by Roy J. Beckemeyer (Johnston Geology Museum, Emporia State University, Emporia, Kansas, USA) and Joseph D. Hall  (Department of Geology, Wichita State University, Wichita, Kansas, USA)

Last updated 17 July 2005

A paper based on the presentation made at the FossilsX3 Congress in Pretoria, South Africa, February, 2005:

  ABSTRACT/INTRO/LOCATION/HISTORY

REVIEW OF INSECTS: Species DIversity / Size Diversity / Nymphal vs Adult Specimens / Specimen Quality 

 CONCLUSIONS/ACKNOWLEDGMENTS/REFERENCES

Abstract:  

The Lower Permian Wellington Formation fossil beds of mid-continent North America are known best for the famous Elmo, Kansas locality discovered by E.H. Sellards in the early 1900’s.  The Elmo site has produced tens of thousands of specimens from which more than 150 species of insects have been described.  Equally productive and more widespread geographically, but less well-known, are the Midco, Oklahoma beds located some 270 km south of Elmo, that were discovered by G.O. Raasch in the late 1930’s.  The Midco beds have also yielded tens of thousands of specimens, but the material has been less well studied, and to date only half as many species have been identified from the Oklahoma localities.  Although little research has been done on the sites or their entomological fauna since the last published work on them by the late F.M. Carpenter in the late 1970’s, renewed attention has been given to both the geology and paleontology of the Wellington Formation in recent years by workers in the region.  The history of these insect beds is recounted and the insect faunal composition is briefly reviewed:  There are nearly 200 species in 106 genera, 53 families and 21 orders. Sizes (as measured by mean fore wing length) range from 1.9 mm to 330 mm, with a mean of 22 mm and a median value of 12 mm. Ten of 13 species with forewings greater than 50 mm in length were Protodonata.  Most species are known from one or a few specimens.  Specimen abundance ranges from 1 to just under 400 specimens per species. Of 10 species for which 50 or more specimens are known, eight are in Grylloblattida (and six of these in Grylloblattida: Lemmatophorina), indicating that these taxa were either quite abundant or were preferentially preserved, or both.  When reviewing the holotype/neotype specimens used to describe the Wellington Formation species, we find that sixty-two percent were specimens of forewings, while nine percent were complete specimens.  However, when considering all known specimens, forty-eight percent of the species are known only by their fore wings, while thirteen percent are now represented by complete specimens, indicating the importance of continued collecting and review of Wellington Formation insect fossils.

Key Words:  Artinskian, Elmo, fossil insects, Kansas, Lower Permian, Midco, Oklahoma, Wellington Formation

INTRODUCTION

The Lower Permian Elmo, Kansas North American locality (Artinskian Age) is an important Paleozoic fossil insect Konservat-Lagerstätt.  Approximately 150 species of insects have been described from these beds, which were discovered just over a hundred years ago.  The Elmo locality insect-bearing strata are found within the Wellington Formation, which extends south from Elmo another 270 km to the vicinity of the city of Perry, Oklahoma, where the Midco fossil insect beds occur in a number of outcrops.  The Midco locality, discovered nearly a half century after Elmo, is less well studied; only half as many species are known from Midco.  In the area between the Elmo and Midco localities the Wellington Formation is covered by quaternary loess and alluvium and is exposed only in a few isolated locations so that the only significant fossil insect beds are those at Elmo and Midco.  Altogether approximately 200 species are known from the mid-continent Wellington deposits of Kansas and Oklahoma.

LOCATION AND EXTENT

The Wellington Formation occurs close to the geographic center of the contiguous United States in North America.  It extends from north-central Kansas south through north-central Oklahoma (Fig. 1).  The insect bearing deposits lie along what was in Permian times a coastal plain or marginal marine environment (Dunbar & Tillyard, 1924, Carpenter, 1947, Raasch, 1946).  The Elmo beds are considered to have been fresh water ponds or lakes while the Midco beds were thought to have been saline, possibly playa lakes (Carpenter, 1947, Tasch, 1964).  Hall’s (2004) recent reassessment of the Oklahoma Wellington Formation stratigraphy assigns the Midco beds to a marginal marine environment.  The insects preserved as fossils were likely allochthonous, and were either blown in or washed into the embayment by streams (Hall, 2004).

  Figure 1.  The location and extent of the North American mid-contintent Lower Permian Wellington Formation insect fossil deposits. The deposits are located near the geographic center of the contiguous United States, in the states of Kansas and Oklahoma.  The formation runs north to south some 270 km along 8 counties, with major sites at the northern (Elmo, Kansas) and southern (Midco, Oklahoma) extremities.

HISTORY OF DISCOVERY

Elias H. Sellards (Fig. 2a) was studying Paleozoic plant fossils under Samuel Williston at Kansas University in 1899 when he found two fossil insects among a collection of plant fossils from the Wellington Formation.  He was interested enough to make trips to the locality, which was located near the small settlement of Elmo, Kansas (Fig. 3).  He eventually collected some two thousand insect specimens.  After completing a Ph.D. at Yale University in 1903, he began the study of the Wellington Formation material, eventually describing many species, 27 of which are today recognized as valid.  Sellards kept most of his specimens, but deposited a small representative collection in the Yale Peabody Museum.

Figure 2.  a. Elias H. Sellards (1875-1961) discovered the Elmo, Kansas fossil beds.  Photo courtesy of the Texas Memorial Museum.  b.  Robin J. Tillyard (1881-1937) studied Elmo fossils for a period of 16 years, carrying out his research in New Zealand and Australia.  Eventually all the fossil specimens he worked with were returned to the Yale Peabody Museum.  Photo courtesy of the late Faith Tillyard Evans, Tillyard’s daughter.  c. Frank M. Carpenter (1902-1994), of the Harvard Museum of Comparative Geology and the dean of American paleoentomologists, published definitive studies of the Elmo specimens over a period of some 60 years.  Photo courtesy of George Byers, University of Kansas.

Figure 3.  The settlement of Elmo, Kansas, as it appeared in the early 1900’s.  The fossil beds, located a few miles southeast, were named for the town.  Photo courtesy of the Wichita State University Libraries Department of Special Collections.

In 1920, the distinguished entomologist Robin J. Tillyard (Fig. 2b), then residing in New Zealand and working at the Cawthron Institute in Nelson, was visiting Yale on a trip through the United States.  He was quite excited to see a series of Sellards’ fossil specimens in the Peabody Museum.  His enthusiasm led Charles Schuchert and Carl Dunbar of Yale to obtain funding and embark on a collecting expedition to the Elmo beds.  Another 2000 specimens were gathered.  These were shipped to Tillyard in New Zealand, and he embarked on a sixteen-year study of the material.  Tillyard described fifty-five valid species of Elmo insects.

Frank M. Carpenter (Fig. 2c) began to work on Elmo fossil specimens in 1925.  He published his last papers in the late 1970’s.  He did prolific and definitive work on the Elmo fossils, including revisions of taxa established by Sellards and Tillyard.  Later, he also published on the Midco palaeopterous specimens (Carpenter, 1947, 1979).  Carpenter named a total of 96 valid species from Elmo and Midco.

Figure 4.  a. Gilbert O. Raasch (1903-1999) discovered the Oklahoma Midco fossil insect beds and documented the stratigraphy of the Oklahoma Wellington Formation in his Ph.D. thesis (Raasch, 1946).  Photo courtesy University of Wisconsin-Madison.  b. Paul Tasch of Wichita State University studied the paleolimnology of the Wellington Formation in Kansas and Oklahoma and discovered additional Wellington Formation fossil insect localities in the late 1950’s and early 1960’s.  Photo courtesy of Wichita State University Department of Special Collections, Wichita, Kansas.

In the late 1930’s, Gilbert O. Raasch (Fig. 4a) was working on a Ph.D. in geology, studying the stratigraphy of the Wellington Formation in Oklahoma (Raasch, 1946).  He discovered a number of localities that were quite rich in fossil insect material, and contacted Frank Carpenter.  In 1940 the two men collected some 5,000 specimens in ten weeks of field time.  Subsequent collecting by Carpenter brought the total number of specimens from Midco in the Harvard Museum of Comparative Zoology collection to about 8,000 (Carpenter, 1979).  Raasch eventually switched his interests to Cambrian trilobites of the Upper Mississippi Valley.

During the late 1950’s and early 1960’s, Paul Tasch (Fig. 4b), of Wichita State University, began working on the paleolimnology of the Wellington Formation in Kansas and Oklahoma.    He discovered insect fossils at a number of sites, and his associate, James Zimmerman, an entomologist from Wichita State University, eventually named eight species from the Oklahoma material (Tasch and Zimmerman, 1962).

During the 1960’s, Jurmilla Kukalová-Peck studied a number of Wellington Formation taxa as part of her research on Paleozoic insects.  She reviewed the nymphal Ephemeroptera fossils collected by Carpenter from the Midco beds, determining them as the Wellington Formation genus Protereisma, but did not assign any specific names (Kukalová-Peck, 1968).  However, in 1970, Demoulin, based solely on Kukalová-Peck’s 1968 paper, assigned generic and specific names to her specimens.  Hubbard and Kukalová-Peck, 1980, and Carpenter (1979, 1992) eventually corrected Demoulin’s work.  The only valid Midco species named by Demoulin is Protereisma americana (Demoulin, 1970). 

A complete list of references to the Wellington Formation publications of the above researchers and a more detailed history of the Elmo locality can be found in Beckemeyer (2000).

REVIEW OF THE ENTOMOFAUNA

Species Diversity

A total of 194 valid insect species are currently known from the Wellington Formation.  One hundred eighteen (118, 61%) are known only from Elmo, 43 (22%) are exclusive to Midco, and 33 (17%) are common to both localities (Fig. 5).  The lower number of species unique to Midco reflects the fact that Carpenter published only on the palaeopterous insects of Midco and not on the Neoptera (Fig. 6), and that there has to this day not been a published review or survey of Midco Neoptera.  

Figure 5.  Proportion of the Wellington Formation entomofauna unique to or shared between the Elmo, Kansas and Midco, Oklahoma localities. 

Figure 6.  Proportion of Wellington Formation entomofauna Neoptera and Palaeoptera subgroups unique to or shared between the Elmo, Kansas and Midco, Oklahoma localities, illustrating that the greater percentage of unique fauna at Elmo arises from the Neoptera.  This reflects the fact that no review of the Neoptera from Midco has yet been published.

Carpenter’s last paper on the Wellington Formation insects appeared in 1979, and little has been added to the knowledge of this fauna until a very recent renewal of interest during the last few years.  Since 1998 seven new Wellington Formation species have been described (Table 1) and seven species previously known from Elmo have been found to occur at Midco as well.  Undoubtedly, many more species will be added upon a complete review of the neopterous material from Midco.

Table 1.  New species added to the Wellington Entomofauna since the publication of Carpenter’s Treatise on Invertebrate Paleontology Hexapoda Volumes (1992).

In Figure 7 we compare the Elmo entomofauna distribution by major group (Holometabolous Neoptera, Paraneoptera, Polyneoptera, Palaeoptera, Apterygota) with the extant World insect fauna (Species numbers for modern Orders were taken from Grimaldi & Engel, 2005).  Insects with complete metamorphosis accounted for only about 7% of Elmo species, while today they are the most numerous insects, amounting to approximately 83% of described species.  Palaeoptera today account for less than one percent of the extant insect fauna while for Elmo, the group comprised nearly 27% of the species. 

Figure 7.  Comparison of the distribution of the Elmo, Kansas Permian and World extant entomofaunas by major groups: Apterygota, Palaeoptera, Polyneoptera, Paraneoptera, and Holometabola.  Only about 6% of the fossil fauna were holometabolous while this group comprises 83% of the extant insects.  Also, some 27% of the Elmo insects as compared to less than 1% of the extant fauna are palaeopterous.

Figure 8 is a bar chart showing species diversity by major taxon (Order in most cases, Order: Suborder for the Grylloblattida) for Elmo and Midco.  The ordinal system used is that of Carpenter (1992), with some modifications.  Carpenter’s “Protorthoptera” are placed in the Grylloblattida, Hypoperlida, and Blattinopseida, following Rasnitsyn & Quicke (2002).  Classification of the Grylloblattida suborders follows Storozhenko (1998, 2002).  Carpenter’s Homoptera are grouped with the Hemiptera.  The family Lophioneuridae is considered a stem group of Thysanoptera, after Kukalová-Peck (1991).  The data are grouped into Apterygota, Pterygota: Palaeoptera and Pterygota: Neoptera sections, with the taxa arranged in each section in order of decreasing species diversity.  

Figure 8.  Bar chart illustrating species diversity of Wellington Formation entomofauna by Order (Suborder for the Grylloblattida).  Orders are placed in three groups, Apterygota, Pterygota: Palaeoptera, and Pterygota: Neoptera, and are arranged in decreasing order of number of species.  Bars show the number of species in each taxon unique to Elmo, Midco and common to both.  The nearly 200 Wellington Formation species represent some 21 orders, 53 families, and 106 genera.

A nearly complete species list of Elmo insects may be found in Beckemeyer (2000).  The Wellington entomofauna is quite diverse with 21 orders, 53 families, and 106 genera represented.  Table 2 lists the known collections that contain Wellington Formation insect fossils.  The Yale Peabody Museum and Harvard Museum of Comparative Zoology are the major repositories for these fossils, but there are a number of smaller collections as well.

Table 2.  Repositories of collections of Elmo and/or Midco Lower Permian Wellington Formation insect fossils.

Size Diversity

The common view of Paleozoic insects is that they were quite large, and, indeed, the largest insect known to have flown, the meganeurid Meganeuropsis permiana Carpenter, 1939, was described from the Elmo deposits.  However, most of the insects of the Wellington formation were small.  In Figure 9 we have plotted the average fore wing lengths in mm of the Wellington insect species from smallest to largest.  Figure 10 contains a bar chart showing the frequency of distribution of forewing lengths in 5 mm size classes.  The charts show that most of the Wellington Formation insects had fore wings less than 25 mm in length.  The mean forewing length was 22 mm, the median (half the species had forewings less than this, half more) length 12 mm, and the mode (the forewing length shared by most of the species) was 6 mm. 

Figure 9.  Distribution of size in the Wellington Formation entomofauna.  Size is based on average forewing length in mm, plotted from smallest species to largest species.  The 13 largest (forewing length > 100 mm) and 2 smallest (forewing length < 2 mm) species are identified.  The two smallest species are both in family Lophioneuridae.  Ten of the largest species are in order Protodonata.  Moravia grandis is in order Palaeodictyoptera, Gigagramma carpenteri is in order Caloneurodea, and Sylvohymen ingens is in order Megasecoptera. 

Figure 10.  Frequency distribution in 5 mm size classes of average forewing lengths for the Wellington Formation entomofauna. 

We have noted the species names of the insects with forewing lengths equal to or greater than 50 mm and those with forewing lengths less than or equal to 2 mm on Figure 9.  Ten of the 13 insects with forewings of 50 mm or more were Protodonata.  Moravia grandis is a palaeodictyopteran (Calvertiellidae), Gigagramma carpenteri is in order Caloneurodea, and Sylvohymen ingens is in order Megasecoptera.  These are all extinct orders.  The two species with the smallest forewings, 1.9 mm in length, are both in the Lophioneuridae, a family considered to be in a primitive stem group of the extant Thysanoptera. 

Figure 11.  Size distributions of Wellington Formation entomofauna by major taxonomic groups.  Size based on average forewing length in mm.

Specimen Abundance

In Figure 12 we have plotted the number of adult specimens known per species in increasing order for the Elmo entomofauna.  Figure 13 contains the frequency distribution data for 10-specimen size classes.  The mean is 21 specimens, but the median is 2 and the mode 1.  The high mean is due to a half dozen species for which large numbers of specimens have been found.  Species for which more than a hundred specimens are known are identified on Figures 12 and 13.  Of these 6 species, 4 (Lemmatophora typa, Artinska clara, A. ovata, and Lisca minuta) are in Grylloblattida: Lemmatophorina: Lemmatophoridae.  One, Probnis speciosa, is in Grylloblattida: Protoperlina: Probnidae, and the last is Psocoptera: Psocidiidae: Dichentomum tinctum.  Of the four species with more than 50 and less than 100 specimens, one (52 specimens) is Ephemeroptera: Protereismatidae: Protereisma permianum, another (86 specimens) is Grylloblattida: Grylloblattina: Liomopteridae: Liomopterum ornatum and the other two are lemmatophorids (66 and 64 specimens, respectively): Lecorium elongatum and Paraprisca fragilis.  Five of the six species with more than a hundred specimens and eight of the ten species with more than fifty specimens are Grylloblattida.  Thus the grylloblattids were either quite abundant in the Elmo Permian, or they were preferentially-preserved, or both.

Figure 12.  Abundance of adult specimens of Elmo entomofauna.  Values are plotted from species with fewest known specimens to species with the most known specimens.  The species with the greatest numbers of specimens are identified.  All labeled taxa are Grylloblattida except for Dichentomum tinctum, which is in order Psocoptera.

Figure 13.  Frequency distribution of adult-specimen abundance in 10-specimen size classes for the Elmo entomofauna.

Figure 14.  Adult-specimen abundance distributions of the Elmo entomofauna by major taxonomic groups.  Taxa are arranged in order of groups with greatest species abundance.

Figure 14 contains plots of specimen abundance per species in increasing order of abundance by taxon.  These taxa are plotted at the ordinal rank, except for the Grylloblattida, which are plotted at the Suborder rank.  It is of interest to note that for the Lemmatophorina, there is a relatively smooth distribution of abundance numbers, with nearly all the species in the family being well-represented.  For the other taxa having a species with a large number of specimens, most species in the relevant taxon have few specimens, and thus the well-represented species is a unique occurrence.  This can be seen in the Protoperlina, the Psocoptera, the Ephemeroptera, and several other groups.  An interesting future study would be to determine if there are any obvious differences between the abundant and rarer species in these taxa that might account for their specimen numbers.

Nymphal vs. Adult Specimens

The great majority of the specimens from the Elmo locality are remains of adult insects.  However, Carpenter (1935) reported eighty-two specimens of nymphs of what he determined to be at least two species of family Lemmatophoridae, the family that is so well-represented at Elmo by adult specimens.  He based the separation of nymphal specimens on size, with nearly half being 9-10 mm in length and the other half 4.5-5.5 mm long.  The larger specimens were considered by Carpenter to most likely be Lemmatophora typa (Carpenter, 1992), although definitive attribution could not be made.  He considered the smaller nymphs to be most likely the species Lisca minuta or Artinska ovata.  The nymphs possessed what appeared to Carpenter to be lateral gills on the first nine abdominal segments, and he stated in his 1935 paper and again in his 1992 monograph that the nymphs were aquatic.  However, later researchers consider these nymphs as semi-aquatic (Storozhenko, 2002) or even terrestrial (Grimaldi & Engel, 2005). 

Specimen Quality

In his initial announcement of his discovery of the Elmo fossil locality, Elias Sellards (1903) noted: “A very large proportion of the wings are complete and their details of structure clear, even the minute hairs often being present.”   Another measure of quality is the amount of information known about the morphology of the various insect species making up the entomofauna.

Figure 15.  Distribution of Wellington Formation species by type of body part(s) for which species are known.  The first bar of every pair is for holotype/neotype specimens, the second bar of every pair is for all known specimens.

Figure 16.  Relative distribution of Wellington Formation species by type of body part(s) by which species are known.  Left bar is for holotype/neotype specimens, right bar for all known specimens.

Figure 16 compares the numbers of species known by various body parts on a percentage basis.  The left bar is for holotype/neotype specimens and the right bar for the total set of specimens.  These two charts illustrate the advantage of continuing to collect and study specimens of a fauna after the species descriptions, and that much more knowledge can potentially be gained about the morphology of these insects. 

Figure 17.  Photograph of fore wing of Grylloblattida: Lemmatophorina: Euryptilonidae: Stereopterum rotundum Carpenter, 1950.  An example of a species known only by its forewing.  The specimen was collected by the authors in March, 2004 from the Midco beds in Noble County, Oklahoma, and is the first specimen of this species to have been recorded from Midco (the species having been described by Carpenter from material collected at Elmo).

Figure 18.  Grylloblattida: Protoperlina: Chelopteridae: Chelopterum peregrinum Carpenter, 1950, originally described from Elmo type material.  a.  Photograph of forewing of a specimen collected by the authors from the Midco beds in Noble County, Oklahoma.  b. Original drawing by Roy Beckemeyer based on reconstruction of the insect by Frank Carpenter, as depicted in the Treatise on Invertebrate Paleontology: Hexapoda (1992).  The insect is depicted as is it were a pinned specimen, with the wings extended in order to illustrate the venation pattern, even though the insect is a polyneopteran, and would have perched with its wings folded over its back.  c. Original drawing by Roy Beckemeyer of the insect as it might have appeared in life, with wings folded.

Figure 18a is a photograph of another fore wing collected from Noble County.  It is one of two forewing specimens of Grylloblattida: Chelopterum peregrinum known to the authors from the Midco, Oklahoma beds.  This species also was described by Frank Carpenter from Elmo material in 1950.  However, additional Elmo specimens allowed Carpenter to reconstruct the entire insect.  Figure 18b is an original drawing by one of the authors (RJB) based on Carpenter’s reconstruction.  As is typical, Carpenter’s reconstruction of the insect shows it as if it were pinned in a display case, with its wings extended to illustrate the wing venation patterns.  Figure 18c is another drawing by RJB showing the species as it may have looked in life, with the wings folded over the abdomen.  As noted in Figure 16, complete insects are known only for 13% of the Wellington Formation species.

CONCLUSIONS

The Kansas/Oklahoma Wellington Formation has been a major source of information on Lower Permian insect fauna for over 100 years, with nearly 200 species described to date.  The Oklahoma beds are poorly-known in comparison with the Kansas beds but both sites meet the criteria for being Konservat-Lagerstätten: the exceptional quality & quantity of the fossil insect specimens found there

A number of species of Grylloblattida and one species of Psocoptera seem to be especially abundant in the Wellington Formation.  A study of the paleoecology of the Wellington Permian might help to decide whether these insects were indeed more abundant or if they have been preferentially preserved, or both.

Large size seemed to be a character of the Wellington Formation Protodonata, with ten of thirteen insects with fore wing lengths greater than 50 mm belonging to this order.  The orders Palaeodictyoptera, Megasecoptera, and Caloneurodea each possessed one large species, with most of the Wellington Formation species in these orders being much smaller in size.  Since Protodonata were predatory and the other orders were not, this may be an indication that the larger insects in these groups were preferentially subject to predation.  A complementary study would be to conduct a similar analysis of Carboniferous fauna to explore whether there was a greater proportion of large species in these orders at an earlier time.

ACKNOWLEDGEMENTS

Thanks to the late Faith Tillyard Evans, daughter of Robin J. Tillyard, for allowing the use of the photo of her father.  Dr. George Byers of Kansas University allowed the use of his photograph of Frank Carpenter.  Marie Dvorzak of the Geology and Geophysics Library, University of Wisconsin – Madison, provided the picture of Gilbert Raasch and a copy of his PhD Thesis.  Ann Molineux of the Texas Memorial Museum, University of Texas, Austin, provided the image of E.H. Sellards.  Mary Nelson of the Wichita State University Library Department of Special Collections allowed the use of the photos of Paul Tasch and of the town of Elmo, Kansas. 

Roy Beckemeyer is supported in his research by Dr. Michael Morales, Director of the Johnston Geology Museum, Emporia State University, Emporia, Kansas.  Joseph Hall acknowledges the guidance of his thesis advisor, Dr. Salvatore Mazzullo of the Department of Geology, Wichita State University; he also acknowledges the financial support of his thesis work provided by the Kansas Geological Foundation and the Kansas Geological Survey. 

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• Storozhenko, S. Yu.  2002.  Section 2.2.2.2.1.  Order Grylloblattida Walker, 1914 (=Notoptera Crampton, 1915, = Grylloblattodea Brues et Melander, 1932 + Protorthoptera Handlirsch, 1906, = Paraplecoptera Martynov, 1925 + Protoperlaria Tillyard, 1928).  Pp. 278-281 in A. P. Rasnitsyn and D. L. J. Quicke (eds.).  Kluwer Academic Publishers.  Dordrecht, The Netherlands.

• Tasch, P.  1964.  Periodicity in the Wellington Formation of Kansas and Oklahoma.  Pp. 481-496 in D.F. Merriam (Ed.), Symposium on cyclic sedimentation.  Bulletin 169, Vol. II.  State Geological Survey of Kansas, Lawrence, Kansas, USA.

• Tasch, P., and J. Zimmerman.  1962.  The Asthenohymen-Delopterum bed – a new Leonardian insect horizon in the Wellington of Kansas and Oklahoma.  Journal of Paleontology 36(6):1319-1333.

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