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A possible hemiphlebiid damsely 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 damsely 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
damselies today that were apparently widespread and diverse during the Cretaceous.
Keywords: Odonata, Zygoptera, damsely, Maastrichtian, amber, Hemiphlebiidae
trAnSActionS of the KAnSAS
AcAdemy of Science
Vol. 113, no. 3-4
p. 231-234 (2010)
introduction
Damselies, 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. Damselies 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 damselies 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 damsely 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 damselies 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
damsely 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.
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