Anz. 23: 660-667.

5 km W of Traralgon (Vic

For example, a field labelled "Year" and containing years should not contain the data item "3 males".

•   Repeated data items are consistently formatted

The same data item should not vary in format within a single dataset, e.g.

Smith, A. (1900) A new species of Aus. Zool. Anz. 23: 660-667.

Smith, A. 1900. A new species of Aus. Zoologischer Anzeiger 23: 660-667.

Smith, A. (1900) A new species of Aus. Zool. Anz. 23, 660-667, pl. ix.

Data compilers have a number of choices when formatting standard data items, but whichever format is chosen, it should be used consistently. A single date field should not, for example, have dates represented as 2005-05-17, May 19, 2005 and 23.v.2005.

If there are fields which contain linked information then these fields should be checked to ensure that they do not conflict with each other. For example, the year or an observation cannot be after the year it was published.  Examples:

Year            Citation

1968            Smith, A. (1966) Polychaete anatomy. Academic Press, New York; 396 pp.

Genus           Subgenus

Aus             Bus (Aus)

This is a rare issue in datasets that have been audited, but occasionally occurs. An example is the Darwin Core "verbatimLocality" field for a record containing a full latitude and longitude, but with the "decimalLatitude" and "decimalLongitude" fields blank.

• Spelling of Darwin Core terms

Darwin Core terms are usually considered case sensitive, therefore you should use their correct spelling ( http://rs.tdwg.org/dwc/ ).

We thank Dr. Robert Mesibov for preparing the Data Quality Checklist draft and Dr. Quentin Groom for reviewing it.

Dryad Repository Submissions

This journal is integrated with the Dryad Digital Repository to make data publication simple and easy for authors. There is a $150 Data Publishing Charge for Dryad submissions, payable via the Dryad website.  For more information, please see their FAQ .

Guidelines for Editors

How to access a manuscript.

Manuscripts can be accessed after login

• Login is possible after registration at the journal's website. Our Editorial Office will register all first-time editors and reviewers. New users will receive an automated notification with a request to confirm registration and account information, and options for setting a password, email alerts and other features.   Note:  All users can use their registration details to login in all three (Book, E-Book and the respective Journal) platforms of  www.pensoft.net . Note:  Please remember that you may have registered with two or more different email addresses, that is why you may have more than one valid account at  www.pensoft.net . We advise using only one email address, hence one password associated with it, for all your operations at  www.pensoft.net . We highly recommend  that in case the user has two or more different accounts, to merge these through the user's profile.

Note:  The users can at any time change the initially set password and correct personal details using their user's profile menu (by clicking on the user's name in the upper right corner of the screen appearing after login).

If you have forgotten your password, please use the function  Forgot your password? or write to request it from  [email protected] .

There are two ways to access a manuscript

After login, please go to the respective journal’s web page and click on My Tasks button in the upper right corner of the screen. This way, you will be able to see all manuscripts you are responsible for as an author or reviewer or editor.

Note: The manuscripts are grouped by categories, e.g., In Review (no.), In layout (no.), Published (no.), and Archived (no.) etc. The number in brackets after each category shows the number of manuscripts that were assigned to you.

Click on the active manuscript link provided in the email notification you have received from the online editorial system. The link will lead you directly to the manuscript.

General Responsibilities of Editors

Subject, or Associate, editors in Pensoft’s journals carry the main responsibility for the scientific quality of the papers. They take the final decision on a manuscript’s acceptance or rejection and their names are listed as Academic Editor  in the header of each published article.

The editorial process is facilitated through an online editorial system and a set of email notifications. The online editorial system informs the Subject Editor about any change in the status of a manuscript from submission to publication.

The online editorial system is designed to save time and effort for Subject Editors in checking the status of the manuscripts. There is no need for editors to visit the journal’s website to keep track on the manuscript they are responsible for. The online system will inform the Subject Editor when an invited reviewer has accepted or declined to review. The email notifications contain stepwise instructions what action is needed at each stage, as well as a link to the respective manuscript (accessible by clicking on the link in the email notification or after login – see How to Access a Manuscript ).

Subject Editors are not expected to provide a thorough linguistic editing or copyediting of a manuscript, but rather focus on its scientific quality and overall style, which should correspond to good practices in clear and concise academic writing. It is the author’s responsibility to submit the manuscript in linguistically and grammatically correct English. The Subject Editor should not hesitate to recommend either Reject , or Reject, but resubmission encouraged  PRIOR to the peer-review process, in cases when a manuscript is scientifically poor and/or does not conform to journal’s style, and/or is written in poor English (see Note under point 1 below how to reject a manuscript prior to peer review). 

Editors-in-Chief, Managing Editors or their deputies are allowed to publish a limited proportion of papers per year co-authored by them, after considering some extra precautions to avoid an impression of impropriety, endogeny, conflicts of interest and ensure that the editorial decision-making process is transparent and fair.

It often happens that even carefully written manuscripts may contain small errors in orthography or stylistics. We shall be thankful if editors spot such errors during the reading process and correct them.

Stepwise Description of the Editorial Process

• Once a manuscript is submitted, the Managing Editor (or the Editor-in-Chief) briefly checks if the manuscript conforms with the journal's Focus, Scope, Policies and style requirements and decides whether it is potentially suitable for publication and can be processed for review, or rejected immediately, or returned to the author for improvement and re-submission. Note: There are two ways to reject/return a manuscript prior to review process: - Through the buttons Reject  or Return to the author for correction  in the Editorial tab. Please note, however, that the buttons will be made active only after a justification for the rejection or return is provided in the text field. - Through an email to the Editorial office explaining the reason for rejection or return. The manuscript will be then rejected/returned through the online editorial system and the respective notification email will be sent from the Editorial Office.
• At this stage, the Managing Editor (or the Editor-in-Chief) can also check the manuscript for plagiarism via the iThenticate service by clicking on the "ïTehnticate report" button. Journals providing a peer review in languages other than English (for example Russian) may use other plagiarsim checking services (for example Antiplagiat).
• When a manuscript is suitable, the Managing Editor (or the Editor-in-Chief) assigns it to the Subject Editor responsible for the respective topic (e.g., science branch or taxon). The Subject Editor receives a notification email on the assignment. Note: The link to the respective manuscript is available in the editorial assignment email and all consequent reminder emails. The manuscript is accessible by clicking on the link in the email notifications, or via the user's dashboard after login. Please see How to Access a Manuscript above in case you have any difficulties.
• The assigned Subject Editor next reads the manuscript to decide whether it is potentially suitable for publication and can be processed for review, or rejected immediately, or returned to the author for improvement and re-submission. Reasons for rejection can be a low scientific quality, non-conformance to the journal’s style/policies, and/or linguistically or grammatically poor English language. Note: There are two ways to reject a manuscript prior to review process: -  Through the buttons  Reject or Reject, but resubmission encouraged  in the Editorial tab. Please note, however, that the buttons become active only after a justification for the rejection is provided in the text field.  -  Through an email to the Editorial office explaining the reason for rejection. The manuscript will be then rejected/returned through the online editorial system and the respective notification email will be sent from the Editorial Office.
• In case the manuscript is acceptable for peer review, the Subject Editor has to invite reviewers by clicking on the Invite reviewers  link. The Subject Editor can select from a list of reviewers, starting with the ones suggested by the authors during the submission process, and followed by the reviewers who are already listed in the database, or add new reviewers.
• Once reviewers are chosen, the Subject Editor has to click the Invite reviewers green button at the end of the page which will generate email templates with review invitations. It is highly recommended that the Subject Editor adds some personal words above the standard email text of the review invitation.
• In case a reviewer is absent from our users' data base, the Subject Editor can add his/her name and email through the Add new reviewer  link, which will appear once the search field reveal no results. It is possible that the needed reviewer has already been registered in the Pensoft database either as customer or author/reviewer of another journal. If this is the case, then his/her name, affiliation and other metadata will automatically appear once the e-mail field is populated in the Create user online form.
• The Subject Editor receives a notification email when the Reviewer agrees or declines to review. The Subject Editor takes care to appoint additional reviewers in case some of the invited reviewers decline.
• Once all Reviewers submit their reviews, the Subject Editor receives an email notification, inviting him/her to consider Reviewers' opinions, read through the manuscript and take a decision through the Proceed  button. Note: Editorial comments can be added in the online editorial form; comments and corrections are expected to be added also in the manuscript file (either on the PDF version or in the text file), that should be uploaded during finalization of the editorial decision process. 
• At this stage, the editor should take a decision either to (1) accept the manuscript, (2) reject it, (3) recommend Major or Minor Revisions or reject it, or open a second review round. In case the manuscript is not rejected, but recommended for Minor Revision, Major Revision, or Acceptance, the author is expected to submit a revised version within a certain period of time (and the Subject Editor will be notified by email about that). Note 1: Authors must submit revised versions as a text file using Track Changes/Comments tools of Word so that the Subject Editor can see their corrections/additions. Authors must reply to the essential critiques and comments of reviewers separately through the online editorial system. Note 2: During the second, or next, review round, the Subject Editor may decide to ask reviewers to evaluate the revised version of the manuscript. He/she may also make a decision based on the author’s responses and the revised version of the manuscript without asking additional Reviewers' support.
• After acceptance, the manuscript will go to proofreading and layout . The Subject Editor will be notified by email when the final proof is uploaded on the journal’s website. The Subject Editor is expected to look at the proofs and notify the Editorial Office through email in case the proofs need improvement.
• The Subject Editor may always access information on the manuscripts which have been edited by him/her through the menu My Tasks –>  Subject Editor  on the journal’s web page – In Review (no.), In Edit (no.), Published (no.), and Archived (no.). The number in brackets after each category shows the number of manuscripts that were assigned.

Editors’ and Reviewers’ Workload Stats

While selecting a Reviewer or a Subject Editor to assign to a manuscript, Editors can access the current and past workload for the person they are considering.

By clicking on the user’s name, an Editor sees how many editorial or review tasks the person is currently assigned with, as well as a record of the user’s previous performance across all ARPHA-hosted journals (i.e. number of accepted and declined editorial and review assignments, as well as the titles of the corresponding journals).

The feature is meant to facilitate and expedite the editorial process by discouraging assignment of tasks to overburdened or inactive users.

Find how to Manage Subject editor assignments and Invite Reviewers in the ARPHA Manual.

Review Quality Rating

Subject Editors should evaluate each review submitted to a manuscript they are handling by using a 5-star rating system. The average score is visible for Subject editors who consider the user as a Reviewer. The feature is meant to expedite the editorial process by aiding Subject Editors in the selection of the most suitable reviewers.

Find how to Rate a peer review in the ARPHA Manual.

Guidelines for Reviewers

Pensoft journals support the open science approach in the peer review and publication process. We encourage our reviewers to open their identity to the authors and consider supporting the peer review oaths, which tend to be short declarations that reviewers make at the start of their written comments, typically dictating the terms by which they will conduct their reviews (see Aleksic et al. 2015, doi:  10.12688/f1000research.5686.2  for more details): Principles of the open peer-review oath

• Principle 1: I will sign my name to my review
• Principle 2: I will review with integrity
• Principle 3: I will treat the review as a discourse with you; in particular, I will provide constructive criticism
• Principle 4: I will be an ambassador for the practice of open science
• Login is possible after registration at the journal's website. Our Editorial Office will register all first-time editors and reviewers. New users will receive an automated notification with a request to confirm registration and account information, and options for setting a password, email alerts and other features.   Note:  All users can use their registration details to login in all three (Book, E-Book and the respective Journal) platforms of  www.pensoft.net . Note:  Please remember that you may have registered with two or more different email addresses, that is why you may have more than one valid account at  www.pensoft.net . We advise using only one email address, hence one password associated with it, for all your operations at  www.pensoft.net . We highly recommend  that, in case the user has two or more different accounts, to merge these through user's profile.   Note: Users can at any time change the initially set password and correct personal details using their user's profile menu (by clicking on the user's name in the upper right corner of the screen appearing after login).

After login, please go to the respective journal’s web page and click on My Tasks button in the upper right corner of the screen. This way, you will be able to see all manuscripts you are responsible for as Author or Reviewer or Subject Editor.

Note: The manuscripts are grouped by categories, e.g., In Review (no.), In layout (no.), Published (no.), and Archived (no.) etc. The number in brackets after each category shows the number of manuscripts assigned to you.

General Responsibilities of Reviewers

This journal uses a single-blind peer review process. The reviewers are encouraged to disclose their identity, if they wish so. The peer review and editorial process is facilitated through an online editorial system and a set of email notifications. The online editorial system sends the Reviewer a review request, initiated by the Subject Editor or the Editorial Office. The online system will also inform about delays in the reviewing and will confirm a successful review submission. The email notifications contain stepwise instructions about the actions needed at each stage along with the link to the respective manuscript (accessible only after login – see section  How to Access a Manuscript ).

Reviewers are not expected to provide a thorough linguistic editing or copyediting of a manuscript, but rather focus on its scientific quality and overall style, which should correspond to the good practices in clear and concise academic writing. If Reviewers recognize that a manuscript requires linguistic edits, we shall be grateful for them to inform both the Author and the Subject Editor in the report. It is the Author’s responsibility to submit the manuscript in linguistically and grammatically correct English.

It often happens that even carefully written manuscripts may contain small errors in orthography or stylistics. We shall be thankful if Reviewers spot such errors during the reading process and correct them.

The manuscripts will generally be reviewed by two or three experts with the aim of reaching a first decision as soon as possible. Reviewers do not need to sign their reports, but are welcome to do so. They are also asked to declare any conflicts of interest.

Reviewers are asked whether the manuscript is scientifically sound and coherent, how interesting it is and whether the quality of the writing is acceptable. Where possible, the final decision is made on the basis of the peer reviews. In cases of strong disagreement between the reports or between the authors and peer reviewers, the editor can assess these according to his/her expertise or seek advice from a member of the journal's Editorial Board.

The ultimate responsibility for editorial decisions lies with the respective Subject Editor and/or, in some journals, with the Editor-in-Chief. All appeals should be directed to the Editor-in-Chief, who may decide to seek advice from the Subject Editors or the Editorial Board.

During a second review round, reviewers may be asked to evaluate the revised version against their recommendations submitted during the first review round.

Reviewers are kindly asked to be polite and constructive in their reports. Reports that may be insulting or uninformative will be rescinded.

Reviewers are asked to start their report with a very brief summary of the reviewed paper. This will help the editor and the authors see whether the reviewer correctly understood the paper or whether a report might be based on misunderstanding.

Furthermore, reviewers are also asked to comment on originality, structure and previous research:

Originality:  Is the paper sufficiently novel and does it contribute to a better understanding of the topic under scrutiny? Is the work rather confirmatory and repetitive?

Structure:  Is the introduction clear and concise? Does it place the work into the context that is necessary for a reader to comprehend aims, hypotheses tested, experimental design or methods? Are Material and Methods clearly described and sufficiently explained? Are reasons given when choosing one method over another one from a set of comparable methods? Are the results clearly, but concisely described? Do they relate to the topic outlined in the introduction? Do they follow a logical sequence? Does the discussion place the paper in scientific context and go a step beyond the current scientific knowledge on the basis of the results? Are competing hypotheses or theories reasonably related to each other and properly discussed? Do the conclusions seem reasonable?

Previous research:  Is previous research adequately incorporated into the paper? Are references complete, necessary and accurate? Is there any sign that substantial parts of the paper are copies of other works?

Peer Review Process

This journal uses a single-blind peer review process. Notwithstanding with that, the Reviewers are encouraged to disclose their identities, if they wish to do so. 

The Reviewer receives a review request generated by the Subject Editor or the Editorial Office and is expected to either agree to provide a review, or decline, through pressing the  Will do the review  or  Unable to do the review  link in the online editorial system. In case the Reviewer agrees to review the manuscript, he/she should submit the review within a certain time frame, which may vary in the different journals. Note:  The link to the respective manuscript is available in the review request email and all consequent reminder emails. The manuscript is accessible by clicking on the link in the email notification, or after login. Please look at the section  How to Access a Manuscript  above in case you have any difficulties.

The review should be submitted through the Proceed  button. The review should consist of:

• a simple online questionnaire to be answered by ticking either Yes , No , or N/A;
• comments addressed to the Author and the Subject Editor in the online form;
• associated files (corrected/commented manuscript file, review submitted in a separate text file, etc.), if any.

Note:  Reviewers can insert corrections and comments in the manuscript review version (PDF) and/or in the manuscript text file (usually Microsoft Word, rarely Open Office file). When working in the PDF, please use either the Text Edits or the Sticky Notes tools (available through the menu Tools -> Comments & Markup of the Acrobat Reader). When editing in Microsoft Word please use the Track Changes / Comments tools. Note: Associated files should be submitted at the end of the review process by clicking on the Browse  button, then selecting the respective file on your computer, and then pressing the Upload  button. A Reviewer may upload as many files to support his/her review as needed.

The Reviewer may decide to stay anonymous or open his/her identity by ticking the Show my name to the author(s)  box at the bottom of the reviewer’s form. Please be aware that your identity might be revealed in the comments or in Track Changes corrections of the Microsoft Word or PDF file you correct. Therefore, please make sure that you delete your name and initials in the Options section of your Word or PDF processor if you want to remain anonymous.

In addition to the above, by checking a box at the bottom of the submission form, reviewers may opt for making their contribution public in the event that the article is accepted and published. The reviewer's name, affiliation and email address will be displayed next to those of the Academic editor (or Subject editor) on the article webpage.

The review process is completed by selecting a recommendation from five options: (1)  Reject;  (2)  Reject, but resubmission encouraged ; (3)  Major Revision ; (4)  Minor Revision ; (5)  Accept . The system will ask for one more confirmation of the selected recommendation before submission. The submitted review cannot be changed after submission. Note:  Reasons for rejection can be a low scientific quality, non-conformance to the journal’s style/policies, and/or grammatically poor English language. Note:  It is also possible for review and associated files (e.g., a corrected manuscript file) to be sent as attached files to the email of the Editorial Office. We strongly recommend  avoiding this option, and instead uploading reviews through the online editorial management system.

Once a Reviewer submits a review of a manuscript, he/she receives an acknowledgement email from the journal.

The submission of the review is also automatically reported to Clarivate  - Web of Science Reviewer Recognition Service (formerly Publons). Reviewers are asked to confirm whether they want their reviews to be recorded on Clarivate.

When all Reviewers have submitted their reviews, the Subject Editor makes a decision to either accept, reject or request further minor/major revision.

After the Subject Editor's decision, the manuscript is sent back to the author for comments and further revision. The Author needs to submit a revised version in due time.

Reviewers are notified via email when the revised version of a manuscript that they have reviewed is submitted by the Author. They receive a link to the revised version along with the editorial decision and all reviews of the manuscript. Reviewers are also provided with a feedback form should they have any comments on the revised version. 

When an article is published, all Reviewers who have provided a review for the respective manuscript receive an email acknowledgment. In the email, there is a link to view/download the published article.

The Reviewer may always access information on the manuscripts that are being / have been reviewed by him/her through the menu My Tasks  –> Reviewer  on the journal’s web page – In Review (no.), In Edit (no.), Published (no.), and Archived (no.). The number in brackets after each category shows the number of manuscripts that have been assigned to you.

Science Communication

Our journal and the PR team at Pensoft invites authors to contribute to the communication and promotion of their published research, thereby increasing the visibility, outreach and impact of their work. 

Authors are welcome to notify us whenever their institution is working on a promotional campaign about their work published in our journal. We are always happy to reshare and/or repost (where appropriate). 

You can contact our PR team at [email protected] to discuss the communication and promotion of your research.

Tailored PR Campaign

(Paid service*)

We encourage authors, who feel that their work is of particular interest to the wider audience, to email us with a press release draft** (see template and guidelines ), outlining the key findings from the study and their public impact. Then, the PR team will work with them to finalise the announcement that will be:

• Issued on the global science news service Eurekalert!
• Sent out to our media contacts from the world’s top-tier news outlets
• Posted on ARPHA’s or Pensoft’s blog
• Shared on social media via suitable ARPHA-managed accounts

Following the distribution of the press announcement, our team will be tracking the publicity across news media, blogs and social networks, in order to report back to the author(s), and reshare any prominent media content.

Request our Tailored PR campaign service by selecting it while completing your submission form and you will be contacted once your manuscript is accepted for publication. Alternatively, contact our PR team  ( [email protected] ), preferably upon the acceptance of your manuscript.

* The Tailored PR campaign is an extra service charged at EUR 150 . However, we would consider discounts and even full waivers for studies of particular interest for the society.

**  Please note that our PR team reserves the right to edit your press release at their discretion. No press announcements will be issued until we receive the author’s final approval to do so. The service is only available for studies published within the past 3 months.

Guest Blog Post

(Free service)

Authors are strongly encouraged to promote their work and its impact on society to the audience beyond their immediate public of fellow scientists by means of storytelling in plain language. Ideally, such guest blog posts will be:

• Written from the author’s own point of view, using conversational tone;
• Written in fluent English;
• Presenting some curious background information, in order to place the discovery in context;
• Including attractive non-copyright imagery.

Request our Guest blog post service by contacting the PR department ( [email protected] ), regardless of the status of your submission, as there are no time constraints for guest blog post publication. Particularly encouraged are follow-up contributions telling the story of, for example, a research paper that has led to an important policy to be set in place; or an article that has met remarkable attention or reactions in the public sphere.

Following the necessary final touches to the guest blog post by the PR team, the contribution will be:

• Shared on social media via multiple and relevant ARPHA-managed accounts

Please note that the PR team reserves the right to refuse publication of a guest blog post on the occasion that it is provided in poor English, uses considerable amount of jargon or does not abide by basic ethical standards. Our PR team reserves the right to request changes to the text related to formatting or language. No blog posts will be issued until we receive the author’s final approval to do so.

Find past guest blog posts on Pensoft’s blog here . 

Video Podcast

To efficiently increase the outreach of their research, authors are suggested to prepare a video contribution (i.e. elevator video pitch, video abstract or topical video), where they present their work to an audience beyond their immediate public of fellow scientists by means of visual storytelling.

To do so, they are expected to send us a short (up to 02’00’’) video clip, presenting their study in a nutshell, in order to spark the viewer’s further interest in their findings and work, as well as the research topic as a whole. Ideally, such contribution will be:

• filmed in high quality, preferably with .mp4 file extension with the H.264 video codec;
• directed from the author’s own point of view, using conversational tone and minimal jargon;
• presented in fluent English or featuring English subtitles;
• accompanied by a transcript in English;
• accompanied by a short text introduction for the purposes of a blog post.

Request our Guest video contribution service by contacting the PR department ( [email protected] ), regardless of the status of your submission, since there are no time constraints for guest blog post publication.

Following the necessary final touches to the guest blog post, the contribution will be:

• Shared on Pensoft’s YouTube channel ;
• Posted on ARPHA’s or Pensoft’s blog;
• Shared on social media via multiple and relevant ARPHA-managed accounts. 

Please note that the PR team reserves the right to refuse distribution of a guest contribution on the occasion that it is provided in poor English, uses considerable amount of jargon or does not abide by basic ethical standards.

Custom Social Media Content

To help increase the visibility and outreach of their research, authors are welcome to suggest custom social media content to be distributed via suitable Pensoft- and ARPHA-managed social media accounts.

Social media posts are expected to:

• Be limited to two short sentences or 280 characters (including links);
• Be written in a conversational tone;
• Contain minimal jargon;
• Include the DOI link of the article;
• Not duplicate the title or abstract of the article;
• Include attractive non-copyright imagery;
• Possibly include up to 10 social media accounts, e.g. co-authors (Twitter only), affiliations, funding bodies etc. relevant to the study.

Request our Custom social media content service by contacting our PR department ( [email protected] ).

Please note that our PR team reserves the right to edit your text at their discretion.

Media Center

Follow Journal of Hymenoptera Research on Twitter and Facebook .

Learn about some of the most notable research published in Journal of Hymenoptera Research on Pensoft's blog.

See top news stories from around the world, mentioning research published in Journal of Hymenoptera Research in Forbes , Business Insider , Fox News , The Sun , The Independent , National Geographic ,  New York Post , Die Welt , The Conversation , The New York Times , Der Spiegel ,  Mongabay  and  РИА Новости (RIA Novosti).

Boost the reach of your paper(s) to a larger audience by making the most of Pensoft's science communication services .

Download journal presentation slides as a PowerPoint or PDF file.

Download journal logo .

Web Services

OAI-PMH - oai_dc: https://jhr.pensoft.net/oai.php?verb=ListRecords&set=jhr&metadataPrefix=oai_dc

OAI-PMH - mods: https://jhr.pensoft.net/oai.php?verb=ListRecords&set=jhr&metadataPrefix=mods

RSS for metadata: https://jhr.pensoft.net/rss.php

Journal Info

This website uses cookies in order to improve your web experience. Read our Cookies Policy

Biodiversity Heritage Library

Journal of Hymenoptera research

Title Variants

Abbreviated: J. Hymenopt. res

Alternative: JHR

International Society of Hymenopterists

Published material

Publication info

Washington, D.C, International Society of Hymenopterists, [1992-

Title from cover.

Hymenoptera , Periodicals

Call Number

QL563 .J68X

Identifiers

CODEN: JHYREJ ISSN (print): 1070-9428 LCCN: https://lccn.loc.gov/98660130 OCLC: 28308627 Wikidata: https://www.wikidata.org/entity/Q19830024

Find in a local library Download MODS

Open Access Research Journal of Life Sciences

Issn 2783-025x (online), biology, ecology, and taxonomy of the parasitoids of the families of austroniidae, peradeniidae, proctorenyxidae, roproniidae, and vanhorniidae (hymenoptera: proctotrupoidea), carlos henrique marchiori * , marco vinícios de oliveira santana and klebert de paula malheiros.

Copyright © 2024 Author(s) retain the copyright of this article. This article is published under the terms of the Creative Commons Attribution Liscense 4.0