ArticlePDF Available

Abstract and Figures

Amber is of great paleontological importance because it preserves a diverse array of organisms and associated remains from different habitats in and close to the amber-producing forests. Therefore, the discovery of amber inclusions is important not only for tracing the evolutionary history of lineages with otherwise poor fossil records, but also for elucidating the composition, diversity, and ecology of terrestrial paleoecosystems. Here, we report a unique find of African amber with inclusions, from the Cretaceous of Ethiopia. Ancient arthropods belonging to the ants, wasps, thrips, zorapterans, and spiders are the earliest African records of these ecologically important groups and constitute significant discoveries providing insight into the temporal and geographical origins of these lineages. Together with diverse microscopic inclusions, these findings reveal the interactions of plants, fungi and arthropods during an epoch of major change in terrestrial ecosystems, which was caused by the initial radiation of the angiosperms. Because of its age, paleogeographic location and the exceptional preservation of the inclusions, this fossil resin broadens our understanding of the ecology of Cretaceous woodlands.
Content may be subject to copyright.
Cretaceous African life captured in amber
Alexander R. Schmidt
, Vincent Perrichot
, Matthias Svojtka
, Ken B. Anderson
, Kebede H. Belete
Robert Bussert
, Heinrich Dörfelt
, Saskia Jancke
, Barbara Mohr
, Eva Mohrmann
, Paul C. Nascimbene
, André Nel
Patricia Nel
, Eugenio Ragazzi
, Guido Roghi
, Erin E. Saupe
, Kerstin Schmidt
, Harald Schneider
, Paul A. Selden
and Norbert Vávra
Courant Research Centre Geobiology, Georg-August-Universität Göttingen, 37077 Göttingen, Germany;
Centre National de la Recherche Scientique, Unité
Mixte de Recherche 6118 Géosciences & Observatoire des Sciences de lUnivers de Rennes, Université Rennes 1, 35042 Rennes, France;
Paleontological Institute,
University of Kansas, Lawrence, KS 66045;
Department of Paleontology, Geozentrum, Universität Wien, 1090 Vienna, Austria;
Department of Geology,
Southern Illinois University, Carbondale, IL 62901-4502;
Golden Prospect Mining Company Ethiopia, Addis Ababa, Ethiopia;
Institut für Angewandte
Geowissenschaften, Technische Universität Berlin, 10623 Berlin, Germany;
Mikrobielle Phytopathologie and
Institut für Ökologie, Friedrich-Schiller-Universität
Jena, 07743 Jena, Germany;
Museum für Naturkunde zu Berlin, 10115 Berlin, Germany;
Division of Invertebrate Zoology, American Museum of Natural History,
New York, NY 10024;
Centre National de la Recherche Scientique, Unité Mixte de Recherche 7205, Entomologie, Muséum National dHistoire Naturelle, 75005
Paris, France;
Department of Pharmacology and Anaesthesiology, University of Padua, 35131 Padua, Italy;
Institute of Geosciences and Earth Resources,
National Research Council and Department of Geoscience, University of Padua, 35137 Padua, Italy; and
Department of Botany and
Palaeontology Department,
Natural History Museum, London SW7 5BD, United Kingdom
Edited* by David L. Dilcher, University of Florida, Gainesville, FL, and approved March 3, 2010 (received for review January 23, 2010)
Amber is of great paleontological importance because it preserves
a diverse array of organisms and associated remains from dif ferent
habitats in and close to the amber-producing forests. Therefore,
the discovery of amber inclusions is important not only for tracing
the evolutionary history of lineages with otherwise poor fossil
records, but also for elucidating the composition, diversity, and
ecology of terrestrial paleoecosystems. Here, we report a unique
nd of African amber with inclusion s, from the Cretaceous of Ethio-
pia. Ancient arthropods belonging to the ants, wasps, thrips, zor-
apterans, and spiders are the earliest African records of these
ecologically important groups and constitute signicant discov-
eries providing insight into the temporal and geographical origins
of these lineages. Together with diverse microscopic inclusions,
these ndings reveal the interactions of plants, fungi and arthro-
pods during an epoch of major change in terrestrial ecosystems,
which was caused by the initial radiation of the angiosperms.
Because of its age, paleogeographic location and the exceptional
preservation of the inclusions, this fossil resin broadens our under-
standing of the ecology of Cretaceous woodlands.
Ambers contain very important terrestrial taphocenoses.
Arthropods and other invertebrates, small reptiles, feathers,
mammal hairs, plants, and various microbes from different
habitats in and close to the amber forests are commonly pre-
served (16). Ambers and other fossil tree resins are found in
hundreds of Old and New World localities (7), with particular
abundance in the Cretaceous and the Eocene to Miocene. Until
now, most amber deposits have been discovered on the former
northern supercontinent Laurasia (SI Appendix,Fig. S1). With
the exception of the Cretaceous Lebanese and Jordanian ambers
(8, 9) and the Eocene Indian and Miocene Amazonian ambers
(10, 11), no fossiliferous amber deposit was known from the
southern continents that formerly formed Gondwana.
Here, we report a unique fossiliferous African amber. The
amber pieces were discovered within the Debre Libanos Sand-
stone Unit from the northwestern Plateau of Ethiopia (Fig. 1).
Because this geologic unit lacks index fossils (12, 13), we used
independent age-relevant information from the amber itself, its
inclusions, and the sporomorphs of the amber-bearing layer to
date the amber. Combined analysis of these physicochemical and
biological parameters revealed a Late Cenomanian age (9395
million years old) for the amber (SI Appendix).
This fossil resin provides a unique window into a mid-Cretaceous
African woodland ecosystem and helps to elucidate the trophic
relationships within this environment. Diverse inclusions of
arthropods, microorganisms, and plant remains reveal manifold
interactions of plants, fungi, and animals during the early phase of
angiosperm evolution.
Results and Discussion
Amber Characteristics. The amber pieces are colorful and trans-
lucent (Fig. 2 Aand B). Chemical analysis of the Ethiopian amber
indicates that it is a Class Ic amber. This result suggests Cheir-
olepidiaceae did not produce the resin, despite the presence of
pollen grains of this Mesozoic conifer family in the amber-bearing
sediment. Infrared spectroscopy shows that this amber is unique
compared with all other fossil resins studied to date. Although we
do not have conclusive information regarding its botanical source,
the resin could have conceivably been produced by a previously
unknown Cretaceous gymnosperm or possibly even by a mid- to
late-Cretaceous angiosperm (SI Appendix).
Arthropods. Arthropod inclusions are abundant in this amber, with
30 fossil specimens preserved in nine of the studied pieces. These
fossils cover an impressive diversity, including the arachnid orders
Acari and Araneae, and at least 13 families of Hexapoda in the
orders Collembola, Psocoptera, Hemiptera, Thysanoptera, Zor-
aptera, Lepidoptera, Coleoptera, Diptera, and Hymenoptera (SI
Appendix,Table S1).
There is a large gap in the Mesozoic terrestrial arthropod
fossil record from the African continent. Compression fossils are
found in only one insect-bearing locality in the Triassic, one in
the Jurassic, and ve in the Cretaceous (14). Thus, the Ethiopian
amber assemblage is of major signicance for understanding the
biogeography and evolutionary history of African biota. Pre-
liminary identication of the inclusions has revealed noteworthy
discoveries: Most specimens represent a unique fossil record of
their group from Africa, and some are among the oldest records
in the world (Figs. 3 and 4, and SI Appendix,Fig. S10).
The most outstanding discovery is a complete, well-preserved
although enrolled, wingless female ant (Formicidae; Fig. 3A).
Visible characters preclude afnities with the extinct Spheco-
myrminae, which is the only subfamily recorded for contempora-
Author contributions: A.R.S., V.P., and N.V. designed research; A.R.S., V.P., M.S., K.B.A.,
K.H.B., H.D., S.J., B.M., E.M., P.C.N., A.N., P.N., E.R., G.R., E.E.S., H.S., P.A.S., and N.V. per-
formed research; A.R.S., V.P., M.S., K.B.A., R.B., H.D., B.M., P.C.N., A.N., P.N., E.R., G.R.,
E.E.S., H.S., P.A.S., and N.V. analyzed data; and A.R.S., V.P., K.B.A., B.M., P.C.N., E.R.,
E.E.S., K.S., P.A.S., and N.V. wrote the paper.
The authors declare no conict of interest.
*This Direct Submission article had a prearranged editor.
To whom correspondence may be addressed. E-mail:
de or
This article conta ins supporting information online at cgi/content/full/
1000948107/DCSupplemental. PNAS Early Edition
neous and older ants in mid-Cretaceous Burmese and French
amber (15, 16). Regardless of the subfamily, this discovery is sig-
nicant because it is one of the oldest records of an ant and the
earliest from Gondwana. It has been suggested that ants arose in
Laurasia during the Early Cretaceous (1618), but the present
discovery challenges this hypothesis. Ants evolved concurrent with
the rise of angiosperms but apparently remained scarce until
radiating into the worlds most diverse and ecologically dominant
eusocial organisms during the Paleogene (19). The discovery will
aid in resolving the phylogeny and timescale of ant lineages.
Other signicant inclusions comprise a male spider that likely
belongs in the Linyphiidae (Fig. 3B), a family of small spiders
that weave sheet-webs (20). This specimen is the second-oldest
linyphiid discovered to date, the oldest being from Hauterivian
lower Aptian Lebanese amber (21), and only the third fossil
spider species to be described from the African continent (22).
Extant linyphiids are globally distributed but most diverse and
common in mesic and hydric regions of the Northern Hemi-
sphere (23). A thrips (Fig. 3C) belonging to the Merothripidae is
only the second Cretaceous merothripid found to date, the other
one being a dubious specimen from Lebanese amber (24), and
only the third known fossil from this group. The merothripids are
a small but worldwide family, with most species residing in the
Neotropics and in North America; these tiny insects feed on
fungal mycelia on the forest oor or under the bark of decaying
trees. The only identiable springtail (Fig. 3D) belongs to the
Isotomidae, which is known from one older record in Burmese
amber and a few younger Cretaceous and Cenozoic ambers (25).
Also identied is a zorapteran (Fig. 3E), an order of minute,
gregarious insects that are exceedingly rare in the fossil record,
with the oldest examples from Lower Cretaceous Jordanian and
Burmese ambers (9, 26). This order exhibits remarkable mor-
phological stability, because most Cretaceous representatives
belong to the extant genus Zorotypus. The fossil and extant dis-
tribution of this genus in virtually all tropical areas indicates a
great antiquity for the group. We found various wasps of the
cosmopolitan families Eulophidae, Mymaridae, Mymar-
ommatidae, and Trichogrammatidae (Fig. 3 FIand SI Appendix,
Fig. S10 Jand K), which are among the earliest fossils of these
small wasps that parasitize eggs and larvae of microlepidoptera,
beetles, various other insects, arachnids, and nematodes. Lep-
idoptera are evidenced by the presence of microscopic scales (SI
Appendix,Fig. S10F). Cretaceous ambers mostly fossilized
microlepidopterans such as Micropterigidae, but the preservation
of scales alone do not allow for a familial determination. Finally,
one leg and a partial head of a beetle were found fossilized in the
amber (SI Appendix,Fig. S10 Hand I), but it was not possible to
assign those fragments to any beetle lineage.
Microorganisms. Microinclusions of bacteria and fungi occur in
almost every studied amber piece, and a few nematodes were
also found (Fig. 2Cand SI Appendix,Fig. S11). Rod-shaped
bacteria are seen as chains of cells attached to detritus and
decayed arthropods (SI Appendix,Fig. S11 AC).
Thousands of septate, mostly four-celled and slightly curved
fungal conidia, which are related to the extant anamorphic asco-
mycete genus Curvularia, are enclosed in the amber (Fig. 2Cand SI
Appendix,Fig. S11F). Extant species of this genus are common
parasites of vascular plants. The fossil spores are attached to sur-
faces of successive resin ows in the amber pieces, indicating that
they dropped down onto the liquid resin. The plethora of conidia
and the occurrence of the related conidiophores in the amber (SI
Appendix,Fig. S11E) suggest that this Cretaceous Curvularia-like
species sporulated plentifully at the site of the resin-bearing trees.
The Ethiopian amber also provides evidence of fungivory in the
Mesozoic. Many Curvularia-like conidia and possible remnants of
the perithecia of its teleomorph were found inside insect fecal pellets
of up to 800 ×250 μm(Fig.2Cand SI Appendix,Fig. S11 GI); these
pellets are too large to have been produced by springtails and mites
trapped close to the fungi in the amber pieces. Instead, beetles or
their larvae are the most probable grazers of this Cretaceous fungus.
Epiphytic fungi with globular, dark-colored hyphae are also
enclosed (SI Appendix,Fig.S11 JL). They belong to the sooty molds,
Fig. 1. Location of the Ethiopian amber deposit. The white rectangle on the
geologic map indicates the area shown in detail. The red asterisk designates
the locality of the outcrop (10°0845’’ N latitude, 38°5756’’ E longitude).
Area map redrawn from Assefa (12).
| Schmidt et al.
an ecological group of saprophytic fungi that are mostly assignable to
the Capnodiales (Ascomycota), which produce colonies on the sur-
face of living plants. Sooty molds typically obtain their nutrients from
the excretions of aphids, scale insects, and other producers of hon-
eydew, or from plant exudates. The discovery of mid-Cretaceous
epiphytic sooty molds is an example of the morphological stability or
stasis of taxa that, once adapted to particular microhabitats, preserve
their morphological features over some 100 million years.
Plant Remains. Remnants of plants are preserved in the form of
spores from homosporous ferns and lycophytes, pollen grains likely
belonging to the gymnosperm family Podocarpaceae and leaf cuticles
that are probably remnants from some early diverging angiosperm
families such as the Lauraceae, Chloranthaceae, or Proteaceae (SI
Appendix,Fig. S12 AC). Some of the more remarkable minute plant
remnants are abundant stellate hairs (Fig. 2 Dand Eand SI Appendix,
Fig. S12 DF). Comparison of these hairs to modern plants excludes
any relation to angiosperms and instead suggests ferns as the likely
source, e.g., the tree fern family Cyatheaceae and the mostly epiphytic
lmy fern family Hymenophyllaceae. As in the fossils, the stellate
hairs of extant representatives of these fern families are composed of
at rays with margins that curve upward when drying and later drop
from the fronds.
Paleoecology. The onset and evolutionary success of angiosperms
led to a reorganization of terrestrial ecosystems in the middle
Cretaceous (27, 28). Coevolutionary events of plants and animals
provided a basis for radiation and speciation. Archaic gymno-
sperms declined and modern gymnosperm and fern families
diversied in the shadow of the angiosperms. As a result, there
was tremendous macroscopic change in terrestrial ecosystems
that inuenced many groups of insects, fungi, and liverworts, and
probably also amphibians and mammals (2831). In the Cen-
omanian, angiosperms were no longer restricted to early suc-
cession stages but became dominant in woodlands worldwide. In
this time of change, the Ethiopian amber forest represents an
ecosystem in which both conifers, such as members of the extinct
Cheirolepidiaceae and the extant Podocarpaceae families, and
early angiosperms co-occurred. The sporomorphs of the amber-
bearing sediment show that other conifers, different angiosperms
and diverse fungi, mosses, lycophytes, and ferns were also
abundant in this amber forest (SI Appendix,Fig. S9 and Fig. S12).
An epiphytic community comprising ferns and fungi developed.
Ascomycetes played a role not only as decomposers but also as
parasites and served as a food source for insects. Although an-
giosperms were present in the Ethiopian amber forest, no insects
were found that would have pollinated their early owers. Many
hexapods trapped by the tree resin occupy other functional
niches in extant forests, such as small aerial parasitoids (wasps)
and fungivores or detritivores (springtails, merothripid, pso-
copteran, zorapteran) living among the leaf litter or under tree
bark. Predators are represented only by the ant, which was
probably foraging on the soil and on plants, and the linyphiid
spider, whose extant relatives typically construct webs close to
the ground in leaf litter, although some build webs in bushes or
other vegetation.
The Ethiopian amber is of great importance for improving our
knowledge of the evolutionary history of terrestrial arthropods,
Fig. 2. Ethiopian amber and its microinclusions. (A) Large translucent piece of amber of 1150 g (NHMW, N3881). (B) Color of amber pieces (MB. Pb. 2009/205).
(C)Curvularia-like fungal spores in an insect fecal pellet (MB. Pb. 2009/201). (D) Stellate hair resembling those of modern tree ferns of the family Cyatheaceae
(NHMW, N6964). (E) Stellate hair resembling those of the modern epiphytic lmy fern family Hymenophyllaceae (MB.Pb. 2009/206). (Scale bars: A, 1 cm, B, 1 mm;
CE, 100 μm.)
Schmidt et al. PNAS Early Edition
plants, and fungi during the initial radiation of angiosperms in
the Cretaceous. Additional discoveries of unique amber deposits
will aid in closing the large gap in the Mesozoic record of ter-
restrial arthropods from the African continent.
Materials and Methods
Locality. The unique amber deposit was discovered near the town of Alem
Ketema in the eastern part of the northwestern Plateau of Ethiopia (Fig. 1). The
amber pieces occur in a siltstone within the Debre Libanos Sandstone Unit,
exposed along the slopes of the Wenchit River valley (SI Appendix,Fig. S2). A
few hundred pieces of amber have been recovered so far; we had access to 62
of them for investigation. Most of the pieces are 15 cm in size, but single
pieces of up to 25 cm were also found (Fig. 2Aand SI Appendix,Fig. S3 Aand B).
Preparation and Microscopy. The pieces of amber were ground and polished
manually with a series of wet silicon carbide abrasive papers [grit from FEPA
P 6004000 (25.8 μmto5μm particle size), rm Struers] to remove the
weathered opaque surface and to minimize light scattering for the inves-
tigation. The original amber pieces were sometimes separated to isolate
inclusions for investigation. The inclusions were studied by using incident-light
microscopes (Carl Zeiss Stemi 2000 and Olympus SZH-ILLD) and transmitted-
light microscopes (Carl Zeiss AxioScope A1 and Olympus BX50) equipped with
Canon 450D and Olympus Color View IIIu digital cameras. In some instances,
incident and transmitted light was used simultaneously. To better illustrate
the three-dimensional inclusions, some photomicrographs were combined by
using the software package HeliconFocus 4.45. Fig. 3 CIand SI Appendix,Fig.
S9 A,C,Dand HN,Fig. S10 Aand CL,Fig. S11 B,D, and E, and Fig. S12 ACand
E, were obtained from several optical sections.
Fig. 3. Assorted arthropods fossilized in CretaceousEthiopian amber. (A) A worker ant (Hymenoptera, Formicidae; NHMW, N6976). (B) A male sheet-web weaver
spider (Araneae ,L inyphiidae; MB. A 1664). (C) A thrips (Thysanoptera, Merothripidae;NHMW, N6974a). (D)A springtail (Collembola,Isotomidae; NHMW,N6969). (E)
A Zoraptera (NHMW,N6991). (F) A wasp of the Eulophidae(Hymenoptera;NHMW, N6966a).(G) A fairy wasp (Hymenoptera,Mymaridae; NHMW,N6970). (H)Afalse
fairy wasp (Hymenoptera, Mymarommatidae; NHMW, N6965a). (I) A representative of the Trichogrammatidae (Hymenoptera; MB. I 5654). (Scale bars: 500 μm.)
| Schmidt et al.
Repository. The amber pieces are deposited in the Museum für Naturkunde
zu Berlin, (MB. Pb. 2009/200 to MB. Pb. 2009/211, MB Pb. 2009/313 to MB Pb.
2009/341, MB. I 5654 to MB. I 5657, and MB. A 1664), in the Department of
Mineralogy and Petrography, Naturhistorisches Museum Wien, (NHMW,
N3881 and NHMW, N6964 to NHMW, N6991) and in the American Museum
of Natural History, New York (three pieces, sine numero). Please note that
some of the original 62 amber pieces were divided to investigate the
inclusions; each obtained fragment now has a separate collection number.
ACKNOWLEDGMENTS. We thank S. and H. Kaiser (Maria Enzersdorf, Austria)
for providing amber pieces for this study, A. Beran (Vienna), A. Chilin (Padua,
Italy), A. Giaretta (Padua, Italy), V. M. F. Hammer (Vienna), S. Salzmann
(Berlin), E. Stenzel (Berlin), and M. Weber (Vienna) for technical support and B.
Kosmowska-Ceranowicz (Warsaw), C. Pott (Stockholm), and E. Schrank (Berlin)
for discussion. This is contribution number 35 from the Courant Research
Centre Geobiology that is funded by the German Initiative of Excellence, and a
contribution to the project AMBRACE (BLAN07-1-184190) of the French
National Research Agency.
1. Grimaldi DA (1996) Amber: Window to the Past (Abrams, New York).
2. Weitschat W, Wichard W (2002) Atlas of Plants and Animals in Baltic Amber (Pfeil,
3. Schmidt AR, Ragazzi E, Coppellotti O, Roghi G (2006) A microworld in Triassic amber.
Nature 444:835.
4. Schmidt AR, Dörfelt H, Perrichot V (2007) Carnivorous fungi from Cretaceous amber.
Science 318:1743.
5. Girard V, et al. (2008) Evidence for marine microfossils from amber. Proc Natl Acad Sci
USA 105:1742617429.
6. Perrichot V, Girard V (2009) A unique piece of amber and the complexity of ancient
forest ecosystems. Palaios 24:137139.
7. Martínez-Delclòs X, Briggs DEG, Peñalver E (2004) Taphonomy of insects in carbonates
and amber. Palaeogeogr Palaeoclimatol Palaeoecol 203:1964.
8. Poinar GO, Jr, Milki R (2001) Lebanese Amber. The Oldest Insect Ecosystem in
Fossilized Resin (Oregon State Univ Press, Corvallis).
9. Kaddumi HF (2007) Amber of Jordan (Eternal River Museum of Natural History,
Fig. 4. Fossil record of signicant arthropods found in Ethiopian amber.
Schmidt et al. PNAS Early Edition
10. Alimohammadian H, Sahni A, Patnaik R, Rana RS, Singh H (2005) First record of an
exceptionally diverse and well preserved amber-embedded biota from Lower Eocene
(52 Ma) lignites, Vastan, Gujarat. Curr Sci 89:13281330.
11. Antoine P-O, et al. (2006) Amber from western Amazonia reveals Neotropical
diversity during the middle Miocene. Proc Natl Acad Sci USA 103:1359513600.
12. Assefa G (1991) Lithostratigraphy and environment of deposition of the Late Jurassic
Early Cretaceous sequence of the central part of the Northwestern Plateau, Ethiopia.
Neues Jahrb Geol P-A 182:255284.
13. Russo A, Assefa G, Atnafu B (1994) Sedimentary evolution of the Abay River (Blue Nil)
Basin, Ethiopia. Neues Jahrb Geol P-M 5:291308.
14. Schlüter T (2003) Fossil insects in Gondwana localities and palaeodiversity trends.
Acta Zool Cracov 46(Suppl):345371.
15. Engel MS, Grimaldi DA (2005) Primitive new ants in Cretaceous amber from Myanmar,
New Jersey, and Canada (Hymenoptera, Formicidae). Am Mus Novit 3485:123.
16. Perrichot V, Lacau S, Néraudeau D, Nel A (2008) Fossil evidence for the early ant
evolution. Naturwissenschaften 95:8590.
17. Grimaldi DA, Agosti D (2000) A formicine in New Jersey Cretaceous amber
(Hymenoptera: Formicidae) and early evolution of the ants. Proc Natl Acad Sci USA
18. Wilson EO, Hölldobler B (2005) The rise of the ants: A phylogenetic and ecological
explanation. Proc Natl Acad Sci USA 102:74117414.
19. Moreau CS, Bell CD, Vila R, Archibald SB, Pierce NE (2006) Phylogeny of the ants:
Diversication in the age of angiosperms. Science 312:101104.
20. Platnick NI (2009) The World Spider Catalog, version 10.0., Available at http://research. (Am Mus Nat Hist New York). Accessed
September 9, 2009.
21. Penney D, Selden PA (2002) The oldest linyphiid spider, in Lower Cretaceous Lebanese
amber (Araneae, Linyphiidae, Linyphiinae). J Arachnol 30:487493.
22. Selden PA, Anderson HM, Anderson JM (2009) A review of the fossil record of spiders
(Araneae) with special reference to Africa, and description of a new specimen from
the Triassic Molteno Formation of South Africa. Afr Invertebr 50:105116.
23. Draney ML, Buckle DJ (2005) Spiders of North America: An Identication Manual, eds
Ubick DP, Paquin P, Cushing PE, Roth V (Am Arachnol Soc, California), pp 124161.
24. Bhatti JS (1979) A revised classication of Thysanoptera. Abstract Volume, the
Workshop on Advances in Insect Taxonomy in India and the Orient (Association for
the study of Oriental Insects, New Dehli), 4648.
25. Christiansen K, Nascimbene PC (2006) Collembola (Arthropoda, Hexapoda) from the
mid Cretaceous of Myanmar (Burma). Cretac Res 27:318363.
26. Engel MS, Grimaldi DA (2002) The rst Mesozoic Zoraptera. Am Mus Novit 3362:120.
27. Dilcher DL (2000) Towards a new synthesis: Major evolutionary trends in the
angiosperm evolution. Proc Natl Acad Sci USA 97:70307036.
28. Wang H, et al. (2009) Rosid radiation and the rapid rise of angiosperm-dominated
forests. Proc Natl Acad Sci USA 106:38533858.
29. Crane PR (1987) The Origin of Angiosperms and their Biological Consequences, eds
Friis EM, Chaloner WG, Crane, PR (Cambridge Univ Press, Cambridge), pp 107144.
30. Schneider H, et al. (2004) Ferns diversied in the shadow of angiosperms. Nature 428:
31. Heinrichs J, Hentschel J, Wilson R, Feldberg K, Schneider H (2007) Evolution of leafy
liverworts (Jungermanniales, Marchantiophyta): Estimating divergence times from
chloroplast DNA sequences using penalized likelyhood with integrated fossil evidence.
Taxon 56:3144.
| Schmidt et al.
... Ethiopian amber from the northwestern Plateau of Ethiopia has been described from an unknown botanical source containing an abundance of well-preserved arthropod inclusions (Schmidt et al., 2010). Perrichot et al. (2016) considered the age likely Miocene, which was later confirmed by Coty et al. (2016) and Bouju and Perrichot (2020). ...
... similar to East African Defaunation resins, copals and Miocene ambers from Mexico or Dominican Republic (also see McCoy et al., 2017). Only a few platypodine inclusions have been found in this amber (Schmidt et al., 2010). Thus, it is not possible to speculate if platypodines are well represented in the amber as is the case in other Miocene deposits (Peris et al., 2015(Peris et al., , 2017. ...
... A testaceous strip on the anterior face of antennal club is not recognizable in the 3D reconstruction of D. ethiopicus; however, this could be an artefact of the fidelity of the scan because the setae of the club were also not reconstructed correctly in the image. Occurrence: According to Schmidt et al. (2010), the amber comes from a region near the town of Alem Ketemain in the eastern part of the northwestern Plateau of Ethiopia and is Miocene in age (Coty et al., 2016;Bouju and Perrichot, 2020). ...
New fossil species of the genus Diapus (Coleoptera: Curculionidae: Platypodinae) are described in Miocene amber from Ethiopia and Zhangpu, China, based on male and female specimens. We report 24 male and 6 female specimens in Zhangpu amber and one male from Ethiopian amber. However, the number of specimens in Ethiopian amber will probably increase when more material is available for study. The large amount of Diapus specimens supports the paleoenvironmental reconstruction of the Zhangpu amber forest as a megathermal seasonal rainforest. The new species highlight that Platypodinae diversification likely associated with angiosperms may have started in the Cretaceous with an increase of diversity in the Miocene. The specimens described here are the first record of Diapus preserved in amber.
... As is often the case with Ethiopian amber gathered by traders, the precise locality remains unknown as the material is collected from at least four localities and can be mixed up in bags of raw pieces available for sale. However, all localities are situated in the North Shewa Zone (NSZ) and South Wollo Zone (SWZ) of the Amhara Region, on the slopes of the major rivers and their tributaries (Schmidt et al., 2010;Coty et al., 2016;Bouju and Perrichot, 2020;Bouju et al., 2021, fig. 2 for a map with localities): a first outcrop was reported by Schmidt et al. (2010) Published by Copernicus Publications on behalf of the Museum für Naturkunde Berlin. 340 V. Bouju et al.: The genus Allodia (Diptera: Mycetophilidae) in Miocene Ethiopian amber from the northwestern part of the NSZ, in a gorge of the river Wenchit, near the town of Alem Ketema; Coty et al. (2016) obtained some amber from another outcrop located near the hamlet of Jema, along the river Jamma; a third outcrop, further east, was accessed by two of us (Vincent Perrichot and Valentine Bouju) in 2019, in a locality named Woll down the gorge of an affluent of the river Jamma; and a fourth one, the westernmost, was reported to us by a trader near the town of Fiche, again in an affluent of the Jamma river (Binyam Teferi, personal communication, 2019). ...
... As is often the case with Ethiopian amber gathered by traders, the precise locality remains unknown as the material is collected from at least four localities and can be mixed up in bags of raw pieces available for sale. However, all localities are situated in the North Shewa Zone (NSZ) and South Wollo Zone (SWZ) of the Amhara Region, on the slopes of the major rivers and their tributaries (Schmidt et al., 2010;Coty et al., 2016;Bouju and Perrichot, 2020;Bouju et al., 2021, fig. 2 for a map with localities): a first outcrop was reported by Schmidt et al. (2010) Published by Copernicus Publications on behalf of the Museum für Naturkunde Berlin. 340 V. Bouju et al.: The genus Allodia (Diptera: Mycetophilidae) in Miocene Ethiopian amber from the northwestern part of the NSZ, in a gorge of the river Wenchit, near the town of Alem Ketema; Coty et al. (2016) obtained some amber from another outcrop located near the hamlet of Jema, along the river Jamma; a third outcrop, further east, was accessed by two of us (Vincent Perrichot and Valentine Bouju) in 2019, in a locality named Woll down the gorge of an affluent of the river Jamma; and a fourth one, the westernmost, was reported to us by a trader near the town of Fiche, again in an affluent of the Jamma river (Binyam Teferi, personal communication, 2019). ...
... The lower layer, more or less covered by the streams, corresponds to Upper Cretaceous sandstones from the geological formation named Debre Libanos Sandstone or Upper Sandstone (Belay et al., 2009). Ethiopian amber was first suggested to originate from this Cretaceous layer, mostly because of erroneous palynomorph determinations (Schmidt et al., 2010;Perrichot et al., 2016Perrichot et al., , 2018. However, further studies of the palynomorphs from the original and additional material enabled a new estimation of the amber age, now considered to be early Miocene (16-23 Myr old; Perrichot et al., 2018;Bouju and Perrichot, 2020). ...
Full-text available
A new, extinct species of Allodia Winnertz is described from early Miocene amber of Ethiopia. Allodia pale-oafricana sp. nov. is mostly characterized by the scutum with strong anteromarginal, dorsocentral, and lateral setae and the wing with the stem of the M-fork slightly shorter than the vein r-m and the base of the M4-CuA fork aligned with the base of r-m. The assignment to any of the two subgenera Al-lodia stricto sensu or Brachycampta Winnertz remains equivocal as the fossil intermingles traits found in both taxa. Allo-dia is known mostly from the Palearctic region, while only a few species have been described from Africa. In this regard , the new fossil species from Ethiopia brings significant new information regarding the Afrotropical distribution and natural history of the genus.
... Amber pieces from this deposit are unique for their transparency and for the life-like preservation of biological inclusions, providing a unique window to the past fauna and flora of the African continent. Ethiopian amber was originally attributed to the mid-Cretaceous period (Schmidt et al. 2010), but it soon became apparent that inclusions were of a more recent age. Evidence coming from chemical characteristics of the resin, analysis of sporomorphs in the associated sediments and the presence of taxa from extant families and genera, suggest that the amber is Miocene in age (Coty et al. 2016;Perrichot et al. 2016;Bouju & Perrichot 2020). ...
... Although the number of taxa recorded in this deposit is still low, efforts have been made in recent years to describe its diversity. Schmidt et al. (2010) listed 30 arthropod specimens found in Ethiopian amber, comprising two arachnid orders and at least 13 families of Hexapoda, as well as microorganisms and plant fragments. Nevertheless, to date, the insects described from the amber comprise a species of the ant genus Melissotarsus Emery, a biting midge of the genus Forcipomyia Meigen and two thrips (Coty et al. 2016;Ulitzka 2020;Szadziewski et al. 2021). ...
Ethiopian amber is one of the few sources of fossiliferous resins in Africa. Evidence from various sources suggests that this amber is Miocene in age, providing a unique view to the past fauna and flora of the African continent. Stingless bees (tribe Meliponini) are among the most important pollinators in present-day tropical environments, and have a relatively rich fossil record in comparison to other bees. Although they are recorded in various resin deposits around the globe, no stingless-bee fossil is known for the African continent. In the present contribution we describe Dactylurina aethiopica sp. nov., the first fossil species of the genus. The new species is another piece in the “puzzle” concerning the age of Ethiopian amber, since the existence of representatives of extant genera indicates a relatively recent age for the amber. Presence of meliponines in Ethiopian amber also supports the existence of tropical forests in the northwestern Plateau of Ethiopia during the Miocene.
... La dominance d'espèces arborescentes productrices de résine et la présence d'eau douce suggèrent l'existence d'une forêt semidécidue mélangée à une communauté de plantes vivaces, dans une région deltaïque(Nel et al., 2004b) myrmicines sont donc ici étudiés pour la première fois. Le matériel est composé d'une centaine de fragments conservés au Muséum national d'Histoire naturelle, Paris, France.Les gisements à ambre fossilifère d'Éthiopie sont une découverte scientifique récente, publiée pour la première fois il y a une décennie parSchmidt et al. (2010). Ces auteurs suggéraient un âge crétacé et mentionnaient la première découverte de multiples bioinclusions 19 animales et végétales, parmi lesquelles une unique fourmi. ...
... Les bioinclusions reportées parSchmidt et al. (2010) contiennent également une grande variété d'organismes protozoaires mais aussi des Fungi, cependant ce type de bioinclusion n'est pas unique à l'ambre éthiopien et il est même plutôt fréquent d'en observer. ...
Avec plus de 7000 espèces, les fourmis Myrmicinae constituent l’un des plus grands succès écologiques de l’histoire. Pourtant, leur histoire évolutive reste mal comprise. Le présent travail tente de retracer l’évolution du groupe, par l’étude taxonomique des récentes découvertes de myrmicines fossiles (ambre éocène de l’Oise, ambre miocène de Zhangpu et d’Éthiopie), pour les utiliser comme nouveaux points de calibration. L’approche combine les plus récents outils permettant de mieux considérer les données paléontologiques (taux de diversification, d’échantillonnage, etc.), et leur intégration dans l’analyse phylogénétique (modèle FBD, CladeAge). L’effet des diverses modalités de calibration (calibration à la racine, du groupe-couronne ou à l’origine, modèles de distribution, node-dating vs. tip-dating) sur les estimations des temps de divergence est également testé et discuté. Enfin, l’histoire biogéographique est revue à l’aune des nouvelles occurrences et des résultats phylogénétiques. Le groupe serait apparu dans le Nouveau Monde au Crétacé Supérieur (85-95 Ma), sans toutefois montrer une appartenance plus marquée au Néarctique ou au Néotropique. Les grandes lignées se seraient ensuite rapidement dispersées, en particulier à l’Éocène à travers l’Antarctique, la Béringie et le Greenland. L’extension des latitudes tropicales à la suite d’évènements hyperthermiques (ETM, MECO) auraient permis des dispersions successives entre Nouveau et Ancien Monde, et expliquent la disparité des distributions actuelles, où les lignées basales sont respectivement restreintes au Néarctique-Paléarctique et au Nouveau Monde, tandis que les lignées plus dérivées montrent des distributions plus larges mais plus hétérogènes.
... The amber inclusions were examined under a Zeiss Stereo Discovery V8 Ethiopia, 1996). Amhara Region, North Shewa Zone: 1, Alem Ketema (from Schmidt et al., 2010); 2, Jema, near Merany (from Coty et al., 2016); 3, Woll, east of Zemero (from Bouju & Perrichot, 2020). Oromia Region, North Shewa Zone: 4, precise locality unknown near the town of Fiche (unpublished pers. ...
... In contrast to the Ethiopian amber described by Schmidt et al. (2010), which was devoid of cryptogam inclusions, the samples obtained more recently contain diverse bryophytes and a lichen. The 14 fossils reported here were found in a relatively small number of eight amber pieces of a maximum size of 3 cm. ...
Amber is renowned for the exceptional preservation state of its inclusions, allowing detailed morphological analysis and providing relevant environmental, palaeoecological, geographical, and geological information. Amber deposits predominantly known from North America, Europe, and Asia, are considered to be rare on the continents that formed Gondwana. The recent discovery of fossiliferous amber deposits in Ethiopia therefore provides an inimitable opportunity to close gaps in the fossil record of African terrestrial biota, and to study organisms otherwise rare in the fossil record. Here we show that diverse cryptogams are preserved in highest fidelity in Miocene Ethiopian amber. We describe gametophyte fragments of four liverworts: Thysananthus aethiopicus sp. nov. (Porellales, Lejeuneaceae), Lejeunea abyssinicoides sp. nov. (Porellales, Lejeuneaceae), Frullania shewanensis sp. nov. (Porellales, Frullaniaceae), and Frullania palaeoafricana sp. nov. (Porellales, Frullaniaceae). Furthermore, we describe a pleurocarpous moss of the extant genus Isopterygium (Hypnales, Pylaisiadelphaceae) and a lichen representing the order Lecanorales. These new specimens represent the first amber fossils of liverworts, mosses, and lichens from the African continent and render Ethiopian amber one of the few worldwide amber deposits preserving bryophytes (mosses and liverworts) or lichens. Fossil species of Thysananthus were recorded in Eocene Baltic and Oligocene Bitterfeld as well as Miocene Dominican and probably also Miocene Mexican ambers. Fossils which can unequivocally be assigned to Lejeunea have only been found in Dominican amber, so far. Neotropical ambers contain only one taxon of Frullania to date, while the genus is most diverse in Baltic, Bitterfeld, and Rovno ambers, formed in temperate regions. The new fossils support a tropical to subtropical origin of Ethiopian amber. The new African liverwort fossils are included in an updated list of leafy liverworts described from worldwide Cenozoic ambers to date. This article is protected by copyright. All rights reserved.
... The Early Cretaceous yields the first amber deposits with a high diversity of inclusions, from various sites around the world (Figs 2, 3): Lebanon (late Barremian), Spain (early Albian), France (late Albian/ early Cenomanian-Santonian), Myanmar: mainly (early Cenomanian), New Jersey (late Turonian), Russia (Taimyr, middle Santonian) and Canada (late Campanian). One amber deposit from Ethiopia was initially described as being of Cenomanian age (Schmidt et al., 2010), but has since been revised to date to the Miocene (Coty et al., 2016). The number of described Coleoptera species varies among these deposits from only a few specimens to diverse and abundant assemblages, but beetles have been found in all of them (Peris, 2016). ...
... Despite this, these lists do (2) Spain (early Albian); (3) France (late Albian/ early Cenomanian-Santonian); (4) Myanmar (early Cenomanian); (5) New Jersey (late Turonian); (6) Russia (Taimyr, middle Santonian); and (7) Canada (late Campanian). Modified from Schmidt et al. (2010). ...
Beetles, the most successful group of invertebrates on Earth, have a worldwide distribution and an outstanding fossil record. In addition, they are well known as inclusions in fossil resin. In historical studies of fossil material, specimens were often named and described without placing the taxa in an ecological context. However, the research philosophy for fossil beetles has changed over the past few years. In this article, we summarize the palaeoecological interpretations of fossil beetles from Cretaceous ambers, which includes species from 69 families, most of which were described during the last 3 years. By analysing current habits of those families, we argue that saproxylicity was the most common feeding strategy for these fossil beetles. More specifically, fungivorous species appear to dominate. In contrast, we find only anecdotal evidence for the presence of wood-boring groups, and it is thus necessary to identify alternative abiotic or biotic processes that are responsible for the copious resin production at this time. Finally, the recent description of some beetles as gymnosperm pollinators during the Cretaceous lends more weight to the importance of amber studies in addressing the role of beetles in the evolution of pollination strategies.
... As pointed out by diverse authors, the amber appearance is determined by the presence of micro-inclusions, mainly of organic origin (e.g. Breton et al., 1999Breton et al., , 2013Breton et al., , 2018Breton and Tostain, 2005;Schmidt and Schäfer, 2005;Schmidt et al., 2010;Girard, 2010;Speranza et al., 2010Speranza et al., , 2015Beimforde and Schmidt, 2011;Saint Martin et al., 2012Girard et al., 2013aGirard et al., , 2013bSaint Martin and Saint Martin, 2018). The more translucent grains often show a reddish peripheral fringe variable in thickness (20-50 mm) corresponding to partial colonization by microbial communities ( Fig. 8A-F). ...
Full-text available
The Upper Cretaceous sedimentary succession cropping out in the Sainte-Baume Massif (Var, southeastern France) is of late Turonian to Campanian age. There, complex imbrication and rapid evolution in space and time of shallow-marine, fluvio-deltaic and lacustrine environments at the front of the North Provence Durancian uplift occur during the Santonian. Marly and/or sandy strata rich in lignite remains and amber grains are interspersed throughout the Santonian series. According to new in situ collections, the features of the amber grains, and their micro-inclusions, are similar to those found in well-known amber-bearing sites from the Bouches-du-Rhône department. This testifies of a widespread resin-producing tree forests in southeastern France at that time.
... discussion in Savoretti & al., 2018). As part of the first shift in diversification rate in mosses in the Cretaceous (Laenen & al., 2014), the evolution of the main haplolepideous lineages may thus be an example supporting the "shadow of angiosperm" hypothesis, as a response to an explosive increase in the structural diversity of flowering plants (Schmidt & al., 2010;Laenen & al., 2014). However, low clade support may also result from using too few markers or markers with little variation and/or considerable homoplasy. ...
Haplolepideous moss systematics has been through major changes based on morpho-molecular studies. However, many taxa remain little studied with still unclear circumscriptions and relationships among the traditionally recognized families. We inferred the phylogenetic relationships of the haplolepideous mosses based on a sample of 37 out of the 38 haplolepideous moss families and using mitochondrial (nad5 G1 intron) and chloroplast (trnS-rps4 spacer/rps4 gene and trnL gene/trnL-trnF spacer) markers. The resolved relationships indicate the need for rearrangements in the circumscription of the Aongstroemiaceae and Dicranellaceae and their genera, which correspond to the variation of morphological characters of leaf shape, rhizoid tuber morphology, and capsule and peristome traits. The two families, although historically considered morphologically close, were resolved as separate lineages in the core haplolepideous clade. The genera Aongstroemia and Dicranella were resolved as polyphyletic across the clades corresponding to the families Aongstroemiaceae and Dicranellaceae and other clades. The characterization of Aongstroemia by julaceous plants is not supported; the three species sampled here belong in three separate families and differ from one another in other (more stable) characters, such as the presence and shape of peristome teeth. Regarding Dicranella, the sampling of 13 of its 167 species suggests the broad morphological variation comprised by the genus under its current circumscription might be narrowed down to smaller ranges corresponding to the phylogenetically separate clades. The present molecular data furthermore suggest changes in the composition of the protohaplolepideous Dicranidae grade, including one Dicranella clade, a new lineage with a ditrichoid morphology, but excluding the Chrysoblastellaceae, which were resolved in the core haplolepideous clade. The latter finding implies the independent evolution of the rare double-opposite peristome type in both the protohaplolepideous grade and the core haplolepideous clade.
... Among extant families, only Mymaridae and Rotoitidae are definitively known from any fossils older than Eocene (Poinar and Huber 2011;Gumovsky et al. 2018). While other families such as Eulophidae and Trichogrammatidae were reported from putatively Cretaceous Ethiopian amber fossils by Schmidt et al. (2010), later data (Perrichot et al. 2016(Perrichot et al. , 2018 have shown Ethiopian amber to be Early Miocene in origin (16-23 Ma). ...
New Baltic amber species of Pteromalidae sensu lato are described, from two different subfamilies, Asaphesinae n. n. and Eunotinae. Asaphesinae is provided as a replacement name for Asaphinae Ashmead 1904, which is a junior homonym of the trilobite family Asaphidae Burmeister 1843. Coriotela lasallei gen. n., sp. n.. and Butiokeras costae gen. n., sp. n.. are described as the first known fossil species of Asaphesinae and Eunotinae, respectively. These species establish the minimum known age of both groups in the Eocene. Taxonomic changes are also proposed for some extant species. The genus Desantisiana Neder de Román syn. n.. is found to be a junior synonym of Notoglyptus, and its only described species is transferred as Notoglyptus jujuyensis (Neder de Román) comb. n.. The tribe Calyconotiscini, previously classified in Eunotinae, is abolished and Calyconotiscus Narendran & Saleem is transferred to Pireninae.
Full-text available
Insect parasites and parasitoids are a major component of terrestrial food webs. For parasitoids, categorization is whether feeding activity is located inside or outside its host, if the host is immobilized or allowed to grow, and if the feeding is done by one or many conspecific or heterospecific individuals, and other features. Fossil evidence for parasitism and parasitoidism consists of taxonomic affiliation, morphology, gut contents, coprolites, tissue damage and trace fossils. Ten hemimetabolous and holometabolous orders of insects developed the parasite condition whereas seven orders of holometabolous insects evolved the parasitoid life habit. Modern terrestrial food webs are important for understanding the Mid Mesozoic Parasitoid Revolution. The MMPR began in late Early Jurassic (Phase 1), in which bottom-to-top regulation of terrestrial food webs dominated by inefficient clades of predators were replaced by top-to-bottom control by trophically more efficient parasitoid clades. The MMPR became consolidated in Phase 2 by the end of the Early Cretaceous. These clades later expanded (phases 3 and 4) as parasitoids became significant ecological elements in terrestrial food webs. Bottom-to-top food webs explained by the resource concentration hypothesis characterize pre-MMPR time. During phases 1 and 2 of MMPR (Middle Jurassic to Early Cretaceous), a shift ensued toward top-to-down food webs, explained by the trophic cascade hypothesis, exemplified by hymenopteran parasitoid clades Stephanoidea and Evanioidea. Clade-specific innovations spurring the MMPR included long, flexible ovipositors (wasps), host seeking, triungulin and planidium larvae (mantispids, beetles, twisted-wing parasites, flies), and extrudable, telescoped ovipositors (flies). After the MMPR, in phases 3 and 4 (Late Cretaceous to Recent), parasitoids increased in taxonomic diversity, becoming integrated into food webs that continue to the present day.
Full-text available
This atlas opens a window onto the world of Baltic amber and the Eocene amber forests. About 650 typical plants and animals are systematically represented in 92 full-page color plates. The detailed text and bibliography allow an up-to-date overview of the taxonomy, paleontology and biology of fossil plants and animals. This book will not only delight everyone interested in Baltic amber and its fascinating inclusions but also serves as an authoritative reference book to the scientific community.
Full-text available
The only window to the diverse terrestrial biotic component comprising insects and spiders is through their preservation in amber nodules commonly found in coals and lignites, from the Permian onwards. Although a few isolated insect remains have been reported from India, the present correspondence represents the first record of a diverse biota and provides insights into fossil insect and spider biodiversity, not available through other processes of fossilization. The present find has the potential of providing significant information on the migration and degree of isolation during the drift phase of the Indian plate prior to the India-Asia collision. Palaeo_biogeographical relationships of insects and spiders have been used in the context of geodynamicplate tectonic models to infer multiple migrational histories for the central Americas and Madagascar_Africa connection. The excellent state of preservation of the DNA characterization and in understanding of the diversity and antiquity of modern-day South Asiatic insect and arachnid communities.
Studies on amber—fossil resin—have attracted much attention from the scientific community in the last decade. Seven Ph.D. theses with amber as the main material of study recently have been completed (Penney, 1999; Azar, 2000; Schmidt, 2003; Perrichot, 2005; Grund, 2006; Solorzano Kraemer, 2007; Girard, 2008), and two more are currently in progress by Malvina Lak and Jaime Ortega Blanco. From our personal bibliographic databases, we estimate that >1080 papers and books on amber have been published between 1999 and 2008 (as of October 2008). These works have dealt not only with the systematics and phylogeny of arthropods, plant remains, and microorganisms preserved as inclusions in the resin but also with the biogeography, taphonomy, paleoecology, paleoenvironmental reconstructions, chemical and physical properties of amber, its archaeological value, and the development of new methodologies for the analysis of its fossil content. A reason for this recent trend, but a consequence of it, too, was the discovery of several new amber deposits during the last 10 years—from the Triassic of Italy (Roghi et al., 2006), the Jurassic of Thailand (Philippe et al., 2005), the Cretaceous of France, South Africa, Spain, and Wyoming in the United States (Alonso et al., 2000; Grimaldi et al., 2000; Gomez et al., 2002; Neraudeau et al., 2002; Guiliano et al., 2006; Pen alver et al., 2007), the Eocene of France, India, and Italy (Nel et al., 1999; Alimohammadian et al., 2005; Trevisani et al., 2005), and the Miocene of Peru (Antoine et al., 2006). In France, Didier Neraudeau, professor of paleontology at the University of Rennes 1, revived studies on Cretaceous amber in 1999 when he discovered the deposit of Archingeay–Les Nouillers, in Charentes, which was soon followed by the discovery of five other deposits …