Advertisement
Browse Subject Areas
?

Click through the PLOS taxonomy to find articles in your field.

For more information about PLOS Subject Areas, click here.

  • Loading metrics

Changes to the Fossil Record of Insects through Fifteen Years of Discovery

  • David B. Nicholson ,

    david.nicholson@nhm.ac.uk

    Current address: Department of Earth Sciences, The Natural History Museum, London, United Kingdom

    Affiliations Department of Biology, University of York, York, United Kingdom, Department of Natural Sciences, National Museum of Scotland, Edinburgh, United Kingdom

  • Peter J. Mayhew,

    Affiliation Department of Biology, University of York, York, United Kingdom

  • Andrew J. Ross

    Affiliation Department of Natural Sciences, National Museum of Scotland, Edinburgh, United Kingdom

Changes to the Fossil Record of Insects through Fifteen Years of Discovery

  • David B. Nicholson, 
  • Peter J. Mayhew, 
  • Andrew J. Ross
PLOS
x

Abstract

The first and last occurrences of hexapod families in the fossil record are compiled from publications up to end-2009. The major features of these data are compared with those of previous datasets (1993 and 1994). About a third of families (>400) are new to the fossil record since 1994, over half of the earlier, existing families have experienced changes in their known stratigraphic range and only about ten percent have unchanged ranges. Despite these significant additions to knowledge, the broad pattern of described richness through time remains similar, with described richness increasing steadily through geological history and a shift in dominant taxa, from Palaeoptera and Polyneoptera to Paraneoptera and Holometabola, after the Palaeozoic. However, after detrending, described richness is not well correlated with the earlier datasets, indicating significant changes in shorter-term patterns. There is reduced Palaeozoic richness, peaking at a different time, and a less pronounced Permian decline. A pronounced Triassic peak and decline is shown, and the plateau from the mid Early Cretaceous to the end of the period remains, albeit at substantially higher richness compared to earlier datasets. Origination and extinction rates are broadly similar to before, with a broad decline in both through time but episodic peaks, including end-Permian turnover. Origination more consistently exceeds extinction compared to previous datasets and exceptions are mainly in the Palaeozoic. These changes suggest that some inferences about causal mechanisms in insect macroevolution are likely to differ as well.

Introduction

A key contribution of palaeontology to the study of the diversity of life has been the elucidation of macroevolutionary patterns and processes through deep time, with fossils providing the only direct temporal evidence of how life has responded to a variety of biotic and abiotic forces [14]. If there are general rules underlying macroevolutionary responses to these forces, studying the past may also inform the future. Palaeontology can therefore, potentially, provide important information on the future progression of the extinction crisis facing the biosphere today, and its likely consequences [1,5].

In addition to such strategic questions, palaeontological data can help solve many basic questions of perennial interest. Comprising over 50% of described species [6], hexapods (insects and their close relatives such as springtails) form a major component of almost all continental ecosystems. An explanation of how and why this group has come to so dominate terrestrial biodiversity is a major challenge in macroevolutionary biology.

Palaeodiversity data are typically compiled in the form of taxonomic databases of fossils that provide either temporal ranges or discrete occurrence data. Commonly, criticisms of such databases focus around the integrity of the data and its resilience to the addition of further information [7]. Substantial additional knowledge, both taxonomic and stratigraphic, of the fossil records of tetrapods [8] and all marine animal families [9], has nonetheless yielded very similar variation in originations and extinctions though time. This supports the notion that broad biological signals can be seen through the statistical noise of an imperfect fossil record, providing the error is randomly distributed [10]. However, the effect of additional data on macroevolutionary patterns has not been tested for the majority of continental groups. This is important because many terrestrial taxa, such as insects, preserved primarily in exceptional conditions (Lagerstätten taxa) are likely to have substantially incomplete fossil records where the potential for change is much greater.

Using data on the temporal ranges of families, Labandeira [11] and Labandeira and Sepkoski [12] considered that, apart from the Late–end-Permian extinction, no other mass extinction event known from other groups appears to have had any major impact on insects at the family level. In addition, a steady increase of insect family-level richness began in the Triassic and was attributable not to particularly high levels of origination, but to consistently low extinction–noticeably lower than that in the Palaeozoic. The rise of angiosperms during the Cretaceous apparently did not cause any increase in family diversity in insects and may even have caused some decline in richness into the Late Cretaceous. However, Labandeira and Sepkoski [12] noted that much of the variation around this long term trend of increasing richness could be linked to specific rich fossil deposits (Lagerstätten) or stages where insect-bearing fossil deposits are poorly known and so were cautious with any such interpretations. Jarzembowski and Ross [13], using data based on but slightly updated from Ross and Jarzembowksi [14], highlighted four major insect origination events; during the Permo-Carboniferous, Early Jurassic, Early Cretaceous and the Eocene. They concurred with Labandeira and Sepkoski [12] that today’s exceptionally high insect diversity is the result of low extinction levels and sustained origination but disagreed that insects were essentially immune to mass extinction after the end-Permian event. Highlighting in particular an apparent decline in family richness seen in the Upper Cretaceous record, they suggest a causal link to the radiation of angiosperms. Additionally, Ross et al. [15] noted the increase in counts of origination and extinction in the Cretaceous as evidence of ecological turnover associated with angiosperms.

The field of palaeoentomology has expanded rapidly in the last two decades, with large increases in the number of active researchers and consequent publication output [16], as well important changes in taxonomy (e.g. the resurrection of the order Cnemidolestodea [17]), the dating of fossil deposits (e.g. the recognition of the mid-Cretaceous age of Burmese amber; see [18,19]) and the exploration of newly discovered insect-bearing formations globally (e.g. the Eocene amber deposits of India [20]).

To take account of these developments, in the first instance, a new dataset of the temporal ranges of hexapod families, compiled from literature (about 2,500 papers) published up to the end of 2009, is compared with that of Ross and Jarzembowski [14] (data from literature published up to the end of 1991) and Labandeira [11] by documenting changes and additions to the data. Then richness time series derived from these datasets are compared to assess any change in the signal provided by the fossil record in light of additional data. Although many recent studies of fossil richness through time have been derived from sample-based occurrence data (e.g. [1]), which facilitates the elimination of biases, studies of the face-value record are still valuable from a comparative perspective: they will, for example, help in the understanding of sampling-based artefacts by comparison with the geological record (e.g. [21,22]). In addition, taxonomic range data still retain considerable utility for the dating of phylogenies. There have to-date been no sample-standardized studies of fossil insect richness, and, although constraints on the completeness and comparability of data may limit what can currently be achieved, we expect that future studies will attempt them. A breakdown of the new data show which main groups of hexapods make a dominant contribution to the signal through time. From the first and last occurrence data, rates of origination and extinction can be calculated per stage indicating the timing of major radiation and extinction events as well as long-term trends and the relative importance of these to hexapod family richness.

Methods

Changes and additions to the hexapod fossil record

We quantified the amount of change in the new dataset (NEW) relative to the fossil insect family datasets presented by Ross and Jarzembowski [14] (downloaded from www.fossilrecord.net 2012-10-05) and Labandeira [11] (referred to herein as FR2 and LAB, respectively). The NEW dataset is dated according to the International Commission on Stratigraphy stages [23]. First, each family in NEW was categorised in the following ways with respect to FR2 and LAB: ‘no change’, ‘new in list’ and ‘range change’. The first of these is self-explanatory with respect to LAB, which, like NEW, presents data at stage resolution. However, FR2 presents data at both epoch and stage level, and no change for a family where data in FR2 were given at epoch or period level represents a case where the data in NEW confirm it was indeed present throughout that epoch or period. ‘New in list’ can refer to newly described families, those brought out of synonymy or Recent families which now have a fossil record. ‘Range change’, used only for comparison with FR2, involves a change in the recorded stratigraphic range of a family, whether an extension or contraction from the finding of new specimens but also includes improved resolution or revised dating of deposits from which previously known specimens occur (i.e. the deposit is now dated to a different stage). Since most of the LAB data is resolved to stage level and so is more directly comparable with the new data, range change was subdivided into three categories: contraction, extension and shift. A contraction is any situation where the NEW range has fewer stages than recorded in LAB, while an extension is any family where the new range covers a greater number of stages. This does not distinguish between whether the first and/or last occurrence has changed to create the contraction or extension and can also include instances where the NEW range has no overlap with that in LAB, e.g. the palaeodictyopteran family Hanidae, P1(Artinskian) in LAB but C2(Gzhelian)–P1(Sakmarian) in the new dataset. Shifts represent when the NEW range for a family covers a different set but the same total number of stages.

Derivation of richness time series from origination and extinction data

Before describing how various time series can be derived from first and last occurrence data, it is worth defining the four classes of taxa which can be counted in a time interval [24] (Fig 1).

thumbnail
Fig 1. Four classes of taxa recorded in an interval using first and last occurrence data.

After Foote [24]. The horizontal axis represents time progressing from left to right. The vertical lines represent the start (left) and end (right) of a specified time interval of interest. Horizontal lines represent the temporal ranges of four types of taxa of interest: FL originates and goes extinct within the interval, bL originates before and becomes extinct within the interval, Ft originates within and continues beyond the interval and bt originates before and continues after the interval.

https://doi.org/10.1371/journal.pone.0128554.g001

Some taxa (bt: bottom, top) originate before the time interval in question and have their last occurrence sometime after it, thus crossing the bottom and top boundaries. Some taxa (bL: bottom, Last) originate before the interval and have their last occurrence within it. Others (Ft: First, top) first appear in the interval and range beyond it. Finally, still others (FL: First, Last–also known as single-interval taxa) appear to originate and go extinct entirely within the interval, never crossing either the bottom or top boundaries. The term ‘single-interval taxon’ is preferable to the commonly used term ‘singleton’ when describing such taxa (as unfortunately done in e.g. [2426]) as the word is already in common usage in ecology for taxa represented by one specimen [27,28].

Two commonly-used counting methods exist for deriving diversity time series from first and last occurrence data–range through (RT) and boundary crossers (BC), and a third employed here, minimum assumption (MIN) [22,28]. These are applied to NEW and LAB data, while with the FR2 data only RT is used but under two assumptions–FR2+ and FR2, explained below.

RT is the classic method of counting a taxon as present in every stage between and including its first and last occurrences in the fossil record (or up to the present day if still extant), as well as those which originate and go extinct within the same time interval (known as single-interval taxa or FL in the notation given above), used, for example, by Sepkoski [9], Labandeira and Sepkoski [12] and Jarzembowski and Ross [13]. This is the sum total of taxa observed and inferred to exist within a time interval and can be written as RT = bt+Ft+bL+FL. For FR2, inconsistent stratigraphic resolution makes it necessary to use maximum and minimum assumptions of the ranges given when comparing with datasets at stage level. FR2+, then, is based on the assumption that the family originates in the first stage of the interval in which lies its first appearance and goes extinct in the last stage of the interval containing its last appearance, while FR2 assumes the origination in the last stage of the interval of first appearance and extinction in the first stage of the interval of last appearance [5]. Consequently, any family which is recorded at epoch or period level but in only one interval is removed from the FR2 series.

The BC series are made up of only those taxa which range between two or more time intervals, i.e. excluding single-interval taxa (FL). However, they are not simply RT minus FL. Rather, BC series represent the number of taxa crossing the bottom boundary into the interval, thereby tying diversity to a single point in time (the boundary) and not adding that diversity to events which occur cumulatively within the interval. It can be written as BC = bt+bL. By restricting the richness count to taxa which cross a single point in time, the data record an actual faunal cohort rather than the accumulation of taxa which exist throughout an interval. The specific advantage of this is that it is immune to changes in interval length, while it might be expected that longer intervals will accumulate more taxa than shorter ones, thereby inflating the richness measurement for that observation point. BC series have found use in some more recent palaeodiversity studies [2931] and have been advocated within the palaeoentomological community more recently by Ponomarenko and Dmitriev [32]. As these are values for interval boundaries, in order to make possible the comparison with data within intervals (placed at stage-midpoint) the geometric mean of the bottom and top boundaries of each interval are used for analyses, i.e. where BC1 and BC2 are the number of bottom and top boundary crossers of a given interval, respectively. Possible drawbacks of excluding single-interval taxa are that it excludes some true biological variation; may increase taxonomic bias by virtue of eliminating particular types of organism from the data; and the data then cease to represent all described variation, which is one of their chief merits.

The MIN series is derived from only the first, last and single-interval taxa, without filling in ranges. Like RT, this is a summation of events within a stage and can be written as MIN = Ft+FL+bL. This is the most conservative of the three as it makes the minimum assumption of what has actually been recorded in each stage and is more directly related to sampling proxies such as formation or collection counts [22]. It can be viewed as a subset of sampled-in-bin counts (counting only taxa which have actually been recorded in a time bin, rather than merely inferred to have existed at that time). Of course, it is a highly truncated version of true sampled-in-bin counts as the original purpose of the dataset was to record only first and last occurrences.

To quantify the similarity between the new and previous datasets, untransformed RT data from FR2, LAB and NEW are assessed using Spearman’s rank correlation because, even when logged, the data were skewed, breaking parametric assumptions. The normal associated probabilities are not reported because autocorrelations in the data invalidate them. Bootstrap estimates for significance of correlations, which reduce the necessary set of assumptions about the data, are instead calculated using the boot.ci function from the boot library in R to re-sample the original data 9999 times, each time recalculating the correlation coefficient, to generate a bootstrapped distribution of the test statistic which indicates the extent of uncertainty in it. Confidence intervals at the 95% and 99% level are calculated using the bca (bias corrected and accelerated or BCa) method due to Efron [33], which corrects for the bias (the difference between the mean of the bootstrap replicates and the true correlation) and asymmetry of the bootstrap distribution [33]. Where the confidence intervals do not bracket zero, the correlation can be said to be significantly different from zero. Note that in the case of “dependent” data such as time-series, bootstrapping may still not accurately estimate confidence intervals around statistical parameters, so, whilst likely more valid than the standard p-values, should still be treated with caution. Correlations were also explored for two detrended versions of each time series: first differencing explores the changes between successive time steps (stages), whilst generalized differencing (first differencing of the residuals from linear regression) quantifies the successive changes after removing the overall long term trend. Differences were calculated using the statistical programming language R [34]. All correlations are on data from the Serpukhovian (top of Early Carboniferous, stage midpoint 323.2Ma) to Piacenzian (top of the Pliocene, stage midpoint ~3.1Ma), as this is the range for which there is a reasonable fossil record of hexapods (i.e. including the long period of almost no record before the Carboniferous would increase all of the coefficients simply from a lack of data).

Calculating origination and extinction rates

The rates of origination and extinction employed here are Foote’s [24] estimated per-capita rates, and respectively. They are derived as follows: where Nt is the total number of taxa crossing the top boundary out of the interval (i.e. bt+Ft), Nb is the total number crossing the bottom boundary into it (i.e. bt+bL) and Nbt is the number of taxa crossing both the bottom and top boundary. The advantage of using these over counts of events within an interval is that they are robust to variation in interval duration, disregard single-interval taxa (which are prone to disproportionately distort the signal) and are independent of each other as they are derived from numbers of taxa passing into and out of intervals rather than the addition of events taking place within them. Due to inconsistent stratigraphic resolution, this is not attempted for the FR2 data.

Results

Changes in the data

The NEW dataset (S1 Appendix and S1 Dataset) contains a total of 1454 families of Hexapoda, of which 1436 are Insecta. In comparison to FR2, a substantial amount of change has left only 8% of families with the same ranges as recorded in 1993; 35% are new to the record, and well over half have a change in the recorded range (Fig 2A). The picture is broadly similar when compared to LAB (Fig 2B), with 10% remaining unchanged and 30% new. The majority of the range changes are made up of roughly equal amounts of extensions and contractions, and only 7% of the total representing a shift in range. Although the NEW dataset has a higher total number of families (1454) than either FR2 (1087 in [14]) or LAB (1272; 1276 if including ‘uncertain’ families), 230 and 263 families listed in FR2 and LAB, respectively, are included in NEW within other families (as synonyms or subfamilies), or no longer have a fossil record, due mostly to taxonomic revisions.

thumbnail
Fig 2. Proportions of changes in new data for family stratigraphic range compared with previous datasets.

(A) FR2 [14] all hexapods and (B) LAB [11] all insects.

https://doi.org/10.1371/journal.pone.0128554.g002

Richness series from new and previous datasets

The richness time series of all three datasets show broad similarities in long-term trends of increasing richness and the synchronicity (or nearly so) of several pulses (Fig 3) but some differences are worth noting.

thumbnail
Fig 3. Family richness of insects through time.

Richness time series derived from (A) NEW data, presented here, (B) LAB data from Labandeira [11] and (C) FR2 data from Ross and Jarzembowski [14]. RT = range through, i.e. all taxa ranging anywhere into an interval, with maximum (+) and minimum (–) assumptions for FR2, plotted at stage-midpoints. BC = boundary crossers, i.e. taxa crossing interval boundaries, plotted at stage boundaries. MIN = minimum richness, representing firm occurrences within stages (i.e. first, last and single-interval taxa records).

https://doi.org/10.1371/journal.pone.0128554.g003

For the Palaeozoic, the RT series from NEW and LAB are more similar to each other than to FR2+. However, the NEW series shows consistently lower richness than LAB and the two main peaks are offset by one stage, reaching a maximum of 109 families by NEW RT (Kungurian: 273 Ma) and 153 by LAB RT (Artinskian: 280 Ma) (Fig 3). FR2+ shows a gradual and steady increase in richness through the Palaeozoic with a dramatic drop at the end-Permian (~250 Ma), after reaching a maximum of 168 families (Fig 3). FR2 shows no such increase and decline but rather remains conspicuously flat through until the Late Triassic at around 210 Ma (Norian). This is also not mirrored by LAB RT and NEW RT, which show slightly less sharp declines from the Early–Middle Permian towards the end-Permian, when a small increase is seen in the final stage (Changhsingian, data point at 252 Ma). The BC series in NEW and LAB mirror the peaks and troughs of the RT curves but they are less pronounced (Fig 3).

In the Triassic (251–200 Ma) all three datasets show a marked increase in richness, with the largest increase in the Carnian (223 Ma) for FR2+ (up to 123 families) and NEW (171 families) and in the Ladinian (233 Ma) for LAB (117 families) (Fig 3). The NEW RT series show the most pronounced Triassic peak followed by an apparent crash in richness, mirrored in the NEW MIN series but NEW BC shows a smooth increase with only a slight decrease after the Carnian, reflecting that many of the records are single-interval taxa.

The Jurassic (200–146 Ma) continues the long-term increase in described richness (Fig 4). The NEW RT series shows a distinct, four-pulsed increase at 193 (Sinemurian), 179 (Toarcian), 158 (Oxfordian), and 148 Ma (Tithonian); the first three are followed by drops in richness, although this is not reflected in the BC series which shows an uninterrupted, fairly smooth increase. An almost identical pattern is seen in LAB RT while FR2+ shows two distinct increases followed by plateaus.

thumbnail
Fig 4. Richness and rates of origination and extinction in fossil hexapods.

(A) Range through time series for NEW, LAB and FR2. (B) Origination (Orig) and extinction (Ext) counts, both including (+) and excluding (–) single interval taxa, from NEW.

https://doi.org/10.1371/journal.pone.0128554.g004

During the Early Cretaceous (146–100 Ma) a more rapid rise is seen, most steeply in NEW RT. LAB and FR2 are similar in then showing a pronounced and sustained drop in richness after their synchronous peaks in the Aptian (point at 119 Ma) in both RT and BC series while the NEW RT series continues to increase, albeit at a decelerated rate until it plateaus across a similar range of stages as LAB and FR2. This plateau is accompanied by very low values in the NEW MIN series. No marked drop in richness is apparent at or near the Cretaceous/Palaeogene boundary (65.5 Ma).

The NEW RT series averages 15% and 26% higher across the Cretaceous and Tertiary compared with LAB and FR2, respectively ending with maxima of 695 (NEW), 549 (FR2) and 625 (LAB) families. All three show the most rapid increase in richness in the entire fossil record through the Tertiary with very little deviation between RT (or +) and BC (or) series.

The major taxa dominating richness in NEW (RT) varied at different times (Fig 5). The earliest known hexapod families are in the ‘Apterygota’. These contribute very little to hexapod fossil richness in the long term. The Carboniferous and Permian peaks and subsequent declines are seen only in the Palaeoptera and Polyneoptera. Paraneoptera and Holometabola had originated before the Permian peak but show no sign of any decline towards the end-Permian, rather a slow but steady increase in richness (Fig 5). The Late Triassic peak seen in the RT (but not BC) series is apparent in all groups except Apterygota. Except for occasional pulses of increased richness, which are synchronous with the other three major contributing groups, Palaeoptera show very slow and steady growth in richness, only attaining their previous Palaeozoic richness in the Palaeogene/Neogene from ~60 Ma onwards. A broadly similar pattern is seen in Polyneoptera. Paraneoptera, however, continue their steady increase from the Palaeozoic and show a pronounced increase during the Early Cretaceous (Berriasian–Albian), between ~150 and 100 Ma (Fig 5). This then levels out until they enter a phase of rapid expansion starting in the Palaeogene, from ~65 Ma onwards. The Holometabola enter a more rapid phase of expansion earlier than the Paraneoptera, starting in the Early Jurassic (Sinemurian, from ~193 Ma onwards). They show a pronounced jump in richness at 128 Ma (Barremian), being the largest contributing group to the rapid rise in richness during the Early Cretaceous seen in the NEW RT series. This is followed by a long plateau and then the most rapid expansion phase seen in the entire hexapod fossil record from the lower Eocene (52.2 Ma) onwards (Fig 5).

thumbnail
Fig 5. Spindle diagram showing NEW range-through family richness in major constituent hexapod groups through time.

Generated using PAST [69].

https://doi.org/10.1371/journal.pone.0128554.g005

Both the FR2+ and LAB RT series are highly correlated (i.e. strongly co-vary) with NEW RT (Table 1), with all values of Spearman’s rho greater than 0.95 and significant at the 99% confidence limit. This decreases substantially with both first and generalised differencing (Table 1), and correlations between NEW RT and LAB RT lose significance, whilst those between NEW RT and FR2+ retain their significance.

thumbnail
Table 1. Spearman rank correlations between richness time series using raw values and after first differencing and generalised differencing.

https://doi.org/10.1371/journal.pone.0128554.t001

Calculated origination and extinction rates

First and last occurrences occur episodically throughout the fossil record of insects (Fig 4B), with an apparent synchrony between origination and extinction through time with origination outstripping extinction. The modal origination occurs in the Palaeogene with large peaks in the Triassic, Late Jurassic and Early Cretaceous. Modal extinction occurs in the Early Cretaceous with large peaks in the late Carboniferous, Permian, Triassic, later Jurassic and Early Cretaceous. Per capita rates of origination and extinction ( and , respectively; Fig 6), however, show distinctly different profiles in the Palaeozoic and post-Palaeozoic (boundary at 251 Ma) in both NEW and LAB data. Greater variance is seen in the Palaeozoic for both rates in both datasets as well as the highest values reached in each. As for raw counts, per capita origination rates stay robustly higher than extinction from the Triassic onwards and both show long term declines towards the present. There are some notable differences between NEW and LAB: the timing and size of Permian origination peaks differs; there is no Late Cretaceous origination peak; the Carboniferous extinction peak is more pronounced, and those in the Permian less pronounced, not exceeding originations by much. As a result, Spearman rank correlations of these rates between NEW and LAB show no significant relationship in origination rates, while the extinction rates are positively correlated in the raw and generalised differenced time series but retain no relationship after first differencing (Table 1). In general, origination rates seem to more consistently exceed extinction rates.

thumbnail
Fig 6. Estimated per-capita rates of origination and extinction .

Rates are from (A) NEW insect family data and (B) Labandeira [11].

https://doi.org/10.1371/journal.pone.0128554.g006

Discussion

Changes in the data

The robustness of described richness through time in the insects, to new discoveries over fifteen years (eighteen years from FR2 data up to end 1991), was tested by compiling a new dataset of fossil hexapod family-richness from literature published up to the end of 2009. Only ten percent of families in the new data remain unchanged over that time, with about 60% of families having different stratigraphic ranges, and 30% of families being completely new to the fossil record. For scientists interested in the details of individual fossil families, for example for dating phylogenies above family level (e.g. [35,36]), the current dataset represents a substantial improvement over previous datasets available. The implication is that the previous fossil insect datasets now have largely historical interest only and should not be used for future macroevolutionary research. Studies based on them ideally require re-assessment. For example, in an analysis based on Labandeira’s [11] data, Yang [37] suggested that differences in origination rates account for the higher diversity of Holometabola compared to Paraneoptera. However, Nicholson et al. [38], using the present data set, found the converse: origination rates between these groups is not significantly different, but extinction rates are significantly reduced in Holometabola compared to Paraneoptera.

While the change in ranges from FR2 in the NEW data (Fig 2A) can be attributed largely to improvement in the stratigraphic resolution of family ranges to stages, the differences from LAB (Fig 2B) require more subtle explanation. Extensions of known ranges in fossil families are to be expected, with continued exploration of fossil sites and descriptions of new finds likely to turn up new first or last occurrences, such as the high rate of discovery in Mesozoic deposits of China (e.g. see [39]). The high proportion of range contractions (25%) seems at first unexpected but can be ascribed to differences in the dates for fossil deposits used (e.g. the Karabastau Formation, Kazakhstan: Kimmeridgian in LAB but Oxfordian in NEW) and extensive changes in taxonomy reducing the number of fossils included in some families, such as in a recent review of termites by Engel et al. [40] wherein several fossil taxa, previously attributed to extant families, were reassigned, thus contracting the known range of some families and removing the Hodotermitidae from the fossil record altogether.

The rate of discovery of new fossil hexapods seems disproportionately concentrated in the Cretaceous, with high numbers of publications on the extensive Jiulongshan and Yixian formations in China [39], continued interest in the Crato Formation in Brazil [41], a new supply of Burmese amber [18,42] and abundant new amber deposits in France [43] and Spain [44], although new material continues to be found across almost the entire temporal range of hexapods [6,16,45]. There are an estimated 1067 extant hexapod families (data compiled from the relevant sections of [46]), implying that ~370 extant families (35%) are not yet known from the fossil record and could in principle be found in future. This sets a broad potential upper limit to the height of the richness curve, indicating substantial, but not excessive, potential for future discovery at the family level. The majority of these (196 families) are from the Holometabola. However, in terms of proportion of extant families represented in the fossil record, Holometabola have the most coverage with ~69% represented, followed closely by Polyneoptera (65%), Paraneoptera (64%) and Palaeoptera (58%). Only 33% of extant Apterygota families have a fossil record, perhaps a result of their small size, habitat requirements, and lack of wings [6].

Other informative ways of assessing the potential for future discovery, beyond the scope of the present study, would be to construct taxon vs. specimen accumulation curves to observe if the number of taxa described through time has asymptoted (e.g. [4749]), or by quantifying the gaps in the record implied by phylogenies (e.g. [47,50,51]). Although some data pertinent to the former (dates of description of extinct families) are present in the current data, one would additionally need to compile the date at which extant families were first described from the fossil record, which is not normally the date of the taxon’s first description.

Changes in the richness series

Despite major changes to the ranges of insect families over fifteen years of discovery, changes to the pattern of described richness through time derived from those data seem less extensive. Correlations between the time series of the new and previous datasets show that the broad pattern of rise in discovered taxa through time is very similar to that previously described. The generally steady rise in richness through time suggests support for the previous conclusion [12] that no strong logistic limits to family richness have yet been met. However, some of the Cenozoic rise may be attributable to the Pull-of-the-Recent [52] whereby the ranges of extant taxa are pulled forward, accentuating the richness rise nearer the present. Sampling may also have been strongly affected by the abundance of suitable deposits, such as Baltic amber and compression deposits such as the Green River and Florissant formations, which coincide with the Eocene rise [53]. These issues will be examined in future papers.

Other important features preserved in the NEW richness series include evidence for a mass extinction at the end-Permian. The Permian drop in richness is however less abrupt than in FR2. This effect is probably due to the improved temporal resolution from epoch to stage, which pulls the ranges of taxa in FR2 forward to the end of the Permian. At stage level resolution, many of these families are instead seen to have last occurrences before the end Permian. In turn, the asynchronicity in extinction may be genuine, but probably is partly an artefact of an incomplete record (the Signor-Lipps effect [54]) which tends to drag extinctions backwards in time. The major turnover in dominant taxa (Fig 5) accompanying the Permian to Triassic interval is strongly reminiscent of the end-Permian extinction in many other taxa (e.g. [55]). In the hexapod case there was a replacement of the Palaeozoic fauna of mainly Palaeoptera and Polyneoptera by a fauna dominated by Paraneoptera and Holometabola, which appear to have suffered little reduction in their richness [13,53]. Studies on the coherence of these different faunas would be useful (see [56]).

Despite the evidence for an end-Permian extinction, the NEW richness data leave no evidence of an end-Cretaceous extinction, in common with previous data [15,53]. Given the known widespread ecosystem impacts of this event, it is difficult to imagine that insects were completely unaffected but extinction may have occurred below the family level. Some genus-level data provide some support for this [13], as do some studies of trophic interactions [57], but others suggest a weaker extinction in insects than in other taxa [58].

Although all datasets show an increase in richness in the Triassic, a subsequent drop is suggested by the NEW RT series (Fig 4A). Many non-insect taxa apparently experienced a mass extinction at the end-Triassic [59,60] but there has never been good evidence for this in insects. However, the drop is lost in the NEW BC series (Fig 3A), indicating that it is due primarily to abundance of single interval taxa and hence may be an artefact of sampling bias. Indeed the total number of extinctions detected at the end-Triassic boundary is close to zero, indicating that it would be premature to suggest an insect extinction then (Fig 4B).

Surprisingly, the overall level of richness in the NEW data is not always higher than the older data. This is mostly the case in the Palaeozoic, where there was an historical tendency by early workers such as Handlirsch and Tillyard to oversplit taxa, while revisions have decreased the number of valid families. Additionally, and perhaps more importantly, of the 324 families in the new data with ranges in the Palaeozoic, 28% of them represent contractions with respect to LAB. This suggests a specific effect of taxonomy on apparent richness that may be important for other researchers.

The correlations between the differenced time series for the new and old data, although positive, are much less strong than for the raw time series, suggesting moderate differences in the shorter term variation in richness from stage to stage. This is potentially important when assessing the drivers behind diversity change, as time series are generally detrended to remove spurious correlations, and it is the short term variation around the long term trends that are analysed (e.g. [5,61]). The Palaeozoic contains much of the discordance between the series (Fig 4A), with FR2 and NEW having very different shapes while the richness peaks of LAB and NEW are offset from each other. Declines seen in both LAB and FR2 during the Early–mid-Cretaceous (~120–85 Ma) are not shared by NEW, which shows more of a plateau. This plateau could simply be a result of the relative paucity in insect-preserving localities in the Upper Cretaceous, however in an analysis of plant-associated insect families Labandeira [62] found that this plateau coincides with a period of transition from gymnosperm-associated families to angiosperm-associated families, with extinctions in the former approximately matching the level of originations in the latter. Future analyses of occurrence-based data, subsampled to remove the effects sampling bias, will help to elucidate the relative importance of these alternative (but not mutually exclusive) explanations.

Patterns of origination and extinction

Labandeira [53] picks out five major periods of originations in the insects and four major extinctions. Of the originations, all are still found in the NEW origination series (Fig 4), namely in order, the Late Carboniferous (Bashkirian–Moscovian–first appearance of winged insects and colonization of forested ecosystems); Early Permian (peaking in the Kungurian–colonization of wider environments and the rise of Paraneoptera and Holometabola); Late Jurassic (Oxfordian–radiation of communities on advanced seed plants); Early Cretaceous (peaking in the Barremian–radiations in decomposer and freshwater systems); and the Eocene (Priabonian–suspected to be a sampling artefact that may represent earlier radiations that are poorly sampled). The main addition to this description in the NEW data is the higher peak in the Triassic, which Labandeira [53] attributes to a rebound from the Permian extinction.

In terms of extinctions, the Late Carboniferous peak is attributed by Labandeira [53] to changes in plant communities and trophic structure. The Permian extinction is high in absolute numbers of extinctions but lower in per capita rates (cf. Fig 4 and Fig 6) and is generally attributed to high continentality and hot dry climates on land [63]. In addition, there were substantial extinctions in the Late Jurassic (attributed to competitive turnover during the simultaneous radiation; [53]) and the Early Cretaceous (attributed to competitive turnover of taxa adapting to new environments, including angiosperms; see [13,15]). The NEW series add to this a large peak in extinctions in the Triassic, as seen for originations. As discussed above, this may represent the detection of the more general end-Triassic mass extinction, although it may also be an artefact of sampling bias.

In general the high agreement between the timing of originations and extinctions in NEW and FR2 is consistent with the findings of similar studies on other taxa [8,9], suggesting that the great potential for change in the insect fossil record has not translated into major changes in pattern. Some previous authors [9] have interpreted this as encouragement that incomplete and partially erroneous data can preserve broad generalizations about the history of life [10]. However, recent experiences with alternative ways of compiling the data suggest that other issues with the data can remain important in correctly describing and interpreting them [29,64,65].

In general there is high synchronicity between the origination and extinction series (Fig 4B), which is the pattern expected if one biologically depends on the other. This pattern is also expected if they are both simply artefacts of sampling, hence determined by the availability of insect-bearing deposits. The pattern is not simply due to the abundance of single interval taxa (Fig 4B), suggesting perhaps some biological signal in the data.

Originations mostly exceed extinctions across intervals, explaining the consistent rise in family level diversity through time, as well as high extant richness [4,15,66]. In terms of rates, the decline from the Palaeozoic to Mesozoic and Cenozoic is the most obvious feature, in common with other family and genus level analyses [60,64]. Explanations for this include lineage sorting, density-dependent processes and the fact that higher taxa are disproportionately described for older groups [64]. Some of the peaks are different in height in the NEW data compared to LAB (Fig 6); a result of taxonomic changes and shifts in the dating of deposits. The Late Cretaceous (Santonian: 85 Ma) LAB origination peak is not seen in NEW, probably from range extensions pulling more first occurrences back to Lower Cretaceous deposits.

An important question now is to what extent the updated richness, origination and extinction series reflect geological and sampling biases, and thus what features may remain once such biases are removed. Previous work on the marine invertebrate record suggests that many features of the face-value fossil record can remain preserved after sample-standardization (e.g. [67]), although diversity curves may become flatter due to the elimination of the Pull-of-the-Recent [65]. In addition, short term patterns after detrending may be altered, potentially altering macroevolutionary inference [64,68]. Because hexapods generally require exceptional preservation conditions, it is possible that sampling and other biases are a more prominent influence on the face-value record, although considerable biological signal may be retained. Occurrence data of community samples will likely help solve this. However, the range data here are likely to retain value in the dating of phylogenetic trees, since one can be more confident about the completeness of the family ranges, and they are easy to update.

In summary, a new compilation of the fossil ranges of insect families shows changes in the ranges of a high proportion of families, and significant changes in described short term richness and some origination and extinction patterns, but little change in broad temporal patterns. Representing an additional 15 years of data in a rapidly expanding field compared with previously available compendia [11,14], it is hoped that this new dataset will form the basis for future work on elucidating the evolutionary history of the hexapods.

Supporting Information

S1 Appendix. Fully referenced family-range dataset, including additional references [701097].

https://doi.org/10.1371/journal.pone.0128554.s001

(PDF)

S1 Dataset. Family-range spreadsheet dataset, with counts and rates of originations and extinctions.

https://doi.org/10.1371/journal.pone.0128554.s002

(XLSX)

Acknowledgments

DBN was supported by a NERC studentship and by the Natural History Museum and National Museums Scotland. We thank the many authors who sent us reprints of their work, and two anonymous reviewers for their helpful comments on the manuscript.

Author Contributions

Conceived and designed the experiments: AJR PJM DBN. Analyzed the data: DBN PJM AJR. Wrote the paper: DBN PJM AJR.

References

  1. 1. Alroy J. The shifting balance of diversity among major marine animal groups. Science. 2010; 329: 1191–1194. pmid:20813951
  2. 2. Benson RBJ, Mannion PD. Multi-variate models are essential for understanding vertebrate diversification in deep time. Biol Lett. 2012; 8: 127–130. pmid:21697163
  3. 3. Ezard THG, Aze T, Pearson PN, Purvis A. Interplay between changing climate and species’ ecology drives macroevolutionary dynamics. Science. 2011; 332: 349–351. pmid:21493859
  4. 4. Mayhew PJ. Why are there so many insect species? Perspectives from fossils and phylogenies. Biol Rev. 2007; 82: 425–454. pmid:17624962
  5. 5. Mayhew PJ, Jenkins GB, Benton TG. A long-term association between global temperature and biodiversity, origination and extinction in the fossil record. Proc R Soc B Biol Sci. 2008; 275: 47–53.
  6. 6. Grimaldi DA, Engel MS. Evolution of the Insects. Cambridge University Press; 2005.
  7. 7. Benton MJ. The history of life: large databases in palaeontology. In: Harper DAT, editor. Numerical palaeobiology: Computer-based modelling and analysis of fossils and their distributions, pp. 249–283. Chichester and New York: John Wiley & Sons; 1991. pp. 249–283.
  8. 8. Maxwell WD, Benton MJ. Historical tests of the absolute completeness of the fossil record of tetrapods. Paleobiology. 1990; 16: 322–335.
  9. 9. Sepkoski JJ Jr. Ten years in the library: new data confirm paleontological patterns. Paleobiology. 1993; 19: 43–51. pmid:11538041
  10. 10. Adrain JM, Westrop SR. An empirical assessment of taxic paleobiology. Science. 2000; 289: 110–112. pmid:10884223
  11. 11. Labandeira CC. A compendium of fossil insect families. Milwaukee Public Museum Contrib Biol Geol. 1994; 88: 1–71.
  12. 12. Labandeira CC, Sepkoski JJ Jr. Insect diversity in the fossil record. Science. 1993; 261: 310–315. pmid:11536548
  13. 13. Jarzembowski EA, Ross AJ. Insect origination and extinction in the Phanerozoic. Geol Soc London Spec Publ. 1996; 102: 65–78.
  14. 14. Ross AJ, Jarzembowski EA. Arthropoda (Hexapoda; Insecta). In: Benton MJ, editor. The Fossil Record 2. London: Chapman and Hall; 1993. pp. 363–426.
  15. 15. Ross AJ, Jarzembowski EA, Brooks SJ. The Cretaceous and Cenozoic record of insects (Hexapoda) with regard to global change. In: Culver SJ, Rawson PF, editors. Biotic Response to Global Change, the Last 145 Million Years. Cambridge University Press; 2000. pp. 288–302.
  16. 16. Ross AJ. The development of Palaeoentomology over the past 25 years. In: 5th International Conference on Fossil Insects, 4th World Congress on Amber Inclusions, 4th international Meeting on Continental Palaeoarthropodology, Beijing, Program & Abstract; 2010. pp. 90–91.
  17. 17. Béthoux O. Cnemidolestodea (Insecta): an ancient order reinstated. J Syst Palaeontol. 2005; 3: 403–408.
  18. 18. Ross AJ, Mellish C, York P, Crighton B. Burmese Amber. In: Penny D, editor. Biodiversity of fossils in amber from the major world deposits. Siri Scientific Press; 2010. pp. 208–235.
  19. 19. Shi G, Grimaldi DA, Harlow GE, Wang J, Wang J, Yang M, et al. Age constraint on Burmese amber based on U–Pb dating of zircons. Cretac Res. 2012; 37: 155–163.
  20. 20. Rust J, Singh H, Rana RS, McCann T, Singh L, Anderson K, et al. Biogeographic and evolutionary implications of a diverse paleobiota in amber from the early Eocene of India. Proc Natl Acad Sci USA. 2010; 107: 18360–18365. pmid:20974929
  21. 21. Peters SE. Geologic constraints on the macroevolutionary history of marine animals. Proc Natl Acad Sci USA. 2005; 102: 12326–12331. pmid:16105949
  22. 22. Peters SE, Foote M. Biodiversity in the Phanerozoic: a reinterpretation. Paleobiology. 2001; 27: 583–601.
  23. 23. Ogg JG, Ogg G, Gradstein FM. The Concise Geologic Time Scale. Cambridge University Press; 2008.
  24. 24. Foote M. Origination and extinction components of taxonomic diversity: general problems. Paleobiology. 2000; 26: 74–102.
  25. 25. Alroy J. New methods for quantifying macroevolutionary patterns and processes. Paleobiology. 2000; 26: 707–733.
  26. 26. Fitzgerald PC, Carlson SJ. Examining the latitudinal diversity gradient in Paleozoic terebratulide brachiopods: should singleton data be removed? Paleobiology. 2006; 32: 367–386.
  27. 27. Preston FW. The commonness, and rarity, of species. Ecology. 1948; 29: 254–283.
  28. 28. Alroy J. Fair sampling of taxonomic richness and unbiased estimation of origination and extinction rates. Paleontol Soc Pap. 2010; 16: 55–80.
  29. 29. Alroy J. Successive approximations of diversity curves: Ten more years in the library. Geology. 2000; 28: 1023–1026.
  30. 30. Alroy J, Marshall CR, Bambach RK, Bezusko K, Foote M, Fürsich FT, et al. Effects of sampling standardization on estimates of Phanerozoic marine diversification. Proc Natl Acad Sci USA. 2001; 98: 6261–6266. pmid:11353852
  31. 31. Bambach RK. Energetics in the global marine fauna: A connection between terrestrial diversification and change in the marine biosphere. Geobios. 1999; 32: 131–144.
  32. 32. Ponomarenko AG, Dmitriev VY. Diversity curves revisited. Paleontol. J. 2009; 43: 226–229.
  33. 33. Efron B. Better bootstrap confidence intervals. J Am Stat Assoc. 1987; 82: 171–185.
  34. 34. R Core Team. R: A Language and Environment for Statistical Computing. Vienna Austria R Foundation for Statistical Computing; 2014.
  35. 35. Davis RB, Nicholson DB, Saunders ELR, Mayhew PJ. Fossil gaps inferred from phylogenies alter the apparent nature of diversification in dragonflies and their relatives. BMC Evol Biol. 2011; 11.
  36. 36. Rainford JL, Hofreiter M, Nicholson DB, Mayhew PJ. Phylogenetic distribution of extant richness suggests metamorphosis is a key innovation driving diversification in insects. PLoS One. 2014; 9: e109085. pmid:25275450
  37. 37. Yang AS. Modularity, evolvability, and adaptive radiations: a comparison of the hemi- and holometabolous insects. Evol Dev. 2001; 3: 59–72.
  38. 38. Nicholson DB, Ross AJ, Mayhew PJ. Fossil evidence for key innovations in the evolution of insect diversity. Proc R Soc B Biol Sci. 2014; 281.
  39. 39. Ren D, Shih C-K, Gao T-P, Yao Y-Z, Zhao Y-Y. Silent Stories–Insect Fossil Treasures from Dinosaur Era of the Northeastern China. Beijing: Science Press; 2010.
  40. 40. Engel MS, Grimaldi DA, Krishna K. Termites (Isoptera): their phylogeny, classification, and rise to ecological dominance. Am Museum Novit. 2009; 3650: 1–27.
  41. 41. Martill DM, Bechly G, Heads SW. Appendix: species list for the Crato Formation. In: Martill DM, Bechly G, Loveridge RF, editors. The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge University Press; 2007. pp. 582–607.
  42. 42. Grimaldi DA, Engel MS, Nascimbene PC. Fossiliferous Cretaceous amber from Myanmar (Burma): Its rediscovery, biotic diversity, and paleontological significance. Am Museum Novit. 2002; 3361: 1–71.
  43. 43. Perrichot V, Néraudeau D. Foreword: Cretaceous ambers from southwestern France: geology, taphonomy, and palaeontology. Geodiversitas. 2009; 31: 7–11.
  44. 44. Delclòs X, Arillo A, Peñalver E, Barrón E, Soriano C, López Del Valle R, et al. Fossiliferous amber deposits from the Cretaceous (Albian) of Spain. Comptes Rendus Palevol. 2007; 6: 135–149.
  45. 45. Nel A, Roques P, Nel P, Prokin AA, Bourgoin T, Prokop J, et al. The earliest known holometabolous insects. Nature. 2013; 503: 257–261. pmid:24132233
  46. 46. Resh VH, Cardé RT. Encyclopedia of Insects. 2nd ed. Elsevier Academic Press; 2009.
  47. 47. Smith AB. Intrinsic versus extrinsic biases in the fossil record: contrasting the fossil record of echinoids in the Triassic and early Jurassic using sampling data, phylogenetic analysis, and molecular clocks. Paleobiology. 2007; 33: 310–323.
  48. 48. Puchalski SS, Eernisse DJ, Johnson CC. The effect of sampling bias on the fossil record of chitons (Mollusca, Polyplacophora). Am Malacol Bull. 2008; 95: 87–95.
  49. 49. Bernard EL, Ruta M, Tarver JE, Benton MJ. The fossil record of early tetrapods: Worker effort and the end-Permian mass extinction. Acta Palaeontol Pol. 2010; 55: 229–239.
  50. 50. Wills MA. How good is the fossil record of arthropods? An assessment using the stratigraphic congruence of cladograms. Geol J. 2001; 36: 187–210.
  51. 51. Ksepka DT, Boyd CA. Quantifying historical trends in the completeness of the fossil record and the contributing factors: an example using Aves. Paleobiology. 2012; 38: 112–125.
  52. 52. Jablonski D, Roy K, Valentine JW, Price RM, Anderson PS. The impact of the pull of the recent on the history of marine diversity. Science. 2003; 300: 1133–1135. pmid:12750517
  53. 53. Labandeira CC. The fossil record of insect extinction: new approaches and future directions. Am Entomol. 2005; 51: 14–29.
  54. 54. Signor PW, Lipps JH. Sampling bias, gradual extinction patterns and catastrophes in the fossil record. Geol Soc Am Spec Pap. 1982; 190: 291–296.
  55. 55. Brusatte SL, Benton MJ, Ruta M, Lloyd GT. 2008 Superiority, competition, and opportunism in the evolutionary radiation of dinosaurs. Science. 2008; 321: 1485–1488. pmid:18787166
  56. 56. Alroy J. Are Sepkoski’s evolutionary faunas dynamically coherent? Evol Ecol Res. 2004; 6: 1–32.
  57. 57. Labandeira CC, Johnson KR, Wilf P. Impact of the terminal Cretaceous event on plant-insect associations. Proc Natl Acad Sci USA. 2002; 99: 2061–2066. pmid:11854501
  58. 58. Wappler T, Currano ED, Wilf P, Rust J, Labandeira CC. No post-Cretaceous ecosystem depression in European forests? Rich insect-feeding damage on diverse middle Palaeocene plants, Menat, France. Proc R Soc B Biol Sci. 2009; 276: 4271–4277.
  59. 59. Raup DM, Sepkoski JJ Jr. Mass extinctions in the marine fossil record. Science. 1982; 215: 1501–1503. pmid:17788674
  60. 60. Benton MJ. Diversification and extinction in the history of life. Science. 1995; 268: 52–58. pmid:7701342
  61. 61. Hannisdal B, Peters SE. 2011 Phanerozoic Earth system evolution and marine biodiversity. Science. 2011; 334: 1121–1124. pmid:22116884
  62. 62. Labandeira C. Why Did Terrestrial Insect Diversity Not Increase During the Angiosperm Radiation? Mid-Mesozoic, Plant-Associated Insect Lineages Harbor Clues. In: Pontarotti P, editor. Evolutionary Biology: Genome Evolution, Speciation, Coevolution and Origin of Life; 2014. pp. 261–299.
  63. 63. Benton MJ. When life nearly died: the greatest mass extinction of all time. London, New York: Thames & Hudson; 2003.
  64. 64. Alroy J. Dynamics of origination and extinction in the marine fossil record. Proc Natl Acad Sci USA. 2008; 105: 11536–11542.
  65. 65. Alroy J, Aberhan M, Bottjer DJ, Foote M, Fürsich FT, Harries PJ, et al. Phanerozoic trends in the global diversity of marine invertebrates. Science. 2008; 321: 97–100. pmid:18599780
  66. 66. Mayhew PJ. Shifts in hexapod diversification and what Haldane could have said. Proc R Soc B Biol Sci. 2002;269:969–974.
  67. 67. Benton MJ. The Red Queen and the Court Jester: species diversity and the role of biotic and abiotic factors through time. Science. 2009;323:728–732. pmid:19197051
  68. 68. Mayhew PJ, Bell MA, Benton TG, McGowan AJ. Biodiversity tracks temperature over time. Proc Natl Acad Sci USA. 2012;109:15141–15145. pmid:22949697
  69. 69. Hammer Ø, Harper DAT, Ryan PD. PAST: Paleontological statistics software package for education and data analysis. Palaeontol Electron. 2001;4:1–9.
  70. 70. Kaddumi HF. Amber of Jordan, the Oldest Prehistoric Insects in Fossilized Resin. Publications of the Eternal River Museum of Natural History, Amman; 2005.
  71. 71. Lubbock JW. Notes on the Thysanura. Part IV. Transactions of the Linnean Society of London. 1871;27:277–297.
  72. 72. Delclòs X, Arillo A, Peñalver E, Barrón E, Soriano C, López-Del-Valle R, et al. Fossiliferous amber deposits from the Cretaceous (Albian) of Spain. Comptes Rendus Palevol. 2007;6(1–2):135–149.
  73. 73. Christiansen K, Pike EM. Cretaceous Collembola (Arthropoda, Hexapoda) from the Upper Cretaceous of Canada. Cretaceous Research. 2002;23(2):165–188.
  74. 74. Riek EF. An entomobryid collembolan (Hexapoda: Collembola) from the Lower Permian of Southern Africa. Paleontologica Africana. 1976;19:141–143.
  75. 75. Christiansen K, Nascimbene P. Collembola (Arthropoda, Hexapoda) from the mid Cretaceous of Myanmar (Burma). Cretaceous Research. 2006;27:318–363.
  76. 76. Ross AJ, York PV. A catalogue of the type and figured specimens of Hexapoda from the Rhynie chert (early Devonian) at The Natural History Museum, London, UK. Transactions of the Royal Society of Edinburgh: Earth Sciences. 2004;94(4):319–395.
  77. 77. McKellar RC, Wolfe AP, Tappert R, Muehlenbachs K. Correlation of Grassy Lake and Cedar Lake ambers using infrared spectroscopy, stable isotopes, and palaeoentomology. Canadian Journal of Earth Sciences. 2008;45(9):1061–1082.
  78. 78. Börner C. Zur Systematik der Hexapoden. Zoologischer Anzeiger. 1904;27:511–533.
  79. 79. Weitschat W, Wichard W. Atlas of plants and animals in Baltic amber. Verlag Dr. Friedrich Pfeil, Mnchen; 2002.
  80. 80. Poinar GO. Life in amber. Stanford University Press, Standford, California; 1992.
  81. 81. Wilson HM, Martill DM. A new japygid dipluran from the Lower Cretaceous of Brazil. Palaeontology. 2001;44(5):1025–1031.
  82. 82. Kukalová-Peck J. New Carboniferous Diplura, Monura, and Thysanura, the hexapod ground plan, and the role of thoracic side lobes in the origin of wings (Insecta). Canadian Journal of Geology. 1987;65:2327–2345.
  83. 83. Engel MS. A new Lower Permian bristletail from the Wellington Formation in Kansas (Archaeognatha: Dasyleptidae). Transactions of the Kansas Academy of Science. 2009;112(1/2):40–44.
  84. 84. Rasnitsyn AP. Taxonomy and morphology of Dasyleptus Brongniart, 1885, with description of a new species (Insecta: Machilida: Dasyleptidae). Russian Entomological Journal. 2000;8[for 1999](3):145–154.
  85. 85. Rasnitsyn AP, Ross AJ. A preliminary list of arthropod families present in the Burmese amber collection at The Natural History Museum, London. Bulletin of The Natural History Museum, Geology Series. 2000;56(1):21–24.
  86. 86. Sturm H, Poinar GO. Cretaceomachilis libanensis, the oldest known bristle-tail of the family Meinertellidae (Machiloidea, Archaeognatha, Insecta) from the Lebanese amber. Deutsche entomologische Zeitschrift. 1998;45(1):43–48.
  87. 87. Bitsch J, Nel A. Morphology and classification of the extinct Archaeognatha and related taxa (Hexapoda). Annales de la Société entomologique de France. 1999;35(1):17–29.
  88. 88. Sinitshenkova ND. A review of Triassic mayflies, with a description of new species from western Siberia and Ukraine (Ephemerida = Ephemeroptera). Paleontological Journal. 2000;34(Suppl. 3):S275–S283.
  89. 89. Kluge NJ. A new suborder of Thysanura for the Carboniferous insect originally described as larva of Bojophlebia, with comments on characters of the orders Thysanura and Ephemeroptera. Zoosystematica Rossica. 1996;4(1):71–75.
  90. 90. Engel MS. A note of the relic silverfish Tricholepidion gertschi (Zygentoma). Transactions of the Kansas Academy of Science. 2006;109(3/4):236–238.
  91. 91. Rasnitsyn AP. Subclass Lepismatona Latreille, 1804. The wingless insects (= Thysanura Latreille 1796, s.l.). In: Rasnitsyn AP, Quicke DLJ, editors. History of Insects. Kluwer Academic Publishers, The Netherlands; 2002. p. 69–74.
  92. 92. Staniczek AH, Bechly G. 11.2 Apterygota: primarily wingless insects. In: Martill DM, Bechly G, Loveridge RF, editors. The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge University Press; 2007. p. 149–154.
  93. 93. Mendes LF, Poinar GO. A new fossil Nicoletiidae (Zygentoma, "Apterygota") in Dominican amber. Proceedings of the Entomological Society of Washington. 2004;106(1):102–109.
  94. 94. Novokshonov VG. New fossil insects (Insecta: Grylloblattida, ordinis incertis) from the Lower Permian of the middle Urals. Paleontological Journal. 2000;34(5):513–518.
  95. 95. Béthoux O, Nel A, Lapeyrie J, Gand G. New data on Paleozoic grylloblattid insects (Neoptera). Journal of Paleontology. 2005;79(1):125–138.
  96. 96. Carpenter FM. Superclass Hexapoda. In: Treatise on Invertebrate Paleontology, Part R, Arthropoda 4 (3&4). Boulder, C. O. and Lawrence, K. A.: Geological Society of America and University of Kansas Press; 1992. p. xxi + 655.
  97. 97. Hong YC. Hebeigramma nom. nov., a new name for Mesogramma Hong, 1984 (Caloneurodea) from the Lower Cretaceous of Hebei Province, China. Geological Bulletin of China. 2003;22(9):686–687.
  98. 98. Rasnitsyn AP. 2.2.1.2.1.2. Order Caloneurida Handlirsch, 1906 (= Caloneurodea Martynov, 1938). In: Rasnitsyn AP, Quicke DLJ, editors. History of Insects. Kluwer Academic Publishers, The Netherlands; 2002. p. 106–108.
  99. 99. Rasnitsyn AP. 2.2 Subclass Scarabaeona Laicharting, 1781. The winged insects (= Pterygota Lang, 1888). In: Rasnitsyn AP, Quicke DLJ, editors. History of Insects. Kluwer Academic Publishers, The Netherlands; 2002. p. 75–82.
  100. 100. Brauckmann C, Hahn G. Ein neuer Insektenfund aus dem Westfalium von Ibbenbüren (Westdeutschland). Paläontologische Zeitschrift. 1980;54(3–4):301–312.
  101. 101. Novokshonov VG. New fossil insects (Insecta: Grylloblattida, Caloneurodea, Hypoperlida?, ordinis incertis) from the Kungurian beds of the middle Urals. Paleontological Journal. 1998;32(4):362–368.
  102. 102. Novokshonov VG. New fossil insects (Insecta: Hypoperlida, Panorpida, ordinis incertis) from the Chekarda locality. Paleontological Journal. 1999;33(1):52–56.
  103. 103. Beckemeyer RJ. The Permian insect fossils of Elmo, Kansas. The Kansas School Naturalist. 2000;46(1):1–16.
  104. 104. Sinitshenkova ND. 2.2.1.2.3 Superorder Dictyoneuridea Handlirsch, 1906 (= Palaeodictyopteroidea). In: Rasnitsyn AP, Quicke DLJ, editors. History of Insects. Kluwer Academic Publishers, The Netherlands; 2002. p. 115–124.
  105. 105. Simon E. Essai d’une classification des Opiliones Mecostethi. Annales de la Société entomologique de Belgique. 1879;22:183–241.
  106. 106. Béthoux O, Nel A, Lapeyrie J. The extinct order Caloneurodea (Insecta: Pterygota: Panorthoptera): wing venation, systematics and phylogenetic relationships. Annales zoologici. 2004;54(2):289–318.
  107. 107. White RD. A type catalog of fossil invertebrates (Arthropoda: Hexapoda) in the Yale Peabody Museum. Postilla. 1995;209:1–55.
  108. 108. Sinitshenkova ND, Marchal-Papier F, Grauvogel-Stamm L, Gall JC. The Ephemeridea (Insecta) from the Grès à Voltzia (early Middle Triassic) of the Vosges (NE France). Paläontologische Zeitschrift. 2005;79(3):377–397.
  109. 109. Hyatt A, Arms JM. Guides for Science-Teaching, no. 8: Insecta. Boston; 1890.
  110. 110. McCafferty WP. Chapter 2. Ephemeroptera. In: Grimaldi DA, editor. Insects from the Santana Formation, Lower Cretaceous, of Brazil. vol. 195. Bulletin of the American Museum of Natural History; 1990. p. 25–50.
  111. 111. McCafferty WP. Higher classification of the burrowing mayflies (Ephemeroptera: Scapphodonta). Entomological News. 2004;115:84–92.
  112. 112. Kluge NJ. The phylogenetic system of Ephemeroptera. Kluwer Academic Publishers, The Netherlands; 2004.
  113. 113. Sinitshenkova ND. The first fossil prosopistomatid mayfly from Burmese amber (Ephemeroptera; Prosopistomatidae). Bulletin of The Natural History Museum, Geology Series. 2000;56(1):25–28.
  114. 114. Godunko RJ, Kłonowska-Olejnik M. The first fossil representative of the genus Analetris Edmunds, 1972 (Insecta: Ephemeroptera: Acanthametropodidae) from the Eocene Baltic amber. Annales zoologici. 2006;56(4):785–790.
  115. 115. Hubbard MD. Ephemeroptera. Fossilium Catalogus 1: Animalia. 1987;129:1–99.
  116. 116. McCafferty WP. Toward a phylogenetic classification of the Ephemeroptera (Insecta): a commentary on systematics. Annals of the Entomological Society of America. 1991;84(4):343–360.
  117. 117. Godunko RJ, Neumann C, Krzeminski W. Fossil mayfly collections of the Museum für Naturkunde, Humboldt University, Berlin. II. Redescription of Baltameletus oligocaenicus Demoulin, 1968 with notes on Ameletidae McCafferty, 1991 (Insecta: Ephemeroptera) from the Eocene Baltic amber. Annales zoologici. 2008;58(1):105–114.
  118. 118. Wichard W, Gröhn C, Seredszus F. Aquatic insects in Baltic amber, Wasserinsketen in Baltischen Bernstein. Verlag Kessel; 2009.
  119. 119. Sinitshenkova ND. New Jersey amber mayflies: the first North American Mesozoic members of the order (Insecta; Ephemeroptera. In: Grimaldi DA, editor. Studies on Fossils in Amber, with Particular Reference to the Cretaceous of New Jersey. Backhuys Publishers, Leiden, The Netherlands; 2000. p. 111–125.
  120. 120. McCafferty WP, Santiago-Blay JA. A new Cretaceous mayfly from Burmese amber (Ephemeroptera: Australiphemeridae). Entomological News. 2009;119(5):492–496.
  121. 121. Kluge NJ. New data on mayflies (Ephemeroptera) from Mesozoic and Cenozoic resins. Paleontological Journal. 1994;27(1A):35–49.
  122. 122. McCafferty WP. Discovery and analysis of the oldest mayflies (Insecta, Ephemeroptera) known from amber. Bulletin de la Société d’Histoire Naturelle de Toulouse. 1997;133:77–82.
  123. 123. Kluge NJ, Godunko RJ, Krzeminski W. A new mayfly family (Insecta: Ephemeroptera) from Eocene Baltic amber. Annales zoologici. 2006;56(1):181–185.
  124. 124. Zamboni JC. Contribution to the knowledge of the aquatic paleoentomofauna from Santana Formation (Araripe Basin, Lower Cretaceous, northeast Brazil) with description of new taxa. Acta Geologica Leopoldensia. 2001;24(52/53):129–135.
  125. 125. Staniczek AH. 11.4 Ephemeroptera: mayflies. In: The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge University Press; 2007. p. 163–184.
  126. 126. Pescador ML, Richard BA, Hubbard MD, Staniczek AH. Evolution of Baetiscidae (Ephemeroptera): current state of knowledge of the family. Aquatic Insects. 2009;31(Supplement 1):137–147.
  127. 127. Kukalová-Peck J. Ephemeroid wing venation based upon new gigantic Carboniferous mayflies and basic morphology, phylogeny and metamorphosis of pterygote insects (Insecta, Ephemerida). Canadian Journal of Zoology. 1985;63:933–955.
  128. 128. Wootton RJ, Kukalová-Peck J. Flight adaptations in Palaeozoic Palaeoptera (Insecta). Biological Review. 2000;75:129–167.
  129. 129. Sinitshenkova ND. Main ecological events in aquatic insects history. Acta zoologica cracoviensia. 2003;46(suppl.—Fossil Insects):381–392.
  130. 130. Sinitshenkova ND. New late Mesozoic mayflies from the Shar-Teeg locality, Mongolia (Insecta, Ephemerida = Ephemeroptera). Paleontological Journal. 2002;36(3):270–276.
  131. 131. Zhang JF, Kluge NJ. Jurassic larvae of mayflies (Ephemeroptera) from the Daohugou Formation in Inner Mongolia, China. Oriental Insects. 2007;41:351–366.
  132. 132. Jacobus LM, McCafferty WP. Revision of Ephemerellidae genera (Ephemeroptera). Transactions of the American Entomological Society. 2008;134(1/2):185–274.
  133. 133. Huang JD, Ren D, Sun JH. Progress in the study of Ephemeroptera (mayfly) fossils. Acta Zootaxonomica Sinica. 2007;32(2):391–404.
  134. 134. Kluge NJ, Sinitshenkova ND. 2.2.1.1.1.3 Order Ephemerida Latreille, 1810. The true mayflies (= Ephemeroptera Hyatt et Arms, 1891 (s.l.); = Euephemeroptera Kluge, 2000). In: Rasnitsyn AP, Quicke DLJ, editors. History of Insects. Kluwer Academic Publishers, The Netherlands; 2002. p. 89–97.
  135. 135. Peñalver E, Martínez-Delclòs X, Arillo A. Yacimientos con insectos fósiles en España. Revista Española de Paleontología. 1999;14(2):231–245.
  136. 136. Ren D, Gao KQ, Guo ZG, Ji S, Tan JJ, Song Z. Stratigraphic division of the Jurassic in the Daohugou area, Ningcheng, Inner Mongolia. Geological Bulletin of China. 2002;21(8–9):584–591. In Chinese with English summary.
  137. 137. Ogden TH, Gattolliat JL, Sartori M, Staniczek AH, Soldán T, Whiting MF. Towards a new paradigm in mayfly phylogeny (Ephemeroptera): combined analysis of morphological and molecular data. Systematic Entomology. 2009;34(4):616–634.
  138. 138. Lewis SE. Two new species of fossil mayflies (Ephemeroptera: Neoephemeridae and Siphlonuridae) from the Ruby River Basin (Oligocene) of southwestern Montana. Proceedings of the Entomological Society of Washington. 1977;79(4):583–587.
  139. 139. Godunko RJ, Krzeminski W. New fossil findings of the mayfly genera Balticobaetisca Staniczek & Bechly, 2002 (Ephemeroptera: Baetiscidae) and Borinquena Traver, 1938 (Leptophlebiidae: Atalophlebiinae). Aquatic Insects. 2009;31(Supplement 1):125–136.
  140. 140. Godunko RJ, Neumann C. Fossil mayfly collections of the Museum für Naturkunde, Humboldt University Berlin. I. Electroletus soldani gen. and sp. nov. (Ephemeroptera: Ameletidae) from the Eocene Baltic amber. Annales zoologici. 2006;56(1):175–180.
  141. 141. Novokshonov VG. New insects (Insecta) from the Lower Permian of Chekarda (Central Urals). Paleontological Journal. 1994;27 [for 1993](1A):172–178.
  142. 142. Novokshonov VG, Aristov DS. New and little-known Permian insects (Insecta: Grylloblattida; Orthoptera) from the Chekarda locality, Central Ural Mountains. Paleontological Journal. 2002;36(6):644–649.
  143. 143. Rasnitsyn AP. 2.2.1.1.1.2. Order Syntonopterida Handlirsch, 1911. In: Rasnitsyn AP, Quicke DLJ, editors. History of Insects. Kluwer Academic Publishers, The Netherlands; 2002. p. 88–89.
  144. 144. Kinzelbach R, Lutz H. Eine neue Eintagsfliege Misthodotes stapfi n. sp. aus dem Rotliegenden des Nahe-Gebietes (Ephemeroptera: Permoplectoptera: Misthodotidae). Paläontologische Zeitschrift. 1984;58(3/4):247–253.
  145. 145. Sinitshenkova ND. A new mayfly species of the extant genus Neoephemera from the Eocene of North America (Insecta: Ephemerida = Ephemeroptera: Neoephemeridae. Paleontological Journal. 1999;33(4):403–405.
  146. 146. Jacobus LM, McCafferty WP. Reevaluation of the phylogeny of the Ephemeroptera infraorder Pannota (Furcatergalia), with adjustments to higher classification. Transactions of the American Entomological Society. 2006;132(1/2):81–90.
  147. 147. Whalley PES, Jarzembowski EA. Fossil insects from the Lithographic Limestone of Montsech (late Jurassic-early Cretaceous), Lerida Province, Spain. Bulletin of the British Museum (Natural History), Geology. 1985;38(5):381–412.
  148. 148. Rowland JM. The late Paleozoic insect assemblage at Carrizo Arroyo, New Mexico p.1-7. In: Lucas SG, Estep JW, Williamson TE, Morgan GS, editors. New Mexico’s Fossil Record 1. vol. Bulletin 11. New Mexico Museum of Natural History and Science, Albuquerque; 1997.
  149. 149. Zhou CF, Peters JG. The nymph of Siphluriscus chinensis and additional imaginal description: a living mayfly with Jurassic origins (Siphluriscidae new family: Ephemeroptera). Florida Entomologist. 2003;86(3):345–352.
  150. 150. Zhang JF. New mayfly nymphs from the Jurassic of northern and northeastern China (Insecta: Ephemeroptera). Paleontological Journal. 2006;40(5):553–559.
  151. 151. Lin QB, Huang DY. New Middle Jurassic mayflies (Insecta: Ephemeroptera: Siphlonuridae) from Inner Mongolia, China. Annales zoologici. 2008;58(3):521–527.
  152. 152. Garrouste R, Nel A, Gand G. New fossil arthropods (Notostraca and Insecta: Syntonopterida) in the continental Middle Permian of Provence (Bas-Argens Basin, France). Comptes Rendus Palevol. 2009;8(1):49–57.
  153. 153. Krzeminski W, Lombardo C. New fossil Ephemeroptera and Coleoptera from the Ladinian (Middle Triassic) of Canton Ticino (Switzerland). Rivista Italiana di Paleontologia e Stratigrafia. 2001;107(1):69–78.
  154. 154. Sinitshenkova ND. New Mesozoic mayflies (Ephemerida) from Mongolia. Paleontological Journal. 1989;23(3):26–37.
  155. 155. Handlirsch A. Revision der paläozoischen Insekten. Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse. 1919;96:511–592.
  156. 156. Rasnitsyn AP, Aristov DS, Gorokhov AV, Rowland JM, Sinitshenkova ND. Important new insect fossils from Carrizo Arroyo and the Permo-Carboniferous faunal boundary. In: Lucas SG, Zeigler KE, editors. Carboniferous-Permian Transition at Carrizo Arroyo, Central New Mexico. New Mexico Museum of Natural History and Science, Albuquerque, Bulletin 25; 2004. p. 215–246.
  157. 157. van Dijk DE, Geertsema H. Permian insects from the Beaufort Group of Natal, South Africa. Annals of the Natal Museum. 1999;40:137–171.
  158. 158. Shcherbakov DE, Makarkin VN, Aristov DS, Vasilenko DV. Permian insects from the Russky Island, South Pimorye. Russian Entomological Journal. 2009;18(1):7–16.
  159. 159. Tasch P, Zimmerman JR. The Asthenohymen-Delopterum bed—A new Leonardian insect horizon in the Wellington of Kansas and Oklahoma. Journal of Paleontology. 1962;36:1319–1333.
  160. 160. Beckemeyer RJ, Engel MS. An enigmatic new genus of biarmohymenid from the early Permian Wellington Formation of Noble County, Oklahoma (Palaeodictyopterida: Diaphanopterodea). Transactions of the Kansas Academy of Science. 2009;112(1/2):103–108.
  161. 161. Béthoux O, Nel A. Revision of Diaphanoptera species and new diagnosis of Diaphanopteridae (Palaeoptera: Diaphanopterodea). Journal of Paleontology. 2003;77(5):1016–1020.
  162. 162. Zajíc J, Štamberg S. Selected important fossiliferous horizons of the Boskovice Basin in the light of the new zoopaleontological data. Acta Musei Reginaehradecensis Series A, Scientiae naturales. 2004;30:5–14.
  163. 163. Carpenter FM. Studies on Carboniferous insects from Commentry, France: Part V. The genus Diaphanoptera and the order Diaphanopterodea. Psyche. 1963;70:240–256.
  164. 164. Rohdendorf BB. Fundamentals of Paleontology Volume 9: Arthropoda, Tracheata, Chelicerta. Orlov YA, editor. Smithsonian Institution Libraries and The National Science Foundation, Washington D.C.; 1991.
  165. 165. Sinitshenkova ND. The peculiar “dipterous" insects from the Permian of Arkhangelsk district and Urals (Hypoperlida, Eukulojidae). Bulletin of Moscow Society of Naturalists, Biological Series. 1981;56(3):116–126. In Russian.
  166. 166. Béthoux O, Nel A, Lapeyrie J, Gand G, Galtier J. New Martynoviidae from the Permian of Southern France (Lodève basin) (Insecta: Palaeoptera: Diaphanopterodea). Geobios. 2003;36:131–139.
  167. 167. Brauckmann C, Herd KJ. Insekten-Funde aus dem Westfalium D (Ober-Karbon) des Piesberges bei Osnabrück (Deutschland). Teil 1: Palaeoptera. Osnabrücker naturwissenschafliche Mitteilungen. 2003;28 (for 2002):27–69.
  168. 168. Kukalová-Peck J, Sinitshenkova ND. The wing venation and systematics of Lower Permian Diaphanopterodea from the Ural Mountains, Russia (Insecta: Paleoptera). Canadian Journal of Zoology. 1992;70:229–235.
  169. 169. Kukalová-Peck J, Brauckmann C. Wing folding in pterygote insects, and the oldest Diaphanopterodea from the early Late Carboniferous of West Germany. Canadian Journal of Zoology. 1990;68:1104–1111.
  170. 170. Carpenter FM. 14A Insecta. In: Shabica CW, Hay AA, editors. Richardson’s Guide to The Fossil Fauna of Mazon Creek. Northeastern Illinois University; 1997. p. 184–193.
  171. 171. Prokop J, Nel A. Systematic position of Triplosoba, hitherto the oldest mayfly, from Upper Carboniferous of Commentry in Central France (Insecta: Palaeodictyopterida). Systematic Entomology. 2009;34(4):610–615.
  172. 172. Brongniart C. Les insectes fossiles des terrains primaires. Coup d’oeil rapide sur la faune entomologique des terrains paléozoïques. Bulletin de la Société des amis des sciences naturelles de Rouen. 1885;1885:50–68.
  173. 173. Kukalová-Peck J. Megasecoptera from the Lower Permian of Moravia. Psyche. 1975;82:1–19.
  174. 174. Carpenter FM. A megasecopteran from Upper Carboniferous strata in Spain. Psyche. 1963;70:44–49.
  175. 175. Brauckmann C. Notiz über Insekten-Reste aus dem Ober-Karbon in Spanien. Jahresberichte des Naturwissenschaften Vereins in Wuppertal. 1993;46:115–119.
  176. 176. Sinitshenkova ND. A new family Aykhalidae from the Upper Palaeozoic of Yakutia Sakha (Insecta: Mischopteridae = Megasecoptera). Paleontological Journal. 1994;27(1993)(1A):131–134.
  177. 177. Brauckmann C, Schöllmann L, Sippel W. Die fossilen Insekten, Spinnentiere und Eurypteriden von Hagen-Vorhalle. Geologie und Paläontologie in Westfalen. 2003;59:5–89.
  178. 178. Zessin W. Zwei neue Insektenreste (Megasecoptera, Odonatoptera) aus dem Westfalium D (Oberkarbon) des Piesberges bei Osnabrück, Deutschland. Virgo, Mitteilungsblatt des Entomologischen Vereins Mecklenburg. 2006;9(1):37–45.
  179. 179. Nelson CR, Tidwell WD. Brodioptera stricklani n. sp. (Megasecoptera: Brodiopteridae), a new fossil insect from the Upper Manning Canyon Shale Formation, Utah (Lowermost Namurian B). Psyche. 1987;94:309–316.
  180. 180. Brauckmann C. Ein neuer Inseckten-Rest (Megasecoptera) aus dem Ober-Karbon von Osnabrück. Osnabrücker naturwissenschafliche Mitteilungen. 1991;17:25–32.
  181. 181. Béthoux O, Galtier J, Nel A. Earliest evidence of insect endophytic oviposition. Palaios. 2004;19(4):408–413.
  182. 182. Carpenter FM. Studies on Carboniferous insects from Commentry, France: Part II. The Megasecoptera. Journal of Paleontology. 1951;25(3):336–355.
  183. 183. Béthoux O, McBride J, Maul C. Surface laser scanning of fossil insect wings. Palaeontology. 2004;47(1):13–19.
  184. 184. Labandeira CC. 1.3.9 Rise and Diversification of Insects. In: Briggs DEG, Crowther PR, editors. Palaeobiology II. Blackwell Science, London; 2001. p. 82–88.
  185. 185. Prokop J, Ren D. New significant fossil insects from the Upper Carboniferous of Ningxia in northern China (Palaeodictyoptera, Archaeorthoptera). European Journal of Entomology. 2007;104(2):267–275.
  186. 186. Béthoux O, Nel A. Révision de Protagrion audouini Brongniart, 1893, du Carbonifère supérieur (Palaeoptera). Bulletin de la Société entomologique de France. 2003;108(3):237–244.
  187. 187. Hong YC. New fossil genera and species of Shanxi Formation in Xishan of Taiyuan. Entomotaxonomia. 1985;7(2):83–91.
  188. 188. Rasnitsyn AP, Sukatsheva ID, Aristov DS. Permian insects of the Vorkuta Group in the Pechora Basin, and their stratigraphic implications. Paleontological Journal. 2005;39(4):404–416.
  189. 189. Pinto ID. Sphecorydaloides lucchesei a new Carboniferous megasecopteran Insecta from Argentina. Pesquisas. 1994;21(2):85–89.
  190. 190. Pinto ID, Adami-Rodrigues K. A revision of South American Paleozoic insects. In: Scoggin M, editor. AMBA projects AM/PFICM98/1.99: Proceedings of the First International Palaeoentomological Conference, Moscow 1998; 1999. p. 117–124.
  191. 191. Novokshonov VG. New insects (Insecta: Hypoperlida, Mischopterida, Jurinida) from the Lower Permian of the Middle Urals. Paleontological Journal. 1998;32(1):46–53.
  192. 192. Pinto ID. Carboniferous insects from Argentina III Familia Xenopteridae Pinto, nov. Ordo Megasecoptera. Pesquisas. 1986;18:23–29.
  193. 193. Goldenberg CF. Fauna Saraepontana fossilis. Die fossilen Thiere aus der Steinkohlenformation von Saarbrücken, Part II. Möllinger: Saarbrücken; 1877.
  194. 194. Prokop J, Nel A. A new genus and species of Homoiopteridae from the Upper Carboniferous of the Intra-Sudetic Basin, Czech Republic (Insecta: Palaeodictyoptera). European Journal of Entomology. 2004;101(4):583–589.
  195. 195. Brauckmann C, Koch L, Kemper M. Spinnentiere (Arachnida) und Insekten aus den Vorhalle-Schichten (Namurium B; Ober-Karbon) von Hagen-Vorhalle (West Deutschland). Geologie und Paläontologie in Westfalen. 1985;3:1–131.
  196. 196. Béthoux O, Nel A, Schneider JW, Gand G. Lodetiella magnifica nov. gen. and nov. sp. (Insecta: Palaeodictyoptera; Permian), an extreme situation in wing morphology of palaeopterous insects. Geobios. 2007;40(2):181–189.
  197. 197. Özdikmen H. Some nomenclatural changes for Blattodea and Dictyoneurida (= Palaeodictyoptera). Munis Entomology and Zoology. 2008;3(2):745–748.
  198. 198. Gorjunova RV. New Carboniferous bryozoans of the Gobi Altai. Sovmestnaya Sovetsko-Mongolskaya Paleontologischeskaya Ekspeditsiya Trudy. 1988;33:10–23.
  199. 199. Sharov AG, Sinitshenkova ND. Novyye Paleodictyoptera s territorii SSSR. [New Palaeodictyoptera from the USSR.]. Paleontologicheskii Zhurnal. 1977;1977:48–63.
  200. 200. Prokop J, Smith R, Jarzembowski EA, Nel A. New homoiopterids from the Late Carboniferous of England (Insecta: Palaeodictyoptera). Comptes Rendus Palevol. 2006;5(7):867–873.
  201. 201. Handlirsch A. Revision of American Paleozoic insects. Proceedings of the United States National Museum. 1906;29:661–820.
  202. 202. Kukalová J. Revisional study of the order Palaeodictyoptera in the Upper Carboniferous Shales of Commentry, France. Part I. Psyche. 1969;76:163–215.
  203. 203. Kukalová J. Revisional study of the order Palaeodictyoptera in the Upper Carboniferous Shales of Commentry, France. Part II. Psyche. 1969;76:349–486.
  204. 204. Bolton H. A Monograph of the Fossil Insects of the British Coal Measures. Part I. Palaeontographical Society, London; 1921.
  205. 205. Jell PA. The fossil insects of Australia. Memoirs of the Queensland Museum. 2004;50(1):1–124.
  206. 206. Brauckmann C, Schneider J. Ein unter-karbonisches Insekt aus dem Raum Bitterfeld/Delitzsch (Pterygota, Arnsbergium, Deutschland). Neues Jahrbuch für Geologie und Paläontologie, Monatshefte. 1996;1996(1):17–30.
  207. 207. Laurentiaux-Vieira F, Laurentiaux D. Paleodictyoptere nouveau du Namurien belge. Annales de la Société Géologique du Nord. 1986;105:187–193.
  208. 208. Sinitshenkova ND. A new species of the Tchirkovaeidae (Insecta, Dictyoneurida) from the Upper Carboniferous of the Tunguska Basin. Paleontological Journal. 1981;15(1):121–123.
  209. 209. Brodsky AK. The evolution of insect flight. Oxford University Press; 1994.
  210. 210. Gutiérrez PR, Muzón J, Limarino CO. The earliest late Carboniferous winged insect (Insecta, Protodonata) from Argentina: geographical and stratigraphical location. Ameghiniana. 2000;37(3):375–378.
  211. 211. Fabricius JC. Entomologicae Systematica, v. 2. Hafniae; 1793.
  212. 212. Fleck G, Nel A, Bechly G, Martínez-Delclòs X. Revision and phylogenetic affinities of the Jurassic Steleopteridae Handlirsch, 1906 (Odonata: Zygoptera). Insect Systematics & Evolution. 2001;32:285–305.
  213. 213. Nel A, Paicheler JC. Les Odonata fossiles: état actuel des connaissances. Deuxième partie: Les Petaluridae et Cordulegastridae fossiles (Odonata, Anisoptera, Petaluroidea). Nouvelle Revue dEntomologie. 1992;9(4):305–323.
  214. 214. Fleck G, Nel A. Revision of the Mesozoic family Aeschnidiidae (Odonata: Anisoptera). Zoologica. 2003;153:1–172.
  215. 215. Nel A, Martínez-Delclòs X, Escuillié F, Brisac P. Les Aeshnidae fossils: Etat actuel des connaissances (Odonata, Anisoptera). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 1994;194(2/3):143–186.
  216. 216. Nel A, Bechly G, Jarzembowski EA, Martínez-Delclòs X. A revision of the fossil petalurid dragonflies (Insecta: Odonata: Anisoptera: Petalurida). Paleontologica Lombarda Nuova serie. 1998;10:1–68.
  217. 217. Bechly G, Nel A, Martínez-Delclòs X, Jarzembowski EA, Coram R, Martill D, et al. A revision and phylogenetic study of Mesozoic Aeshnoptera, with description of numerous new taxa (Insecta: Odonata: Anisoptera). Neue Paläontologische Abhandlungen. 2001;4:1–219.
  218. 218. Bechly G, Ueda K. The first fossil record and first New World record for the dragonfly clade Chlorogomphida (Insecta: Odonata: Anisoptera: Araripechlorogomphidae n. fam.) from the Crato Limestone (Lower Cretaceous, Brazil). Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie). 2002;328:1–11.
  219. 219. Bechly G. 11.5 Odonata: damselflies and dragonflies. In: Martill DM, Bechly G, Loveridge RF, editors. The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge University Press; 2007. p. 184–222.
  220. 220. Bechly G. Morphologische Untersuchungen am Flügelgeäder der rezenten Libellen und deren Stammgruppenvertreter (Insecta: Pterygota: Odonata), unter besonderer Berücksichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata. Petalura, Special Volume. 1996;2:1–402.
  221. 221. Fleck G, Nel A, Bechly G, Delclòs X, Jarzembowski EA, Coram RA. New Lower Cretaceous ‘libelluloid’ dragonflies (Insecta: Odonata: Cavilabiata) with notes about estimated divergence dates for this group. Palaeodiversity. 2008;1:19–36.
  222. 222. Bechly G. New fossil dragonflies from the Lower Cretaceous Crato Formation of northeast Brazil (Insecta: Odonata. Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie). 1998;264:1–66.
  223. 223. Jarzembowski EA. Chapter 10. Arthropods 2: Insects p.149–160. In: Swift A, Martill DM, editors. Fossils of the Rhaetian Penarth Group. The Palaeontological Association, London; 1999.
  224. 224. Sukatsheva ID, Rasnitsyn AP. Jurassic insects (Insecta) from the Sai-Sagul locality (Kyrgyzstan, Southern Fergana). Paleontological Journal. 2004;38(2):182–186.
  225. 225. Nel A, Paicheler JC, Henrotay M. Les “Anisozygoptera” fossiles. Phylogénie et classification (Odonata). Martinia. 1993;3:1–311.
  226. 226. Petrulevicius JF, Nel A. Austroperilestidae, a new family of damselflies from early Eocene of Argentina (Insecta: Odonata). Phylogenetic relationships within Odonata. Journal of Paleontology. 2005;79(4):658–662.
  227. 227. Nel A, Papier F, Grauvogel-Stamm L, Gall JC. Voltzialestes triasicus gen. nov., sp. nov., le premier Odonata Protozygoptera du Trias inférieur des Vosges (France). Paleontologica Lombarda Nuova serie. 1996;5:25–36.
  228. 228. Nel A, Gand G, Garric J, Lapeyrie J. The first recorded protozygopteran insects from the Upper Permian of France. Palaeontology. 1999;42(1):83–97.
  229. 229. Jarzembowski EA, Nel A. The earliest damselfly-like insect and the origin of modern dragonflies (Insecta: Odonatoptera: Protozygoptera). Proceedings of the Geologists’ Association. 2002;113:165–169.
  230. 230. Zessin W. Überblick über die paläozoischen Libellen (Insecta, Odonatoptera). Virgo, Mitteilungsblatt des Entomologischen Vereins Mecklenburg. 2008;11(1):5–32.
  231. 231. Gentilini G. Fossil damselflies and dragonflies from the Eocene of Monte Bolca, Italy (Insecta: Odonata). Studi e Ricerche sui Giaimenti Terziari di Bolca. 2002;9:7–22.
  232. 232. Fleck G, Waller A, Serafin J, Nel A. The oldest Calopterygidae in the Eocene Baltic amber (Odonata: Zygoptera). Zootaxa. 2009;1985:52–56.
  233. 233. Nel A, Bechly G, Delclòs X, Huang DY. New and poorly known Mesozoic damseldragonflies (Odonata: Isophlebioidea: Campterophlebiidae, Isophlebiidae). Palaeodiversity. 2009;2:209–232.
  234. 234. Huguet A, Nel A, Martínez-Delclòs X, Bechly G, Martins-Neto R. Preliminary phylogenetic analysis of the Protanisoptera (Insecta: Odonatoptera). Geobios. 2002;35(5):537–560.
  235. 235. Petrulevicius JF, Nel A. First Cordulephyidae dragonfly in America: A new genus and species from the Paleogene of Argentina (Insecta: Odonata). Comptes Rendus Palevol. 2009;8(4):385–388.
  236. 236. Bechly G. A new fossil dragonfly (Anisoptera: Corduliidae) from the Paleocene Fur Formation (Mo clay) of Denmark. Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie). 2005;358:1–7.
  237. 237. Jarzembowski EA, Martínez-Delclòs X, Bechly G, Nel A, Coram R, Escuillié F. The Mesozoic non-calopterygoid Zygoptera: description of new genera and species from the Lower Cretaceous of England and Brazil and their phylogenetic significance (Odonata, Zygoptera, Coenagrionoidea, Hemiphlebioidea, Lestoidea). Cretaceous Research. 1998;19(3–4):403–444.
  238. 238. Coram RA, Nel A. A new petalurid dragonfly from the Lower Cretaceous of southern England (Odonata: Petalurida:? Cretapetaluridae). Palaeodiversity. 2009;2:205–208.
  239. 239. Nel A, Bechly G. The third petalurid dragonfly from the Lower Cretaceous of Brazil (Odonata: Cretapetaluridae). Annales zoologici. 2009;59(3):281–285.
  240. 240. Bechly G. New fossil odonates from the Upper Triassic of Italy, with a redescription of Italophlebia gervasuttii Whalley, and a reclassification of Triassic dragonflies (Insecta: Odonata). Rivista del Museo Civico di Scienze Naturali “Enrico Caffi”. 1997;19:31–70.
  241. 241. Nel A, Huang DY. First Chinese Cymatophlebiidae from the Middle Jurassic of Inner Mongolia (Odonata: Anisoptera: Aeshnoptera). Palaeodiversity. 2009;2:199–204.
  242. 242. Rust J, Petrulevicius JF, Nel A. The first damselflies from the lowermost Eocene of Denmark, with a description of a new subfamily (Odonata, Zygoptera: Dysagrionidae). Paleontology. 2008;51(3):709–713.
  243. 243. Nel A, Arillo A. The first Baltic amber dysagrionine damselfly (Odonata: Zygoptera: Thaumatoneuridae: Dysagrioninae). Annales de la Société entomologique de France (Nouvelle série). 2006;42(2):179–182.
  244. 244. Nel A, Néraudeau D, Perrichot V, Girard V, Gomez B. A new dragonfly family from the Upper Cretaceous of France. Acta Palaeontologica Polonica. 2008;53(1):165–168.
  245. 245. Pritykina LN. Two new dragonflies from the Lower Cretaceous deposits of west Mongolia (Anisoptera: Sonidae fam. nov., Corduliidae). Odonatologica. 1986;15(2):169–184.
  246. 246. Nel A, Paicheler JC. Les Odonata fossiles: état actuel des connaissances. Huitième partie: Les Calopterygoidea fossiles (Odonata, Zygoptera). Bulletin de la Société entomologique de France. 1993;97(4):381–396.
  247. 247. Petrulevicius JF, Nel A, Rust J, Bechly G, Kohls D. New Paleogene Epallagidae (Insecta: Odonata) recorded in North America and Europe. Biogeographic implications. Alavesia. 2007;1:15–25.
  248. 248. Fleck G, Bechly G, Martínez-Delclòs X, Jarzembowski E, Coram R, Nel A. Phylogeny and classification of the Stenophlebioptera (Odonata: Epiproctophora). Annales de la Société entomologique de France (Nouvelle série). 2003;39(1):55–93.
  249. 249. Bridges CA. Catalogue of the family-group, genus-group and species-group names of the Odonata of the world (3rd edition). Urbana, Illinois, USA; 1994.
  250. 250. Jarzembowski EA. Early Cretaceous zygopteroids of southern England, with the description of Cretacoenagrion alleni gen. nov., spec. nov. (Zygoptera: Coenagrionidae; “Anisozygoptera”: Tarsophlebiidae, Euthemistidae). Odonatologica. 1990;19(1):27–37.
  251. 251. Petrulevicius JF, Nel A. Frenguelliidae, a new family of dragonflies from the earliest Eocene of Argentina (Insecta: Odonata): phylogenetic relationships within Odonata. Journal of Natural History. 2003;37(24):2909–2917.
  252. 252. Petrulevicius JF, Nel A. Enigmatic and little known Odonata (Insecta) from the Paleogene of Patagonia and northwest Argentina. Annales de la Société entomologique de France (Nouvelle série). 2007;43(3):341–347.
  253. 253. Prokop J, Fikacek M. An annotated list of early Oligocene insect fauna from Seifhennersdorf (Saxony, Germany). Acta Musei Nationalis Pragae, Series B Historia Naturalis. 2007;63(2–4):209–217.
  254. 254. Schlüter T. Fossil insects in Gondwana—localities and palaeodiversity trends. Acta zoologicacracoviensia. 2003;46(suppl.- Fossil Insects):345–371.
  255. 255. Bechly G, Nel A, Martínez-Delclòs X, Fleck G. Four new dragonflies from the Upper Jurassic of Germany and the Lower Cretaceous of Mongolia (Anisoptera: Hemeroscopidae, Sonidae, and Proterogomphidae fam. nov.). Odonatologica. 1998;27(2):149–187.
  256. 256. Ren D, Liu JY, Cheng XD. A new hemeroscopid dragonfly from the Lower Cretaceous of Northeast China (Odonata: Hemeroscopidae). Acta Entomologica Sinica. 2003;46(5):622–628.
  257. 257. Vasilenko DV. New damselflies (Odonata: Synlestidae, Hemiphlebiidae) from the Mesozoic Transbaikalian locality of Chernovski Kopi. Paleontological Journal. 2005;39(3):280–283.
  258. 258. Zalessky G. Two new Permian dragonfly-like insects of the order Permodonata [in Russian]. Doklady Akademii Nauk SSSR. 1955;104(4):630–633.
  259. 259. Ansorge J. Heterophlebia buckmani (Brodie 1845) (Odonata: "Anisozygoptera")–das erste Insekt aus dem untertoarcischen Posidonienschiefer von Holzmaden (Wüttemberg, SW Deutschland). Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie). 1999;275:1–9.
  260. 260. Bechly G, Wichard W. Damselfly and dragonfly nymphs in Eocene Baltic amber (Insecta: Odonata), with aspects of their palaeobiology. Palaeodiversity. 2008;1:37–73.
  261. 261. Nel A, Petrulevicius JF, Jarzembowski EA. New fossil Odonata from the European Cenozoic (Insecta: Odonata: Thaumatoneuridae, Aeshnidae,? Idionychidae, Libellulidae). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 2005;235(3):343–380.
  262. 262. Pritykina LN. Isophlebiid dragonflies from the late Mesozoic of eastern Transbaikalia (Odonata: Isophlebiidae). Paleontological Journal. 2006;40(6):636–645.
  263. 263. Bybee SM, Ogden TH, Branham MA, Whiting MF. Molecules, morphology and fossils: a comprehensive approach to odonate phylogeny and the evolution of the odonate wing. Cladistics. 2008;24:477–514.
  264. 264. Bechly G. Juracordulia schiemenzi gen. et. sp. nov., Eine neue Libelle aus den
  265. 265. Solnhofener Plattenkalken (Insecta: Odonata: Anisoptera). Archaeopteryx. 1998;16:29–36.
  266. 266. Nel A, Bechly G, Martínez-Delclòs X, Fleck G. A new family of Anisoptera from the Upper Jurassic of Karatau in Kazakhstan (Insecta: Odonata: Juragomphidae n. fam.). Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie). 2001;314:1–9.
  267. 267. Huang DY, Nel A. Oldest ‘libelluloid’ dragonfly from the Middle Jurassic of China (Odonata: Anisoptera: Cavilabiata). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 2007;246(1):63–68.
  268. 268. Nel A, Béthoux O, Bechly G, Martínez-Delclòs X, Papier F. The Permo-Triassic Odonatoptera of the “Protodonate” grade (Insecta: Odonatoptera). Annales de la Société entomologique de France (Nouvelle série). 2001;37(4):501–525.
  269. 269. Petrulevicius JF, Nel A. A new damselfly family from the Upper Palaeocene of Argentina. Palaeontology. 2004;47(1):109–116.
  270. 270. Nel A, Paicheler JC. Les Lestoidea (Odonata, Zygoptera) fossils: un inventaire critique. Annales de Paléontologie. 1994;80(1):1–59.
  271. 271. Huang DY, Nel A, Lin QB. A new genus and species of aeshnopteran dragonfly from the Lower Cretaceous of China. Cretaceous Research. 2003;24(2):141–147.
  272. 272. Etter W, Kuhn O. An articulated dragonfly (Insecta, Odonata) from the Upper Liassic Posidonia Shale of northern Switzerland. Palaeontology. 2000;43(5):967–977.
  273. 273. Nel A, Petrulevicius JF. A new genus and species of damsel-dragonfy from the early Liassic of Germany (Odonata, Liassophlebiidae). Bulletin de la Société entomologique de France. 2005;110(2):185–188.
  274. 274. Nel A, Paicheler JC. Les Libelluloidea autres ue Libellulidae fossils. Un inventaire critique (Odonata, Corduliidae, Macromiidae, Synthemistidae, Chlorogomphidae et Mesophlebiidae). Nouvelle Revue d’Entomologie. 1994;11(4):321–334.
  275. 275. Fleck G, Nel A, Martínez-Delclòs X. The oldest record of libellulid dragonflies from the Upper Cretaceous of Kazakhstan (Insecta: Odonata, Anisoptera). Cretaceous Research. 1999;20:655–658.
  276. 276. Bechly G. Two new fossil dragonfly species (Insecta: Odonata: Anisoptera: Araripegomphidae and Lindeniidae) from the Crato Limestone (Lower Cretaceous, Brazil). Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie). 2000;296:1–16.
  277. 277. Azar D, Nel A. First Baltic amber megapodagrionid damselfly (Odonata: Zygoptera). Annales de la Société entomologique de France (Nouvelle série). 2008;44(4):451–457.
  278. 278. Bechly G. Description of a new species of Nannogomphus (Insecta: Odonata: Nannogomphidae) from the Upper Jurassic Solenhofen Limestone in Germany. Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie). 2003;339:1–6.
  279. 279. Lin QB, Huang DY, Nel A. A new family of Cavilabiata from the Lower Cretaceous Yixian Formation, China (Odonata: Anisoptera). Zootaxa. 2007;1469:59–64.
  280. 280. Carle FL. Evolution, taxonomy, and biogeography of ancient Gondwanian libelluloides, with comments on anisopteroid evolution and phylogenetic systematics (Anisoptera: Libelluloidea). Odonatologica. 1995;24(4):383–424.
  281. 281. Petrulevicius JF, Nel A, Muzón J. A new libelluloid family from the upper Palaeocene of Argentina. Palaeontology. 1999;42(4):677–682.
  282. 282. Petrulevicius JF, Nel A. New palaeomacromiid dragonflies from the upper Palaeocene of Argentina. Palaeontology. 2002;45(4):751–758.
  283. 283. Nel A. A new Odonata family from the Jurassic of Central Asia (Odonata: Epiproctophora). Journal of Natural History. 2009;43(1–2):57–64.
  284. 284. Bechly G. A re-description of “Stenophlebiacasta (Insecta: Odonata: Parastenophlebiidae n. fam.) from the Upper Jurassic Solenhofen Limestone in Germany. Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie). 2005;359:1–12.
  285. 285. Vasilenko DV, Rasnitsyn AP. Fossil ovipositions of dragonflies: review and interpretation. Paleontological Journal. 2007;41(11):1156–1161.
  286. 286. Petrulevicius JF, Nel A. Oldest petalurid dragonfly (Insecta: Odonata): a Lower Cretaceous specimen from south Patagonia, Argentina. Cretaceous Research. 2003;24(1):31–34.
  287. 287. Nel A. Piroutetia liasina Meunier, 1907, Insecte du Lias de France, espèce-type des Piroutetiidae nov. fam. (Odonatoptera, Meganeurina). Bulletin du Muséum National d’Histoire Naturelle, Série 4, Section C. 1989;11(1):15–19.
  288. 288. Wappler T, Petrulevicius JF. Priscalestidae, a new damselfly family (Odonata: Lestinoidea) from the Middle Eocene Eckfeld maar of Germany. Alavesia. 2007;1:69–73.
  289. 289. Nel A, Bechly G, Martínez-Delclòs X. A new fossil dragonfly from the Upper Jurassic in Germany [Odonata, Anisoptera, Protolindeniidae]. Revue française d’Entomologie. 2001;23(4):257–261.
  290. 290. Nel A, Petrulevicius JF, Martínez-Delclòs X. New Mesozoic Protomyrmeleontidae (Insecta: Odonatoptera: Archizygoptera) from Asia with a new phylogenetic analysis. Journal of Systematic Palaeontology. 2005;3(2):187–201.
  291. 291. Nel A, Jarzembowski EA. New protomyrmeleontid dragonflies from the Lower Cretaceous of southern England (Insecta, Odonata, Archizygoptera). Cretaceous Research. 1998;19(3–4):393–402.
  292. 292. Lin QB, Zhang S, Huang DY. Fuxiaeschna hsiufnia gen. nov., spec. nov., a new Lower Cretaceous dragonfly from northwestern China (Aeshnoptera: Rudiaeschnidae). Odonatologica. 2004;33(4):437–442.
  293. 293. Nel A, Gand G, Fleck G, Béthoux O, Lapeyrie J, Garric J. Saxonagrion minutus nov. gen. et sp., the oldest damselfly from the Upper Permian of France (Odonatoptera, Panodonata, Saxonagrionidae nov. fam.). Geobios. 1999;32(6):883–888.
  294. 294. Zessin W. Eine unwahrscheinliche Erfolgsbilanz: die Evolution der Libellen. Virgo, Mitteilungsblatt des Entomologischen Vereins Mecklenburg. 2005;8(1):54–66.
  295. 295. Nel A, Petrulevicius JF, Gentilini G, Martínez-Delclòs X. Phylogenetic analysis of the Cenozoic family Sieblosiidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios. 2005;38(2):219–233.
  296. 296. Nel A, Marie V, Schmeißner S. Revision of the lower Mesozoic dragonfly family Triassolestidae Tillyard, 1918 (Odonata: Epiproctophora). Annales de Paléontologie. 2002;88:189–214.
  297. 297. Fleck G, Bechly G, Martínez-Delclòs X, Jarzembowski EA, Nel A. A revision of the Upper Jurassic-Lower Cretaceous dragonfly family Tarsophlebiidae, with a discussion on the phylogenetic positions of the Tarsophlebiidae and Sieblosiidae (Insecta, Odonatoptera, Panodonata). Geodiversitas. 2004;26(1):33–60.
  298. 298. Huang DY, Nel A. The first Chinese Tarsophlebiidae from the Lower Cretaceous Yixian Formation, with morphological and phylogenetic implications (Odonatoptera: Panodonata). Cretaceous Research. 2009;30(2):429–433.
  299. 299. Martins-Neto RG, Gallego OF, Zavattieri AM. A new Triassic insect fauna from Cerro Bayo, Potrerillos (Mendoza Province, Argentina) with descriptions of new taxa (Insecta: Blattoptera and Coleoptera). Alcheringa. 2007;31(2):199–213.
  300. 300. Jarzembowski EA, Nel A. New fossil dragonflies from the Lower Cretaceous of SE England and the phylogeny of the superfamily Libelluloidea (Insecta: Odonata). Cretaceous Research. 1996;17:67–85.
  301. 301. Bechly G. New fossil damselflies from Baltic amber, with description of a new species, a redescription of Litheuphaea carpenteri Fraser, and a discussion on the phylogeny of Epallagidae (Zygoptera: Caloptera). International Journal of Odonatology. 1998;1(1):33–63.
  302. 302. Brongniart C. Recherches pour servir à l’histoire des insectes fossiles des temps primaires, précédées d’une étude sur la nervation des ailes des insectes. Thèse présentée à la Fauclté des Sciences de Paris. 1893;821:495.
  303. 303. Nel A, Huguet A. Revision of the enigmatic Upper Carboniferous insect Campyloptera eatoni Brongniart, 1893 (Insecta: Odonatoptera). Organisms Diversity & Evolution. 2002;2(4):313–318.
  304. 304. Nel A, Fleck G, Garrouste R, Gand G, Lapeyrie J, Bybee SM, et al. Revision of Permo-Carboniferous griffenflies (Insecta: Odonatoptera: Meganisoptera) based upon new species and redescription of selected poorly known taxa from Eurasia. Palaeontographica Abteilung A. 2009 September;289(4–6):89–121.
  305. 305. Nel A, Gand G, Garric J. A new family of Odonatoptera from the continental Upper Permian: The Lapeyriidae (Lodève Basin, France). Geobios. 1999;32(1):63–72.
  306. 306. Béthoux O. The insect fauna from the Permian of Lodève (Hérault, France): state of the art and perspectives. Journal of Iberian Geology. 2008;34(1):109–113.
  307. 307. Ren D, Nel A, Prokop J. New early griffenfly, Sinomeganeura huangheensis from the late Carboniferous of northern China (Meganisoptera: Meganeuridae). Insect Systematics & Evolution. 2008;39:223–229.
  308. 308. Rehn AC. Phylogenetic analysis of higher-level relationships of Odonata. Systematic Entomology. 2003;28(2):181–239.
  309. 309. Martynov AV. Über eine neue Ordnung der fossilen Insekten, Miomoptera nov. Zoologischer Anzeiger. 1927;72:99–109.
  310. 310. Rasnitsyn AP. 2.2.1.3.1. Superorder Palaeomanteidea Handlirsch, 1906. Order Palaeomanteida Handlirsch, 1906 (= Miomoptera Martynov, 1927). In: Rasnitsyn AP, Quicke DLJ, editors. History of Insects. Kluwer Academic Publishers, The Netherlands; 2002. p. 161–164.
  311. 311. Rasnitsyn AP. On skimming hypothesis of the insect flight origin. Acta zoologica cracoviensia. 2003;46(suppl.- Fossil Insects):85–88.
  312. 312. Prokop J, Nel A. An enigmatic Palaeozoic stem-group: Paoliida, designation of new taxa from the Upper Carboniferous of the Czech Republic (Insecta: Paoliidae, Katerinkidae fam. n.). African Invertebrates. 2007;48(1):77–86.
  313. 313. Brunner von Wattenwyl K. Prodromus der europäischen orthoptèren. Engelmann, Leipzig; 1882.
  314. 314. Vršanskyý P. Phyloblatta grimaldii sp. nov.—a new Triassic cockroach (Insecta: Blattaria) from Virginia. Entomological Problems. 2003;32(1–2):51–53.
  315. 315. Shcherbakov DE. Madygen, Triassic Lagerstätte number one, before and after Sharov. Alavesia. 2008;2:113–124.
  316. 316. Martins-Neto RG, Mancuso AC, Gallego OF. The Triassic insect fauna from Argentina. Blattoptera from Los Rastros Formation (Bermejo Basin) La Rioja province. Ameghiniana. 2005;42(4):705–723.
  317. 317. Liang JH, Ren D, Ye QP, Liu M, Meng XM. The fossil Blattaria of China: a review of present knowledge. Acta Zootaxonomica Sinica. 2006;31(1):102–108.
  318. 318. Lin QB. Mesozoic insects from Zhejiang and Anhui provinces, China. In: of Geology NI, Palaeontology, editors. Division and correlation of the Mesozoic volcano-sedimentary formations in the provinces of Zhejiang and Anhui. Science Press, Beijing; 1980. p. 211–234.
  319. 319. Lin QB. Cretaceous insects of China. Cretaceous Research. 1994;15:305–316.
  320. 320. Li XH, Chen S, Cao K, Chen YH, Xu BL, Ji Y. Paleosols of the Mid-Cretaceous: a report from Zhejiang and Fujian, SE China. Earth Science Frontiers. 2009;16(5):63–70.
  321. 321. Bechly G. 11.8 ‘Blattaria’: cockroaches and roachoids. In: Martill DM, Bechly G, Loveridge RF, editors. The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge University Press; 2007. p. 239–249.
  322. 322. Béthoux O, Klass KD, Schneider JW. Tackling the Protoblattoidea problem: Revision of Protoblattinopsis stubblefieldi (Dictyoptera; late Carboniferous). European Journal of Entomology. 2009;106:145–152.
  323. 323. Hörnschemeyer T, Stapf H. Review of Blattinopsidae (Protorthoptera) with description of new species from the Lower Permian of Niedermoschel (Germany). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 2001;221(3):81–132.
  324. 324. Rasnitsyn AP. 2.2.1.2.1.1. Order Blattinopseida Bolton, 1925. In: Rasnitsyn AP, Quicke DLJ, editors. History of Insects. Kluwer Academic Publishers, The Netherlands; 2002. p.106.
  325. 325. Béthoux O, Nel A. Venation pattern and revision of Orthoptera sensu nov. and sister groups. Phylogeny of Palaeozoic and Mesozoic Orthoptera sensu nov. Zootaxa. 2002;96:1–88.
  326. 326. Cifuentes-Ruiz P, Vršanskyý P, Vega FJ, Cevallos-Ferriz SRS, González-Soriano E, Delgado de Jesús CR. Campanian terrestrial arthropods from the Cerro del Pueblo Formation, Difunta Group in northeastern Mexico. Geologica Carpathica. 2006;57(5):347–354.
  327. 327. Vršanskyý P. Decreasing variability—from the Carboniferous to the present! (Validated on independent lineages of Blattaria). Paleontological Journal. 2000;34(Suppl. 3):S374–S379.
  328. 328. Vršanskyý P, Ansorge J. Lower Jurassic cockroaches (Insecta: Blattaria) from Germany and England. African Invertebrates. 2007;48(1):103–126.
  329. 329. Vršanskyý P, Vishniakova VN, Rasnitsyn AP. 2.2.2.1.1. Order Blattida Latreille, 1810. The cockroaches (= Blattodea Brunner von Wattenvill, 1882). In: Rasnitsyn AP, Quicke DLJ, editors. History of Insects. Kluwer Academic Publishers, The Netherlands; 2002. p. 263–270.
  330. 330. Vršanskyý P, Ansorge J. New Lower Cretaceous polyphagid cockroaches from Spain (Blattaria, Polyphagidae, Vitisminae subfam. nov.). Cretaceous Research. 2001;22(2):157–162.
  331. 331. Handlirsch A. Neue Untersuchungen über die fossilen Insekten mit Ergänzungen und Nachträgen sowie Ausblicken auf phylogenetische, palaeogeographische und allgemein biologische Probleme. I Teil. Annalen des Naturhistorischen Museums in Wien. 1937;48:1–140.
  332. 332. Clifford E, Coram RA, Jarzembowski EA, Ross AJ. A supplement to the insect fauna from the Purbeck Group of Dorset. Proceedings of the Dorset Natural History and Archaeological Society. 1994;115:143–146.
  333. 333. Vršanskyý P. Albian cockroaches (Insecta, Blattida) from French amber of Archingeay. Geodiversitas. 2009;31(1):73–98.
  334. 334. Liang JH, Vršanskyý P, Ren D, Shih CK. A new Jurassic carnivorous cockroach (Insecta, Blattaria, Raphidiomimidae) from the Inner Mongolia in China. Zootaxa. 2009;1974:17–30.
  335. 335. Ross AJ. The cockroaches (Blattodea) of the Purbeck Limestone Group and Wealden Supergroup (Lower Cretaceous) of southern England p.59-60. In: 2nd International Congress on Palaeoentomology, Krakow, abstract volume; 2001.
  336. 336. Vršanskyý P, Liang JH, Ren D. Advanced morphology and behaviour of extinct earwig-like cockroaches (Blattida: Fuziidae fam. nov.). Geologica Carpathica. 2009;60(6):449–462.
  337. 337. Vršanskyý P. Origin and the early evolution of mantises. AMBA projekty. 2002;6(1):1–16.
  338. 338. Vršanskyý P. Mesozoic relative of the common synanthropic German cockroach (Blattodea). Deutsche entomologische Zeitschrift. 2008;55(2):215–221.
  339. 339. Jarzembowski EA, Schneider JW. The stratigraphical potential of blattodean insects from the late Carboniferous of southern Britain. Geological Magazine. 2007;144(3):449–456.
  340. 340. Li Z, Hong YC, Yang D. A new Middle Triassic genus and species of Mylacridae (Blattodea) from China. Zootaxa. 2007;1660:53–59.
  341. 341. Schneider J. Die Blattodea (Insecta) des Paläzoikums, Teil 1: Systematik, Ökologie und Biostratigraphie. Freiberger Forschungenshefte, Reihe C. 1983;382:106–145.
  342. 342. Hong YC. The discovery of Late Palaeozoic insecta in Shanxi Province. Geological Review. 1980;26(2):89–95.
  343. 343. Zhang ZL, Sun KQ, Yin JR. Sedimentology and sequence stratigraphy of the Shanxi Formation (Lower Permian) in the northwestern Ordos Basin, China: an alternative sequence model for fluvial strata. Sedimentary Geology. 1997;112:123–136.
  344. 344. Hong YC. Establishment of fossil entomofaunas and their evolutionary succession in north China. Entomologia Sinica. 1998;5(4):283–300.
  345. 345. Vršanskyý P. New blattarians and a review of dictyopteran assemblages from the Lower Cretaceous of Mongolia. Acta Palaeontologica Polonica. 2008;53(1):129–136.
  346. 346. Schneider JW, Lucas SG, Rowland JM. The Blattida (Insecta) fauna of Carrizo Arroyo, New Mexico—biostratigraphic link between marine and nonmarine Pennsylvanian/Permian boundary profiles. In: Lucas SG, Zeigler KE, editors. Carboniferous-Permian Transition at Carrizo Arroyo, Central New Mexico. vol. Bulletin 25. New Mexico Museum of Natural History and Science, Albuquerque; 2004. p. 247–261.
  347. 347. Schneider J. Die Blattodea (Insecta) des Palaozoikums Teil II: Morphogenese der Flugelstrukturen und Phylogenie. Freiberger Forschungenshefte, Reihe C. 1984;391:5–34.
  348. 348. Vršanskyý P. Jumping cockroaches (Blattaria, Skokidae fam. n.) from the late Jurassic of Karatau in Kazakhstan. Biologia. 2007;62(5):588–592.
  349. 349. Schneider JW, Werneburg R. Insect biostratigraphy of the Euramerican continental late Pennsylvanian and early Permian. In: Lucas SG, Cassinis G, Schneider JW, editors. Non-Marine Permian Biostratigraphy and Biochronology. Geological Society of London Special Publications 265; 2006. p. 325–336.
  350. 350. Papier F, Nel A. Les Subioblattidae (Blattodea, Insecta) du Trias d’Asie Centrale. Paläontologische Zeitschrift. 2001;74(4):533–542.
  351. 351. Gorokhov AV. New and little known orthopteroid insects (Polyneoptera) from fossil resins: Communication 1. Paleontological Journal. 2006;40(6):646–654.
  352. 352. Chen SX, Tan JJ. A new family of Coleoptera from the Lower Cretaceous of Kansu. Acta Entomologica Sinica. 1973;16(2):169–178.
  353. 353. Vršanskyý P. Umenocoleoidea—an amazing lineage of aberrant insects (Insecta, Blattaria). AMBA projekty. 2003;7(1):1–32.
  354. 354. Perrichot V, Néraudeau D, Nel A, de Ploëg G. A reassessment of the Cretaceous amber deposits from France and their palaeontological significance. African Invertebrates. 2007;48(1):213–227.
  355. 355. Beckemeyer RJ. Ligogramma wichita, a new species of Caloneurodea (Polyneoptera: Orthopterida) from the Lower Permian Wellington Formation of Noble County, Oklahoma. Journal of the Kansas Entomological Society. 2009;82(4):300–304.
  356. 356. Aristov DS. A new family of the order Grylloblattida (Insecta) from the Middle Permian of Russia. Paleontological Journal. 2009;43(2):178–182.
  357. 357. Béthoux O, Nel A, Lapeyrie J, Gand G. The Permostridulidae fam. n. (Panorthoptera), a new enigmatic insect family from the Upper Permian of France. European Journal of Entomology. 2003;100:581–585.
  358. 358. Martins-Neto RG, Gallego OF, Brauckmann C, Cruz JL. A review of the South American Palaeozoic entomofauna part I: the Ischnoneuroidea and Cacurgoidea, with description of new taxa. African Invertebrates. 2007;48(1):87–101.
  359. 359. Béthoux O. Revision and phylogenetic affinities of the lobeattid species bronsoni Dana, 1864 and silvatica Laurentiaux & Laurentiaux-Vieira, 1980 (Pennsylvanian; Archaeorthoptera). Arthropod Systematics & Phylogeny. 2008;66(2):145–163.
  360. 360. Béthoux O, Nel A. Some Palaeozoic ‘Protorthoptera’ are ‘ancestral’ orthopteroids: major wing braces as clues to a new split among the ‘Protorthoptera’ (Insecta). Journal of Systematic Palaeontology. 2005;2 [for 2004](4):285–309.
  361. 361. Béthoux O, Nel A, Galtier J, Lapeyrie J, Gand G. A new species of Tococladidae Carpenter, 1966 from the Permian of France (Insecta: Archaeorthoptera). Geobios. 2003;36(3):275–283.
  362. 362. Rasnitsyn AP. 2.2.2. Infraclass Gryllones Laicharting, 1781. The Grylloneans (= Polyneoptera Martynov, 1938). In: Rasnitsyn AP, Quicke DLJ, editors. History of Insects. Kluwer Academic Publishers, The Netherlands; 2002. p. 254–262.
  363. 363. Martins-Neto RG. Estágio atual da paleoartropodologia brasileira: Hexápodes, Miriápodes, Crustáceos (Isopoda, Decapoda, Eucrustacea e Copepoda) e quelicerados. Arquivos do Museu Nacional, Rio de Janeiro. 2005;63(3):471–494.
  364. 364. Béthoux O. Ordinal assignment of the genus Tococladus Carpenter 1996 (Insecta: Archaeorthoptera). Alavesia. 2007;1:3.
  365. 365. Rasnitsyn AP. 2.2.1.2.2. Superorder Hypoperlidea Martynov, 1928. Order Hypoperlida Martynov, 1928. In: Rasnitsyn AP, Quicke DLJ, editors. History of Insects. Kluwer Academic Publishers, The Netherlands; 2002. p. 111–115.
  366. 366. de Geer C. Mémoires pour servir à l’histoire des insectes, v. 3. Stockholm; 1773.
  367. 367. Engel MS, Haas F. Family-group names for earwigs (Dermaptera). American Museum Novitates. 2007;3567:1–20.
  368. 368. Haas F. 11.6 Dermaptera: earwigs. In: Martill DM, Bechly G, Loveridge RF, editors. The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge University Press; 2007. p. 222–234.
  369. 369. Wappler T, Engel MS, Haas F. The earwigs (Dermaptera: Forficulidae) from the middle Eocene Eckfeld maar, Germany. Polskie Pismo Entomologiczne. 2005;74(3):227–250.
  370. 370. Zhang JF. The most primitive fossil earwigs (Archidermaptera, Dermaptera, Insecta) from the Upper Jurassic of Nei Mongol Autonomous Region, northeastern China. Acta Micropalaeontologica Sinica. 2002;19(4):348–362.
  371. 371. Spahr U. Ergänzungen und Berichtigungen zu R. Keilbach’s Bibliographie und Liste der Bernsteinfossilien–“Apterygota”. Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie). 1990;166:1–23.
  372. 372. Engel MS, Grimaldi DA. A primitive earwig in Cretaceous amber from Myanmar (Dermaptera: Pygidicranidae). Journal of Paleontology. 2004;78(5):1018–1023.
  373. 373. Engel MS. The earwigs of Kansas, with a key to gener