ArticlePDF Available

A Possible Hemiphlebiid Damselfly in Late Cretaceous Amber from South Dakota (Odonata: Zygoptera)

Authors:

Abstract and Figures

The first damselfly in Late Cretaceous amber from South Dakota is described and figured.The specimen preserves the forewing apex of a possible hemiphlebiid, a group of relict damselflies today that were apparently widespread and diverse during the Cretaceous.
Content may be subject to copyright.
A possible hemiphlebiid damsely in Late Cretaceous amber from
South Dakota (Odonata: Zygoptera)
André nel1, robert A. dePAlmA2, And michAel S. engel3,4
1. CNRS UMR 7205, Muséum National d’Histoire Naturelle, Entomologie, CP 50, 45 rue
Buffon, F-75005 Paris, France (anel@mnhn.fr)
2. Division of Vertebrate Paleontology, Natural History Museum, and Department of
Geology, 1475 Jayhawk Boulevard, University of Kansas, Lawrence, Kansas 66045-
7613 (paleogen@aol.com)
3. Division of Entomology (Paleoentomology), Natural History Museum, and Department
of Ecology & Evolutionary Biology, 1501 Crestline Drive – Suite 140, University of
Kansas, Lawrence, Kansas 66045-4401 (msengel@ku.edu)
4. Corresponding author
The rst damsely in Late Cretaceous amber from South Dakota is described and gured.
The specimen preserves the forewing apex of a possible hemiphlebiid, a group of relict
damselies today that were apparently widespread and diverse during the Cretaceous.
Keywords: Odonata, Zygoptera, damsely, Maastrichtian, amber, Hemiphlebiidae
trAnSActionS of the KAnSAS
AcAdemy of Science
Vol. 113, no. 3-4
p. 231-234 (2010)
introduction
Damselies, suborder Zygoptera (Odonata),
have a long geological history extending
back to at least the Late Jurassic (e.g.,
Fleck et al., 2001). Most of this material is
preserved as compressions with little or no
relief and frequently only on the basis of
wings. Damselies in amber understandably
preserve greater detail owing to the delity
of amber preservation, but such inclusions
are uncommon and the majority are in resins
of Tertiary age (e.g., Bechly, 1998a; May
and Carle, 2005; Nel and Arillo, 2006).
Interestingly, those damselies recovered in
Cretaceous ambers have all proved to be of the
relic family Hemiphlebiidae (e.g., Lak et al.,
2009; Azar et al., in press).
Herein we provide a brief account of a
fragmentary damsely forewing preserved
in amber recovered from the Hell Creek
Formation in South Dakota. Unfortunately,
the sole specimen deteriorated quickly but
given the degree to which damselies are
uncommonly found in amber, we provide the
following account in the hopes that further
material may some day come to light. In the
description the wing venation nomenclature
follows that of Riek (1976) and Riek and
Kukalová-Peck (1984) as emended by Nel et
al. (1993), Bechly (1996), and Fleck and Nel
(2003).
geologicAl Setting
The amber in which the possible hemiphlebiid
wing was discovered was excavated from the
“Bone Butte” dig site, a uvial sequence in the
Cretaceous strata of Harding County, South
Dakota’s Upper Hell Creek Formation. The
Bone Butte site is estimated to be no more
than 20 m below the Cretaceous-Tertiary
contact (Rothrock, 1937; Cox, 1962; DePalma,
unpubl. data) and detailed observations on
the microstratigraphy, vertebrate, and plant
fossils indicate that the total sedimentary
sequence at the site represents no more than
ve years of deposition (DePalma, unpubl.
data). The majority of amber from Bone Butte
was recovered from a ning-upward sequence
deposited in a paleo oxbow lake.
The sediments at Bone Butte are comprised of
laminated sandstones, siltstones, and clay-
pebble conglomerates and form a section
232 Nel, DePalma and Engel
approximately 2.2 m thick. Two crevasse-splay
deposits (facies 1) consisting of sandstone,
clay-pebble conglomerate, and vertebrate
remains, overlay two additional facies:
laminated sandstone (facies 2) from a paleo
stream channel and a westwardly adjacent
ning-upward sequence (facies 3) from a paleo
oxbow lake. Whereas vertebrate remains from
dinosaurs, mammals, reptiles, amphibians,
sh, and birds are found in all three facies,
amber is found in greatest abundance only
in the lower two strata of facies 3. Facies
3 is primarily subdivided into strata 11 and
15, corresponding to shifts in lithologic
composition. Stratum 15, the lowermost of the
two strata, averages 23 cm in thickness and
consists of nely bedded, buff-tan, medium to
ne-grained sandstone punctuated by laminae
composed of medium to coarse-grained
sandstone and small (<2 cm) rip-up mud clasts.
The sediments of stratum 15 exhibit a ning-
upward trend associated with the formation of
an oxbow lake (Lehman, 1987; Behrensmeyer
and Hook, 1992; Derstler and Myers, 2008;
Henderson and Harrison, 2008) and grade
into nely laminated brownish-black siltstone
rich in plant material at stratum 15’s upper
bounding surface. Stratum 11 directly overlays
stratum 15, averages 15 cm in thickness, and is
composed of nely bedded dark brown-black
siltstone punctuated by laminae composed of
medium-grained sandstone and rip-up mud
clasts typically not exceeding 2 cm in diameter,
as well as lenses of medium to coarse-grained
sandstone enriched by abundant plant matter.
The organic-rich sediments of stratum 11 were
deposited as silt and autochthonous plant debris
lled the paleo oxbow lake, and coarser-grained
laminae and sandstone lenses in strata 11 and
15 represent periodic increases of sedimentation
related to rain storms (Lehman, 1987;
Behrensmeyer and Hook, 1992). Amber is most
common within the upper third of stratum 15
and throughout stratum 11, typically found as
small to moderate-sized (0.5–7 cm) nuggets that
exhibit surface morphology produced as the
resin was originally exuded from its host tree,
such as runnel ows and droplets.
SyStemAtic PAleontology
Family HemipHlebiidae? Tillyard
Genus et Species Indeterminate
(Fig. 1)
Description―Length of preserved fragment
6 mm, width 4.2 mm. Wing hyaline, with only
apical part preserved; preserved veins are
RA, RP1, IR1, apical parts of RP2, IR2, and
RP3/4, all straight except for IR2 zigzagged at
apex; no intercalary longitudinal vein between
preserved main vein; three preserved postnodal
crossveins not aligned with postsubnodal
crossveins; IR1 very short, beginning only
one cell basal of pterostigmal level; RP1 with
a very strong angle below pterostigmal brace;
pterostigmal brace strong but not very oblique;
pterostigma covering one cell, 0.6 mm long,
0.4 mm wide; only three cells between C and
RA distal of pterostigma.
Material―Wing fragment; Late Cretaceous
(Maastrichtian) amber; Hell Creek Formation;
Bone Butte site, Harding County, South
Dakota. During hand preparation of the amber
piece (which measured 9 mm in length) by
R.A.D. the specimen disintegrated.
diScuSSion
The preserved portions of the wing fragment
reported herein are nearly identical to
those of the Early Cretaceous hemiphlebiid
Jordanhemiphlebia electronica Kaddumi from
Jordan (Lak et al., 2009). In those comparable
portions preserved the only discernible
differences are the slightly longer IR1 that
begins below the pterostigma instead of one
cell basal and the presence of only three cells
in area distal of the pterostigma, instead of ve
in J. electronica. Further characters shared by
this fossil and Hemiphlebiidae are as follows:
postnodal and postsubnodal crossveins not
aligned (a symplesiomorphy, sensu Bechly,
1998b; which is probably likely owing to
the ‘basal’ position of Hemephlebiidae in
damsely phylogeny, vide Dumont et al.,
Transactions of the Kansas Academy of Science 113(3-4), 2010 233
2010), all intercalary veins (except IR1 and
IR2) suppressed, and RP1 with a strong
angle below the pterostigmal brace (character
convergently present in Recent Hemiphlebia
and some Coenagrionoidea). Thus this new
fossil is a possible hemiphlebiid although the
lack of information regarding the wing base
and body structures forbids us to conclusively
place it within this family.
It is interesting to note that while Hemiphlebia
survives today only in Tasmania and extreme
southeastern Australia, the family as a whole is
of some considerable antiquity. Hemiphlebiids
are known from as far back as the Late
Jurassic and were clearly widespread during
the Cretaceous (Lak et al., 2009; Azar et
al., in press), even perhaps being dominant
among Zygoptera during the early part of
the Cretaceous. What factors contributed to
the eventual turnover of this fauna, whereby
hemiphlebiids are ecologically replaced by
coenagrionids, is uncertain but may relate to
ecological conditions of the aquatic naiads.
Unfortunately, nothing is known of the
immature stages of these early hemiphlebiids
and until such time as fossil evidence is
recovered we can only speculate. Regardless,
from the record available it is clear that
hemiphlebiids enjoyed a far greater diversity
and distribution during the Mesozoic.
AcKnowledgementS
We are grateful to two anonymous reviewers
for comments on the manuscript. The second
author is grateful to Terry Smith, Dr. Fred
Cichocki, Robert Feeney, Fallon Cohen, the
Lindsey Family, the Palm Beach Museum of
Natural History, and the University of Kansas
for assistance with eldwork and access
to specimens. This is a contribution of the
Division of Entomology, University of Kansas
Natural History Museum.
literAture cited
Azar, D., Prokop, J. and Nel, A. In press.
The rst damsely from Lebanese amber
(Odonata, Zygoptera). Alavesia
Bechly, G. 1996. 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
2:1–402.
Bechly, G. 1998a. New fossil damselies
from Baltic amber with description of a
new species, a redescription of Litheuphae
carpenteri Fraser, and a discussion on the
phylogeny of Epallagidae. International
Journal of Odonatology 1(1):33–63.
Bechly, G. 1998b. New fossil dragonies from
the Lower Cretaceous Crato Formation
of north-east Brazil (Insecta: Odonata).
Suttgarter Beiträge zur Naturkunde, Serie B,
Geologie und Paläontologie 264:1–66.
Behrensmeyer, A.K. and Hook, R.W. 1992.
Paleoenvironmental contexts and taphonomic
modes. pp. 15–93 in Behrensmeyer, A.K.,
Damuth, J.D., DiMichelle, W.A., Potts, R.,
Sues, H.-D. and Wing, S.L. (eds.), Terrestrial
Ecosystems through Time: Evolutionary
Paleoecology of Terrestrial Plants and
Animals. University of Chicago Press;
Chicago, Illinois; xix+568 pp.
Figure 1. Photomicrograph of a probable
hemiphlebiid from the Late Cretaceous of South
Dakota prior to disintegration of the specimen
(Photograph by R.A.D.).
234 Nel, DePalma and Engel
Cox, E.J. (ed). 1962. Geology of selected
highway strips in South Dakota. South
Dakota State Geological Survey, Report of
Investigations 93:1–184.
Derstler, K. and Myers, J. 2008. Taphonomy
of the Tyrannosaurus rex “Peck’s Rex”
from the Hell Creek Formation of Montana.
pp. 67-74 in Larson, P. and Carpenter, K.
(eds.), Tyrannosaurus rex: The Tyrant King.
Indiana University Press, Bloomington,
Indiana, xvi + 435 pp.
Dumont, H.J., Vierstraete, A. and Vaneteren,
J.R. 2010. A molecular phylogeny of the
Odonata (Insecta). Systematic Entomology
35(1):6–18.
Fleck, G. and Nel, A. 2003. Revision of the
Mesozoic family Aeschnidiidae (Odonata:
Anisoptera). Zoologica 153:1–172.
Fleck, G., Nel, A., Bechly, G. and Martínez-
Delclòs, X. 2001. Revision and
phylogenetic afnities of the Jurassic
Steleopteridae Handlirsch, 1906 (Insecta:
Odonata: Zygoptera). Insect Systematics
and Evolution 32(3):285–305.
Henderson, M., Harrison, W. (2002).
Taphonomy and environment of deposition
of a juvenile tyrannosaurid skeleton
from the Hell Creek Formation (latest
Maastrichtian) of eoutheastern Montana.
p. 83–92 in Larson, P. and Carpenter, K.
(eds.), Tyrannosaurus rex: The Tyrant King.
Indiana University Press, Bloomington,
Indiana, xvi + 435 pp.
Lak, M., Fleck, G., Azar, D., Engel, M.S.,
Kaddumi, H.F., Néraudeau, D., Tafforeau,
P. and Nel, A. 2009. Phase contrast x-ray
synchrotron microtomography and the
oldest damselies in amber (Odonata:
Zygoptera: Hemiphlebiidae). Zoological
Journal of the Linnean Society 156(4):913–
923.
Lehman, T.M. 1987. Late Maastrichtian
paleoenvironments and dinosaur
biogeography in the Western Interior
of North America. Palaeogeography,
Palaeoclimatology, Palaeoecology 60:189–
217.
May, M.L. and Carle, F.L. 2005. Pamita
hannahdaltonae gen. nov., sp. nov. from
Baltic amber (Odonata : Amphipterygida).
International Journal of Odonatology
8(2):213–221.
Nel, A. and Arillo, A. 2006. The rst
Baltic amber dysagrionine damsely
(Odonata: Zygoptera: Thaumatoneuridae:
Dysagrioninae). Annales de la Société
Entomologique de France 42(2):179–182.
Nel, A., Martínez-Delclòs, X., Paicheler,
J.-C. and Henrotay, M. 1993. Les
‘Anisozygoptera’ fossiles. Phylogénie et
classication (Odonata). Martinia, Numéro
Hors Série 3:1–311.
Riek, E.F. 1976. A new collection of insects
from the Upper Triassic of South Africa.
Annals of the Natal Museum 22:791–820.
Riek, E.F. and Kukalová-Peck, J. 1984. A new
interpretation of dragony wing venation
based upon Early Carboniferous fossils
from Argentina (Insecta: Odonatoidea) and
basic character states in pterygote wings.
Canadian Journal of Zoology 62(6):1150–
1166.
Rothrock, E.P. 1937. Structural conditions
in Harding County. South Dakota State
Geological Survey, Report of Investigations
28:1–30.
... It currently comprises twelve genera and sixteen fossil species, plus an undetermined genus and species, recovered from the Upper Jurassic to Upper Cretaceous as rock impressions and amber inclusions (Table 1). In particular, the hemiphlebiids are found as rock impressions in the Upper Jurassic of Mongolia and Germany (Pritykina and Vassilenko, 2014;Bechly, 2019), the Lower Cretaceous of England, Brazil, Transbaikalia (Jarzembowski et al., 1998;Vassilenko 2005;Felker and Vasilenko, 2018), and the Upper Cretaceous of Israel (Vassilenko 2014), plus in the amber from the Lower Cretaceous of France and Jordan (Lak et al., 2009) and the Upper Cretaceous of America and Myanmar (Nel et al., 2010). ...
... nov. with a weak angle (Jarzembowski et al., 1998;Bechly, 1998;Vasilenko, 2005Vasilenko, , 2014Lak et al., 2009;Nel et al., 2010;Pritykina and Vassilenko, 2014;Zheng et al., 2017;Felker & Vasilenko, 2018;Bechly, 2019;Zheng, 2021). The alleged hemiphlebiid Pantelusa krassilovi Vasilenko, 2014 ...
Article
Hemiphlebiidae are the most basal lestomorphan family following the latest phylogenetic analysis of the Zygoptera: this unique damselfly family today contains one relict species found in the wetlands of Australia. It was, however, very diverse and widespread during the Mesozoic. Nevertheless, very few species were known obscuring the origination and early evolution of the family. Here we propose a new stem hemiphlebioid taxon (Protohemiphlebiidae Zheng, Jarzembowski & Nel, fam. nov.) based on a new genus and two species: Protohemiphlebia zhangi Zheng, Jarzembowski & Nel, sp. nov. and Protohemiphlebia meiyingae Zheng, Jarzembowski & Nel, sp. nov. The new family shares the characters of both Hemiphlebiidae and Coenagrionoidea, but it is more closely related to Hemiphlebiidae in having the pterostigma with a ‘star-shaped’ microsculpture, and AA originating from the wing base slightly distal of Ax0. Protohemiphlebia Zheng, Jarzembowski & Nel, gen. nov. is further considered to belong to the stem group of Hemiphlebioidea, instead of belonging to the Hemiphlebiidae, in possessing pretibial combs and a weakly kinked RP1 below the Pt-brace. The new damselflies will help to calibrate the origin of Hemiphlebiidae, which could be earlier than their current oldest records in the Kimmeridgean (Late Jurassic).
... Ma; Figure 1B), dipterans account for all but one of the 22 documented inclusions (95% of all inclusions). 4,15 As more samples are recovered from all these deposits, including Big Muddy, it will be noteworthy to see if the assemblage compositions change and whether these initial trends are representative of the overall deposit assemblages or the result of sampling bias. The ordinal fossil diversity and richness of the initial Big Muddy amber assemblage also differs significantly from Grassy Lake amber ($79-78 Ma), a large and well-studied Late Cretaceous deposit located nearby in Alberta ( Figure 1B). ...
... Despite the very important recent discoveries of Odonata of the early Late Cretaceous age, especially in the Burmese amber (Zheng 2020), the Cretaceous/Paleocene transition remains a crucial period for the understanding of the ecological conditions and timing of the diversification of the modern odonatan fauna. The current knowledge on the Turonian-Maastrichtian Odonata is very fragmentary with less than 10 species (Fossilworks Database at http://fossilworks.org/; Nel et al. 2010b), but the recent unexpected discoveries of Maastrichtian Aeschnidiidae confirms the importance of this period for the turnover between the Mesozoic and a more modern fauna (Nel 2021). With 18 described species, the Paleocene Odonata are better known but the majority of taxa have been found in upper Paleocene deposits: Wighton and Wilson (1986) described some Gomphaeschnidae from the Paskapoo Formation (ca. ...
Article
Full-text available
The current knowledge on the Paleocene Odonata is rather limited despite the fact that it is a crucial period for the history of this order. An overview of the fossil odonatans from the Paleocene of Menat (France) is provided. We describe the anisopteran Macrogomphus menatensis sp. nov., first fossil representative of the family Epigomphidae, together with two zygopteran, viz. the dysagrionid Menatagrion hervetae gen. et sp. nov., and the new family Menatlestidae fam. nov., with its type species Menatlestes palaeocenicus gen. et sp. nov. The genus Menatagrion gen. nov. is the first Paleocene record of the Dysagrionidae, otherwise known by a putative Cretaceous genus and several Eocene to Miocene genera. Menatlestes gen. nov., putatively attributed to the stem-group of the Lestinoidea (Megalestidae and Lestidae), would correspond to the oldest record of this clade. With these three new taxa, and the previously described Thanetophilosina menatensis, Valerea multicellulata, “Lestes” zalesskyi, and an Aeshna species indet., the total number of Odonata from Menat goes up to seven species in total; two Anisoptera and five Zygoptera. Furthermore, we propose new evidences showing that the head characters defining the putative suborder Cephalozygoptera are due to deformations, very frequent among the fossil Odonatoptera. We treat the Cephalozygoptera as a junior synonym of Zygoptera.
... The odonatans are rare compared to the other insects in the fossil records especially in the amber deposits. There are scarce records of the odonatans in Cretaceous ambers outside of Kachin amber: two fragmentary wings in Lebanese amber (early Barremian; Azar et al., 2010), a single wing in Jordanian amber (Barremian; Lak et al., 2009), a well-preserved damselfly in Charentes amber (latest Albian; Lak et al., 2009), and a wing apex from amber of South Dakota (Maastrichtian; Nel et al., 2010), all belonging or closely related to the family Hemiphlebiidae. In Kachin amber, odonatans are especially abundant with 38 species of 16 families having been described (Table 1). ...
Article
Odonatans are rare in Cretaceous amber deposits, and only four fragmentary specimens were recorded in Lebanese, Jordanian, Charentes and South Dakota ambers except for Kachin amber. This study reviewed the odonatans in lowermost Cenomanian Kachin amber and commented 11 odonatans, which were previously placed in the family level. Until now, over 350 odonatans have been found in Kachin amber, including 38 species of three extant suborders, and representing the most diverse and abundant dragonfly assemblage in amber inclusions. The dominated odonatans include the damselflies Burmahemiphlebia zhangi and Mesomegaloprepus magnificus, and the damsel-dragonfly Burmaphlebia reifi, accounting for ca. 79% of all odonatans. The Kachin odonatans include the earliest records of the damselfly Perilestidae, Platycnemididae and Platystictidae, and the youngest records of the true dragonfly Araripegomphidae and Stenophlebiidae, contributing to exploring the origination and evolution of these families. Hemiphlebiidae is the dominant family in Kachin amber, represented by the species Burmahemiphlebia zhangi with ca. 160 species been recorded. It is surprising that the hemiphlebiid damselfly is abundant, but the diversity is very low with only two species been previously reported. Herein, a new hemiphlebiid damselfly, Kachinhemiphlebia lini gen. et sp. nov., is described by the unique characters: Arc slightly distal of Ax2 in both fore- and hindwings; and a well-defined AA ending on posterior wing margin below Ax2. This is the second genus of the Hemiphlebiidae in Kachin amber.
... It is distinguished from all other Mesozoic Hemiphlebiidae by the following diagnostic characters: From Burmahemiphlebia Zheng et al., 2016b by veins MP and CuA not shortened; from Cretarchistigma Jarzembowski et al., 1998 by the lower number of postnodals, shorter IR1, and the arculus not aligned with ax2 (also see Bechly 1998aBechly , 2007a; from Cretacoenagrion Jarzembowski, 1990 by the shorter pterostigma, shorter IR1, lower number of 4 non-aligned postnodals (instead of 9 aligned ones), very different shape of the discoidal cell (less acute); from Cretahemiphlebia Jarzembowski et al., 1998 by the lower number of postnodals, shorter IR1, and distally divergent RP2 and IR2; from Electrohemiphlebia Lak et al., 2009 by the very different shape of the forewing (also see Bechly, 1998a, 2007a by the absence of a mesothoracic interpleural suture, only 2 cells between RP1 and RP2 basal of IR1, bigger distance between arculus and ax2, distinct kink in arculus before the origin of the discoidal vein MAb, and very short vein MAb; and from Thairia Felker & Vasilenko, 2018 by a shorter IR1 and straight MP at discoidal cell. Finally, the new genus differs from an unnamed possible hemiphlebiid from the Late Cretaceous Hell Creek Formation of South Dakota (Nel et al., 2010) by the shorter IR1, and the distally divergent RP2 and IR2. ...
Article
Three new taxa of odonates are described from the Upper Jurassic Solnhofen limestone from Eichstätt and Painten in Bavaria (Germany), including the first two genuine Zygoptera (Andrephlebia buergeri gen. et sp. nov. in fam. inc. sed. and Jurahemiphlebia haeckeli gen. et sp. nov. in Hemiphlebiidae) and a new taxon of Stenophlebioptera (Reschiostenophlebia koschnyi gen. et sp. nov. in Stenophlebiidae). With an age of about 152 million years, the holotype of Jurahemiphlebia from the Painten locality represents the oldest fossil record and thus a new calibration point for crown group Zygoptera, Lestoidea, and Hemiphlebiidae, and the oldest record for any living odonate family. Furthermore, the first relatively complete specimen of the dragonfly Prohemeroscopus kuehnapfeli (Prohemeroscopidae) is described, which was previously known only from a pair of isolated hind wings. A revised diagnosis is provided for the species and genus.
... Fossil odonatans are quite rare in amber compared to sedimentary deposits with sporadic records in Baltic, Dominican, Burmese, French, Jordanian, South Dakotan and Lebanese amber (Bechly, 1996a(Bechly, , 1998Fleck et al., 2000;Bechly and Wichard, 2008;Lak et al., 2009;Nel et al., 2010;Azar et al., 2010;Poinar et al., 2010;Bechly et al., 2013). In Asia, many odonatans have been discovered in Mesozoic-Cenozoic sedimentary rocks, but only a few have been recently described from Burmese amber despite a century of study of insect inclusions (Poinar et al., 2010;Bechly et al., 2013;Huang et al., 2015;Schä del and Bechly, 2016;Zheng et al., 2016a,b,c,d). ...
Article
A new dragonfly, Cretagomphaeschnaoides jarzembowskae gen. et sp. nov., is described from mid-Cretaceous Burmese amber. Cretagomphaeschnaoides gen. nov. is of a small size, has a three-celled discoidal triangle, and more undulating vein MAb than other genera in the extinct tribe Gomphaeschnaoidini of the extant family Gomphaeschnidae. This fossil is the second record of Anisoptera in Cretaceous amber.
... Finally, Parahemiphlebia and Enteropia both have a long MP and CuA, and a nonrectangular discoidal cell, and can be excluded from further comparison. Cretarchistigma Jarzembowski et al., 1998, Cretahemiphlebia Jarzembowski et al., 1998 and an unnamed damselfly described by Nel et al. (2010) are all considered to be closely related to Hemiphlebiidae (Jarzembowski et al. 1998, Bechly 1998) but differ from Burmahemiphlebia in having IR1 originating before, instead of distal of, Pt. The extant genus Hemiphlebia differs from Burmahemiphlebia in the presence of a long IR1 originating slightly basal of Pt, the long MP and CuA, a short cell between the bases of RP3/4 and IR2 below Sn, and in developing more crossveins between the main veins (Münz 1919). ...
Article
Two damselflies, Burmahemiphlebia zhangi gen. et sp. nov. and Palaeodysagrion cretacicus gen. et sp. nov., are described from the mid-Cretaceous Burmese amber. Burmahemiphlebia zhangi is the first record of Hemiphlebiidae from this amber, although the family was cosmopolitan during the Mesozoic. It can be readily distinguished from all other members of Hemiphlebiidae in having very short MP and CuA veins, and in its rectangular discoidal cell. The new fossils support the view that hemiphlebiid damselflies were one of the dominant groups of Zygoptera during the Mesozoic. Palaeodysagrion cretacicus is the first dysagrionid damselfly from Burmese amber and the second Mesozoic representative of this predominantly Paleogene group. It differs from other members of Dysagrionidae in having a unique elongate discoidal cell. These new finds increase the diversity of damselflies in mid-Cretaceous Burmese amber.
Research
Full-text available
A census of Hell Creek and Lance Formation dinosaur remains was conducted from April, 2017 through February of 2018. Online databases were reviewed and curators and collections managers interviewed in an effort to determine how much material had been collected over the past 130+ years of exploration. The results of this new census has led to numerous observations regarding the quantity, quality, and locations of the total collection, as well as ancillary data on the faunal diversity and density of Late Cretaceous dinosaur populations. By reviewing the available data, it was also possible to make general observations regarding the current state of certain exploration programs, the nature of collection bias present in those collections and the availability of today's online databases. A total of 653 distinct, associated and/or articulated remains (skulls and partial skeletons) were located. Ceratopsid skulls and partial skeletons (mostly identified as Triceratops) were the most numerous, tallying over 335+ specimens. Hadrosaurids (Edmontosaurus) were second with at least 149 associated and/or articulated remains. Tyrannosaurids (Tyrannosaurus and Nanotyrannus) were third with a total of 71 associated and/or articulated specimens currently known to exist. Basal ornithopods (Thescelosaurus) were also well represented by at least 42 known associated and/or articulated remains. The remaining associated and/or articulated specimens, included pachycephalosaurids (18), ankylosaurids (6) nodosaurids (6), ornithomimids (13), oviraptorosaurids (9), dromaeosaurids (1) and troodontids (1). Over 41,800 isolated bones and teeth, were also located. This number represents only a small fraction of the actual total collection as many of the museums and institutions surveyed were unable to provide complete numbers on isolated elements. Over 46% of these isolated bones and teeth were identified as hadrosauridae, usually identified as Edmontosaurus. Isolated elements identified as ceratopsids made up just over 21% of the total. These were generally identified as Triceratops. Isolated bones and teeth of tyrannosaurids were significantly less, at only 4.6%. The large difference between the associated and/or articulated remains and the isolated bones and teeth of tyrannosaurids (10.9% down to 4.6%) and ceratopsids (51.3% to 21.5%) is likely due to both a preservational and collection bias towards the larger, more likely to be fossilized, and more likely to be collected, Tyrannosaurus and Triceratops skeletons and skulls. Even though small theropods accounted for less than 0.6% of the total recovered associated and/or articulated remains, their teeth and isolated elements were encountered quite frequently. Isolated bones and teeth of dromaeosaurids, troodontids and "unidentified small theropods" accounted for as much as 16% of the total number of isolated remains. This data suggests that there is a tremendous level of collection and preservational bias in the current Hell Creek and Lance sample set Actual small theropod diversity and populations in the Late Cretaceous were most likely much higher than previously considered. It is highly likely that the fluvial and geochemical environment that dominated the Late Cretaceous of this region was simply too rough and tumble for bones of most genera under the 400 kg live weight
Article
Dragonflies (odonatans) are comparatively rare as amber inclusions, and most are not well preserved on account of their size. Here, we report a single piece of Mexican amber with one complete dragonfly and two damselflies. The dragonfly is attributed to the extant gomphid Erpetogomphus Selys Longchamps, and the damselflies belong to the extant coenagrionid Argia Rambur. Both genera are nowadays distributed widely in Mexico. The new discovery dates the origins of these two genera to the Miocene at least.
Article
Full-text available
The fossil dragonfly Burmalindenia imperfecta gen. et sp. nov. is described from mid-Cretaceous Burmese amber as the first record of the odonate suborder Anisoptera for this locality and one of the few records from amber in general. The inclusion comprises two fragments of the two hind wings of a dragonfly. The fossil can be attributed to a new genus and species of the family Gomphidae, presumably in the subfamily Lindeniinae, and features a strange teratological phenomenon in its wing venation.
Article
Full-text available
A new genus and species Electrophenacolestes serafini is described. It is the first Thaumatoneuridae recorded from an amber deposit and the second record of the family in the European Paleogene. A comparison with related genera and families is done. Résumé. Le premier Zygoptera Dysagrioninae de l'ambre balte (Odonata : Zygoptera : Thaumatoneuridae : Dysagrioninae). Le premier Thaumatoneuridae connu de l'ambre balte est décrit sous le nom d'Electrophenacolestes serafini. Il s'agit du second représentant connu de cette famille dans le Paléogène d'Europe. Le fossile est comparé aux familles et genres proches.
Article
Full-text available
The oldest known odonatoid wings are described from the Namurian of Argentina: Eugeropteron lunatum Riek n.g. et sp. and Geropteron arcuatum Riek n.g. and sp. (Meganisoptera: Meganeurina: Eugeropteridae n. fam.). The wings are generalized and support a reinterpretation of the venation of living Odonata as being fully homologous to that of other pterygotes and closely related to Ephemeropteroidea, but different from Neoptera. Therefore, Paleoptera is a valid phylogenetic unit, and Odonatoidea and Ephemeropteroidea are sister groups.
Article
Full-text available
The Jurassic odonate family Steleopteridae is revised. Two new genera and species Parasteleopteron guischardi and Euparasteleopteron viohli are described. The phylogenetic affinities of this group are discussed. The Steleopteridae are excluded from the Epiproctophora and transferred into the Zygoptera (stemgroup). Euphaeopsis multinervis is redescribed and transferred to Epiproctophora: Isophlebioidea, and the genus Pseudoeuphaea with its four species is considered as a nomen dubium in Odonata incertae sedis.
Article
We estimated the phylogeny of the order Odonata, based on sequences of the nuclear ribosomal genes 5.8 S, 18S, and ITS1 and 2. An 18S-only analysis resolved deep relationships well: the order Odonata, as well as suborders Zygoptera and Epiprocta (Anisoptera + Epiophlebia), emerged as monophyletic. Some other deep clades resolved well, but support for more recently diverged clades was generally weak. A second, simultaneous, analysis of the 5.8S and 18S genes with the intergenic spacers ITS1 and 2 resolved some recent branches better, but appeared less reliable for deep clades with, for example, suborder Anisoptera emerging as paraphyletic and Epiophlebia superstes recovered as an Anisopteran, embedded within aeshnoid-like anisopterans and sister to the cordulegastrids. Most existing family levels in the Anisoptera were confirmed as monophyletic clades in both analyses. However, within the corduliids that form a major monophyletic clade with the Libellulidae, several subclades were recovered, of which at least Macromiidae and Oxygastridae are accepted at the family level. In the Zygoptera, the situation is complex. The lestid-like family groups (here called Lestomorpha) emerged as sister taxon to all other zygopterans, with Hemiphlebia sister to all other lestomorphs. Platystictidae formed a second monophylum, subordinated to lestomorphs. At the next level, some traditional clades were confirmed, but the tropical families Megapodagrionidae and Amphipterygidae were recovered as strongly polyphyletic, and tended to nest within the clade Caloptera, rendering it polyphyletic. Platycnemididae were also non-monophyletic, with several representatives of uncertain placement. Coenagrionids were diphyletic. True Platycnemididae and non-American Protoneurids are closely related, but their relationship to the other zygopterans remains obscure and needs more study. New World protoneurids appeared relatively unrelated to old world + Australian protoneurids. Several recent taxonomic changes at the genus level, based on morphology, were confirmed, but other morphology-based taxonomies have misclassified taxa considered currently as Megapodagrionidae, Platycnemididae and Amphipterygidae and have underestimated the number of family-level clades.
Article
Electrohemiphlebia barucheli gen. et sp. nov. and Jordanhemiphlebia electronica gen. et sp. nov., two new genera and species are described, based on exceptional inclusions of hemiphlebiid damselflies in Cretaceous amber from France and Jordan. The type specimen of E. barucheli was studied using phase contrast X-ray synchrotron microtomography, giving exceptional images and detailed information. Its comparison with the recent Hemiphlebia mirabilis confirms the attribution of several Cretaceous damselflies to the Hemiphlebiidae, showing that this particular group was widespread in the Early Cretaceous and probably originated in the Late Jurassic or earlier. The ecological niches today occupied by the small coenagrionoid damselflies were occupied during the Triassic and Jurassic by Protozygoptera, hemiphlebiids during the Early Cretaceous, and modern taxa in the Cenozoic.
Article
During Late Maastrichtian time, three major sedimentary depositional provinces existed in continental environments of the western interior region of North America. The sediments accumulated within these provinces are herein termed the piedmont, alluvial plain, and coastal lowland lithosomes. These sedimentary lithosomes correspond to broad paleoenvironmental and paleophysiographic provinces that resulted from the westward shift of the Pacific coast, the bisection and retreat of the interior epeiric sea, and the development of an intermontane basin system during Maastrichtian time. These dramatic physiographic changes created environments markedly different from those that had prevailed during most of the Late Cretaceous, when western North America consisted of a narrow peninsula flanked on the east by a remarkably stable and monotonous coastal plain. In conjunction with these physiographic and environmental changes, three diverse dinosaur-dominated continental vertebrate faunas arose, herein termed the Leptoceratops, Triceratops, and Alamosaurus faunas. Each was peculiar to a sedimentary/environmental province and among them there was little interchange. The separation of these faunas indicates that the dinosaur-dominated communities had adapted to changing or “deteriorating” environmental conditions at the close of Cretaceous time. The Alamosaurus fauna characterized markedly seasonal, semi-arid, environments of the intermontane basins south of about 35°N latitude. The Triceratops fauna inhabited humid coastal floodplains and swamps bordering the retreating northern embayment of the epeiric sea, north of latitude 35°N. The Leptoceratops fauna inhabited cool piedmont environments flanking the mountainous Cordilleran Overthrust Belt north of 35°N latitude. Recognition of this biogeographic zonation indicates that the Triceratops biozone is only of local utility, and that the biostratigraphy of Upper Cretaceous continental deposits in North America must reflect the provincialism now recognized. Moreover, the evident adaptability of dinosaur faunas to varied and changing environmental conditions suggests that climatic change alone may not have been responsible for their demise.
Structural conditions in Harding County
  • E P Rothrock
Rothrock, E.P. 1937. Structural conditions in Harding County. South Dakota State Geological Survey, Report of Investigations 28:1–30.
  • A Nel
  • X Martínez-Delclòs
  • J.-C Paicheler
  • M Henrotay
Nel, A., Martínez-Delclòs, X., Paicheler, J.-C. and Henrotay, M. 1993. Les 'Anisozygoptera' fossiles. Phylogénie et classification (Odonata). Martinia, Numéro Hors Série 3:1–311.