◄ Carnets Geol. 22 (16) ►
Outline:
[1. Introduction]
[2. Historical background]
[3. Geological settings]
[4. Importance of the Lebanese fossil insects]
[5. Conclusions]
[Bibliographic references]
[Plates] and ...
[Appendix]
ORCID iD: 0000-0003-4004-6735
Lebanese University, Faculty of Sciences II, Department of Natural Sciences,
Fanar, P.O. Box 26110217, Fanar-Matn (Lebanon);
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of
Geology and Palaeontology, and Center for Excellence in Life and
Paleoenvironment, Chinese Academy of Sciences, Nanjing, Jiangsu 210008 (People's
Republic of China)
ORCID iD: 0000-0001-9468-2353
2 impasse Charles Martel, 29217 Plougonvelin (France)
ORCID iD: 0000-0002-4485-197X
* corresponding author
Lebanese University, Faculty of Sciences II, Department of Natural Sciences,
Fanar, P.O. Box 26110217, Fanar-Matn (Lebanon);
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of
Geology and Palaeontology, and Center for Excellence in Life and
Paleoenvironment, Chinese Academy of Sciences, Nanjing, Jiangsu 210008 (People's
Republic of China)
Published online in final form (pdf) on September 14, 2022
DOI
10.2110/carnets.2022.2216
[Editor:
Michel Moullade;
technical editor: Bruno R.C. Granier]
With 35 Cretaceous outcrops yielding fossil insects, either in amber or as rock (marls, limestones, cinerite, or dysodile) impressions-compressions (adpressions), Lebanon has continuously contributed significantly to the advance of palaeoentomology and to our understanding of entomological evolution and palaeobiodiversity. Compared to its small surface area, this country can be considered among the richest of fossil insect outcrops. This is due to its geological history and mainly to its forest, fluvial - lacustrine tropical and proximal marine subtropical palaeoenvironments plus Peritethys equatorial and subequatorial palaeogeography during the Lower and "Middle" Cretaceous. Herein, an exhaustive review of all outcrops with insects is given. A list of all fossil insects described from Lebanon is provided.
• amber;
• Cretaceous;
• dysodile;
• fossil insects;
• adpression;
• palaeoenvironment;
• palaeobiodiversity
Maksoud S., Granier B.R.C. & Azar D. (2022).- Palaeoentomological (fossil insects) outcrops in Lebanon.- Carnets Geol., Madrid, vol. 22, no. 16, p. 699-743.
Affleurements paléoentomologiques (insectes fossiles) au Liban.- Avec 35 affleurements crétacés recelant des insectes fossiles, soit dans de l'ambre, soit sous forme d'impressions-compressions (adpressions) de roches (marnes, calcaires, cinérite ou dysodile), le Liban a depuis toujours significativement contribué aux progrès de la paléoentomologie et à l'amélioration de notre compréhension de l'évolution des insectes et de leur paléobiodiversité. En dépit de sa petite superficie, ce pays peut être considéré comme l'un des plus riches en gisements fossilifères ayant fourni des insectes. Ceci est dû principalement à son histoire géologique dans le cadre paléogéographique de la Péritéthys au cours du Crétacé inférieur et "moyen", i.e., dans un domaine équatorial ou subéquatorial avec notamment des paléoenvironnements fluvio-lacustres à proximité de forêts tropicales ou des paléoenvironnements marins proximaux subtropicaux. Un examen exhaustif de tous les affleurements avec des insectes a été réalisé et une liste détaillée de tous les insectes fossiles du Liban est fournie.
• ambre ;
• Crétacé ;
• dysodile ;
• insectes fossiles ;
• adpression ;
• paléoenvironnement ;
• paléobiodiversité
Insects are the most diverse group of animals on the planet and as such are present in a wider variety of habitats than most other complex organisms (Grimaldi & Engel, 2005).
Palaeoentomology (a branch of entomology that deals with fossil insects and related terrestrial arthropods) started in its present scientific and taxonomic form in the late eighteenth century, shortly after the foundation of modern taxonomy with the 10th edition of Linnaeus' "Systema Naturae", when some papers commencing with one by Blochs (1776) on the curiosities of insects entombed in fossil resins were published. It is however noteworthy to state that before this, fossil insects were mentioned several times, viz. in Aristotle's "Zoologia", in Marcus Valerius Martialis' "Epigrammaton libri", in Plinius Secundus' "Naturalis Historia", in Sir Francis Bacon's "The historie of life and death…", in famous Emmanuel Kant's quotes, and especially in Nathanael Sendelius' "Historia succinorum…" on amber and its inclusions and many others (Szwedo, 2011).
The beginning of the nineteenth century (with the growing interest in geological sciences and prehistoric life) witnessed the first attempts to study and describe insects from sedimentary rocks (D. Azar et al., 2018). This discipline then developed during the nineteenth and beginning of the twentieth centuries, slowly but constantly, and resulted in some major works and reviews (e.g., Handlirsch, 1906-1908).
At the beginning of the twenty-first century, with the growing interest in fossil insects and globalisation, several serious multidisciplinary and collaborative scientific teams have been formed in many countries resulting in a noticeable increase in the number of annually published works during the past two decades, form dozens to hundreds.
Palaeoentomology is nowadays developing significantly and exponentially. This discipline is undergoing an intellectual radiation with the discovery of new rock and amber outcrops with fossil insects of different geological ages and in various parts of the world (D. Azar et al., 2018). It is noteworthy to state that since its beginning, palaeoentomology covered not only descriptive aspects of terrestrial arthropods (including Insecta, Chelicerata, Myriapoda, etc.) but also reconstructions of ancient environments, ecology, evolution and phylogenies.
Herein we present an exhaustive review of all the 35 outcrops yielding
fossil insects in Lebanon, either as rock adpressions or amber inclusions (Fig. 1 
) and we provide an updated list of hitherto described Lebanese fossil insects.
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Figure 1: Location map of Lebanese outcrops with fossil insects. Green areas indicate the distribution of the amber localities. Yellow
circular spots indicate the locations of Lower Cretaceous amber outcrops with
insect inclusions. Red squares indicate the locations of the outcrops with
fossil insects preserved as compression-impression. Amber outcrops with insects:
(1) Mechmech (Ain El-Khyar); (2) Nimrin (El-Dabsheh); (3) Brissa;
(4) near Bcharreh; (5) Beqaa Kafra; (6) Hadath El-Joubbeh; (7)
Tannourine;
(8) Mazraat Kfardibiane; (9) Ouata El-Jaouz; (10) Bqaatouta
(El-Shqif); (11) Baskinta (Qanat Bakish); (12) Daychouniyyeh; (13) Kfar
Selouan; (14) Kfar Selouan (Khallet Douaiq); (15) Mdeyrij-Hammana; (16)
Falougha; (17) Ain Zhalta;
(18-19) Ain
Dara (two localities); (20) Sarhmoul; (21) Roum - Aazour -
Homsiyeh;
(22)
Bkassine (Jouar Es-Souss); (23) Wadi Jezzine; (24) Maknouniyeh; (25)
Rihane;
(26) Esh-Sheaybeh; (27) Bouarij; (28) Aita El-Foukhar; (29) Ain Zhalta
(Ain Azimeh). Outcrops with fossil
insects preserved as compression-impression: (A) Qnat; (B) Hjoula;
(C) Nammoura;
(D) Qahmez; (E) Jdeidet Bkassine; (F) Sniyya. Red curves: boundaries of
Governorates; blue curves: boundaries of districts. |
Lebanon officially joined "the club" of the countries with fossil
insect localities in 1888, when Hermann Julius Kolbe (b.1855-d.1939) (Fig.
2.A ) described the trace of an insect larva (to which he gave a
scientific name, Curculionites senonicus
Kolbe, 1888: 136, Pl. XI, fig. 8; herein: Fig. 2.B ) in silicified wood from the
late Santonian lithographic limestone of Sahel Alma. Anton Handlirsch
(b.1865-d.1935) later (1906-1908, p. 665) changed the name of the insect that is
supposed to make this trace to Curculidium
senonicum. The outcrop of Sahel Alma is world famous for its fossil fishes (Davis,
1887). The oldest written evidence of this site dates back to the fourth century
AD when Eusebius of Caesarea (circa b.263-d.339) (often called Eusebius Pamphili), the bishop of Caesarea
Palaestina, evoked these mysterious stones found in Lebanon and
considered them as the witnesses of Noah's deluge. The most famous
mention of this site probably appears in the
writings of Jean de Joinville (b.1224-d.1317) -one of the great chroniclers of Medieval France- who tells how a
fossil fish was presented to King Louis IX ("Saint Louis") (b.1214-d.1270) during one of his crusades to the Middle
East.
|
|
Figure 2:
A-
Hermann Julius Kolbe (1855-1939), a German entomologist from
Halle,
Westphalia. He was curator at the Berlin Zoological Museum from 1890 until 1921
specialising in Coleoptera, Psocoptera and Neuroptera. B- Fossil insect trace in
silicified wood from Sahel Alma. C- Professor Aftim Acra (1922-2007), with his
well-known collection from the outcrop of Jouar Es-Souss (Bkassine). |
Phoenicians were probably the first tradesmen of amber in the Mediterranean (McDonald, 1940) and also the pioneers of the amber maritime route towards the shores of Northern Europe (Baltic area) to obtain the golden fossil resin in exchange for bronze between the thirteenth and fourteenth centuries BC. According to some authors, based on recent archeological discoveries, the amber was collected in Phoenicia (today Lebanon, Syrian coast and Northern Israel) and marketed in the Middle East by Phoenicians until Baltic amber, which is of better gemological quality, became available (Williamson, 1932; Nissenbaum, 1975). The oldest reliable publications describing fossil insects from Lebanon are those of Willi Hennig (b.1913-d.1976) and Dieter Schlee, both in 1970. These concerned fossil insects in Lower Cretaceous amber. Prior to 1994, only one amber outcrop with fossil insects was known, viz. Jouar Ess-Souss in Bkassine (Caza Jezzine, Southern Lebanon). Recent field work increased the number of amber localities with biological inclusions to 29 (D. Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021a, 2021b).
In modern times, although the presence of amber in Lebanon has been documented several times by several authors since the beginning of the nineteenth century, occasionally while describing coal or lignite extraction in mines (Desmarest, 1811; Kastner, 1831; Botta, 1831; Brucchi, 1842; Russegger, 1843; Ritter, 1854; Fraas, 1876, 1878; John, 1878; Cuinet, 1896) and in the twentieth century (Zumoffen, 1926; Dubertret, 1950, 1951a, 1953, 1955), it was only late in the nineteen-sixties that fossil insects were recorded in this source (Schlee, 1970; Schlee & Dietrich, 1970; Hennig, 1970). It is noteworthy that the first geological maps of Lebanon made by Paul-Émile Botta (b.1802-d.1870), then by Joseph Ritter von Russegger (b.1802-d.1863), had the aim of locating the lignite and iron mines in order to exploit them for both fuel and industry.
In
1962, Aftim Acra (b.1922-d.2007) (Fig. 2.C ), while leading a
palaeontological expedition in Daher-El-Baydar (Mount-Lebanon, Central Lebanon)
and accompanied by his son Fadi (and Raif Milki), found a piece of amber.
From then till the 1970s, they found several amber outcrops including the
well-known one of Bkassine (Jouar Ess-Souss) in the Jezzine area, which was
discovered independently and at the same time by a German expedition organised
in 1968-1969.
The German expedition was carried out after M. Warth gave Willi Hennig (in 1967) some samples of amber from Bkassine (Southern Lebanon) housed in the Ludwigsburg collection of Stuttgart Museum (Germany). These samples were the remains of the collection of Oscar Fraas (b.1824-d.1897), a German geologist who was tasked by Rustem Pasha (Rustem Mariani b.1810-d.1885), the Italian Governor of Mount Lebanon (1873-1883), to study the geology of the region in order to establish coal mines.
From 1994 until the present day, the team of one of us (DA) has found about 450 amber-bearing deposits, ranging from the Late Jurassic to the Cenomanian (Late Cretaceous). Among these numerous outcrops only 29 hitherto yielded fossil insects. It is noteworthy that the clay and shale in one of the amber outcrops (Qahmez, in Kesserouan District, Central Lebanon, unpublished data) includes some fragments of insects originally floating with other organic and plant remains, deposited in an abandoned reaches of a fluvial system.
The rediscovery of dysodiles, unusual Lower Cretaceous continental deposits in Lebanon, brought important and exceptional palaeontological assets (El Hajj et al., 2019, 2021a, 2021b; D. Azar et al., 2019a). Dysodiles are sedimentary rocks characterised by finely laminated (micrometric) layers (papershales), with high organic content (Cordier, 1808) and well-preserved fossils (Fraas, 1878). Lebanese dysodiles were formed in different lacustrine deposits in different areas, during the lower Barremian and Albian. Their presence was mentioned in some nineteenth century publications in the Lower Cretaceous sandstones of Mount Lebanon (Botta, 1831), and in the South (Fraas, 1878) and Centre of Lebanon (Janensch, 1925). Fraas (1878) was the first author to point out their richness in fossils including fish and plant debris, and he identified some plant species that were later reviewed by Edwards (1929) and attributed to Zamites sp. and Weichselia reticulata. In 1925, Janensch identified two small actinopterygian fishes as Pleuropholis koerti and Thrissops sp. Then, since the 1930s, dysodiles were forgotten until their recent rediscovery through the extensive geological fieldwork of our team which resulted in the finding of seven localities, five of them in the lower Barremian and two in the Albian. Among those dysodile outcrops, two lower Barremian ones (Jdeidet Bkassine and Sniyya) and one Albian (Qnat, where dysodiles are associated with cinerite, unpublished data) produced fossil insects. Recently D. Azar et al. (2019a) described an ephemeropteran larva and El Hajj et al. (2021a) illustrated several fossil insects from the lower Barremian dysodiles of Jdeidet Bkassine.
Apart from amber, Lebanon is world-wide famous in palaeontology for its rich Upper Cretaceous marine fish deposits in Sahel Alma, Haqel, Nammoura, and Hjoula. Recently, the two latter outcrops surprisingly yielded some complete and undisarticulated fossil insects (A. Nel et al., 2004; Vršanský & Makhoul, 2013; D. Azar et al., 2019b; Maksoud & D. Azar, 2021), indicating a depositional marine palaeoenvironment, close to a palaeo-shoreline during the mid-Cenomanian.
Lower Barremian fossil insect outcrops
Amber
Of the 450 amber
outcrops in Lebanon, 430 outcrops contain early Barremian amber,
among which 29 have yielded amber with bioinclusions. Up to 10% of Lebanon's total
land surface could potentially yield amber (Fig. 1 ).
The list of the fossiliferous amber outcrops is given in Table 1 ranging from the North to South of Lebanon.
Table 1: Barremian
(Lower
Cretaceous) amber outcrops with arthropod (mainly insect) inclusions. The
numbers in bold between brackets correspond to the placement of the localities
on the general map of Figure 1 .
| Governorate | District | Outcrop | Inclusions | References |
| Akkar | Akkar | Mechmech (Ain El-Khyar) [1]; Fig. 3.A-B ;
Pl. 1 , fig. A-D |
4 | Maksoud et al., 2019, 2021b, 2021c |
| North Lebanon | Sir Ed-Danniyeh | Nimrin (El-Dabsheh)
[2]; Fig. 3.A-B ;
Pl. 1 , fig. E |
~250 | D. Azar et al., 2010b; Szwedo et al., 2013; Maksoud & D. Azar, 2020; Maksoud et al., 2021c |
| Brissa [3]; Fig. 3.A, D ;
Pl. 1 , fig. F |
~100 | Kirejtchuk & D. Azar, 2013; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | ||
| Bcharreh | near Bcharreh [4]; Fig. 4.A ;
Grimaldi
& Engel, 2005:
80, Fig. 2.56 |
~1000 | Grimaldi & Engel, 2005; Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | |
| Beqaa Kafra [5]; Fig. 4.A-B ;
Pl. 1 , fig. G |
6 | D. Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | ||
| Hadath El-Joubbeh [6]; Fig. 4.A, C ;
Pl. 1 , fig. H |
~5 | D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | ||
| El-Batroun | Tannourine [7]; Fig. 4.A, D ;
Pl. 1 , fig. I |
47 | D. Azar & Ziadé, 2005; Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | |
| Jbeil - Kesserouan | Kesserouan | Mazraat
Kfardibiane [8]; Fig. 6.A-B ;
Pl. 1 , fig. J |
2 | Maksoud et al., 2020, 2021c |
| Ouata El-Jaouz [9]; Fig. 6.A, C ;
Pl. 2 ,
figs. A-B |
6 | D. Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | ||
| Bqaatouta [10]; Fig. 6.A, E ;
Pl. 2 ,
figs. C-E |
~40 | Maksoud et al., 2021a | ||
| Mount Lebanon | El-Matn | Baskinta [11]; Fig. 6.A, D ;
Pl. 2 ,
figs. F-G |
~40 | Maksoud et al., 2021a |
| Daychouniyyeh [12]; Fig. 7.A-B ;
Pl. 2 , fig. H |
11 | D. Azar et al., 2010b; Szwedo et al., 2011; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | ||
| Baabda | Kfar
Selouan [13]; Fig. 8.A-B ;
Pl. 2 , figs. I-J |
69 | D. Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | |
| Kfar Selouan (Khallet Douaiq) [14]; Fig.
8.A, C ;
Pl. 2 , figs. K-M |
37 | Maksoud et al., 2021c | ||
| Mdeyrij-Hammana
[15]; Fig. 8.A, E ;
Pl. 3 ,
figs. A-E |
3200 | D. Azar et al., 1999, 2010b, 2011a; D. Azar, 2012; Szwedo et al., 2013; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | ||
| Falougha [16]; Fig. 8.A, D ;
Pl. 3 , fig. F |
~40 | D. Azar et al., 2015; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | ||
| Esh-Shouf | Ain Zhalta [17]; Fig. 9.A-B ;
Pl. 3 , fig. G |
20 | D. Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | |
| Ain Zhalta (Ain Azimeh) [29]; Fig. 9.A-B ;
Maksoud
et al.,
2022: 401, Fig. 1; 402, Fig. 2 |
30 | Maksoud et al., 2022 | ||
| Aley | Ain
Dara (two localities) [18-19]; Fig. 9.A, F ;
Pl. 3 , fig. H |
130 | D. Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | |
| Sarhmoul [20]; Fig. 10.A-B ;
Pl. 3 , fig. I |
29 | D. Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | ||
| South Lebanon | Jezzine | Roum - Aazour - Homsiyeh [21]; Fig. 11.A,
C ;
Pl. 3 , fig. J |
37 | D. Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c |
| Bkassine (Jouar Es-Souss) [22]; Fig. 11.A,
D ;
Pl. 3 , fig. K |
~3000 | Schlee & Dietrich, 1970; Azar et al., 2010b; Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | ||
| Wadi Jezzine [23]; Fig. 11.A-B ;
Pl. 3 , fig. L |
~20 | Maksoud & D. Azar, 2020; Maksoud et al., 2021c | ||
| Maknouniyeh [24]; Fig. 11.A, E ;
Pl. 3 , fig. M |
14 | D. Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | ||
| Rihane [25]; Fig. 12.A-B ;
Pl. 3 , fig. N |
~40 | D. Azar & Nel, 2013; Szwedo et al., 2013; Maksoud & D. Azar, 2020; Maksoud et al., 2021c | ||
| Baalbeck - El Hermel | Baalbeck | Esh-Sheaybeh [26]; Fig. 13.A-B ;
Pl. 3 , fig. O |
22 | D. Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c |
| Beqaa | Zahleh | Bouarij [27]; Fig. 9.A, G ;
Pl. 3 , fig. P |
220 | D. Azar et al., 2010b; D. Azar, 2012; Maksoud et al., 2021c |
| Rashaiya | Aita El-Foukhar
[28]; Fig. 14.A-B |
1 | D. Azar et al., 2010b; D. Azar, 2012; Maksoud & D. Azar, 2020; Maksoud et al., 2021c |
|
|
Figure 3: A-
Geological map: fossiliferous outcrop of Mechmech (Ain El-Khyar) (red arrow)
outcrop (modified after Dubertret, 1951c;
Guerre, 1975). J6 = uppermost Jurassic; C2a = lower Barremian "Grès
du Liban" sandstone; C2b = Barremian clay and oolitic deposition of the upper
part of the "Grès du Liban" and oolitic deposition of the lower part of the
Jezzinian; C3
= micritic part of the Jezzinian (uppermost Barremian-lowermost Aptian); C4 =
Albian; C4-5 = Albian-Cenomanian; C5 = Cenomanian; Q = Quaternary scree; βC3 = Jezzinian
volcanic deposition. Thick lines represent faults. Scale bar = 1 km. B-
General view of the outcrop of Mechmech (Ain El-Khyar). |
|
|
Figure 4: A-
Geological map: fossiliferous outcrops of Nimrin (El-Dabsheh) (yellow arrow) and
Brissa (red arrow) outcrops (modified after Dubertret, 1951c). J6 =
uppermost Jurassic; C2 = lower Barremian "Grès du Liban"; C3 = Jezzinian
(uppermost Barremian-lowermost Aptian); C4-5 = Albian-Cenomanian; C5 =
Cenomanian; Q = Quaternary scree; βC3 = Jezzinian
volcanic deposition. Thick lines represent faults. Scale bar = 1 km. B-
General view of Nimrin (El-Dabsheh) outcrop. C- General view of Brissa outcrop. |
|
|
Figure 5: A-
Geological map: fossiliferous outcrops of Bcharreh (yellow arrow), Beqaa Kafra
(blue arrow), Hadath El-Joubbeh (orange arrow), Tannourine (brown arrow) and
Qnat (clear blue star) outcrops (modified after Dubertret & Wetzel,
1945, 1951; Dubertret,
1949b, 1951c). J6 = uppermost Jurassic; C2a = lower Barremian
"Grès
du Liban" sandstone; C2b = Barremian clay and oolitic deposition of the upper
part of the "Grès du Liban" and oolitic deposition of the lower part of the
Jezzinian; C3
= micritic part of the Jezzinian (uppermost Barremian-lowermost Aptian); C4 =
Albian; C5 = Cenomanian; Q = Quaternary scree; βJ6 =
Kimmeridgian volcanic deposition; βC3 = Jezzinian volcanic deposition. Thick lines represent faults. Scale bar = 1 km.
B- General view of Beqaa
Kafra outcrop. C- General view of Hadath El-Joubbeh outcrop. D- General view of
Tannourine outcrop. |
|
|
Figure 6: A- Geological map: Ouata
El-Jaouz
(yellow arrow), Mazraat Kfardibiane (Red Rock) (orange arrow), Baskinta (Qanat
Bakish) (red arrow), Bqaatouta (blue arrow) and Qahmez (purple star) outcrops
(modified from Dubertret, 1945, and Dubertret & Wetzel,
1945). J6 = uppermost Jurassic; C1 = Salima Formation (lower Valanginian); C2a =
lower Barremian "Grès du Liban" sandstone; C2b = Barremian clay and oolitic
deposition of the upper part of the "Grès du Liban" and oolitic deposition
of the lower part of the Jezzinian; C3 = micritic part of the Jezzinian
(uppermost Barremian-lowermost Aptian); C4 = Albian; C5 = Cenomanian; Q =
Quaternary scree; βJ6 = Kimmeridgian volcanic deposition; βC2 = lower
Barremian volcanic deposition. Thick lines represent faults. Scale bar = 1
km. B- General view of Mazraat
Kfardibiane outcrop. C- General view of Ouata El-Jaouz
outcrop. D- General view of Baskinta (Qanat Bakich) outcrop. E- General view of Bqaatouta outcrop. |
|
|
Figure 7: A- Geological map: Daychouniyyeh (red
arrow) outcrop (modified from Dubertret, 1951a). J6 = uppermost Jurassic; C2a =
lower Barremian "Grès du Liban" sandstone; C2b = Barremian clay and oolitic
deposition of the upper part of the "Grès du Liban" and oolitic deposition
of the lower part of the Jezzinian; C3 = micritic part of the Jezzinian
(uppermost Barremian-lowermost Aptian); C4 = Albian; C5 = Cenomanian; Q =
Quaternary scree; M = mid-Miocene. Thick lines represent faults. Scale bar =
1 km. B- General view of Daychouniyyeh outcrop. |
|
|
Figure 8: A- Geological map: Kfar Selouan
(orange arrow), Kfar Selouan (Khallet Douaiq) (purple arrow), Bouarij,
Mdeyrij-Hammana (brown arrow), Falougha (red arrow) and Ain Dara (yellow arrow)
outcrops (modified from Dubertret, 1951a,
1953). J6 = uppermost Jurassic;
C2a = lower Barremian "Grès du Liban" sandstone; C2b = Barremian clay and
oolitic deposition of the upper part of the "Grès du Liban" and oolitic
deposition of the lower part of the Jezzinian; C3 = micritic part of the
Jezzinian (uppermost Barremian-lowermost Aptian); C4 = Albian; C5 = Cenomanian; Q = Quaternary
scree; βC2 = lower Barremian volcanic
deposition. Thick lines represent faults. Scale bar = 1 km. B- General
view of Kfar
Selouan outcrop. C- General view of Kfar
Selouan (Khallet Douaiq) outcrop. D-
General view of Falougha outcrop. E- General view of Mdeyrij-Hammana
outcrop. F- General view of Ain Dara outcrop. G- General view of Bouarij
outcrop. |
|
|
Figure 9: A-
Geological map: Ain Zhalta (red arrow) and Ain Zhalta (Ain Azimeh, blue arrow) (modified from Dubertret,
1950, 1951a). J6 = uppermost Jurassic; C2a = lower Barremian "Grès du Liban"
sandstone; C2b = Barremian clay and oolitic deposition of the upper part of the
"Grès du Liban" and oolitic deposition of the lower part of the
Jezzinian; C3 = micritic part of the Jezzinian (uppermost Barremian-lowermost
Aptian); C4 = Albian; C5 = Cenomanian; Q = Quaternary scree. Thick lines
represent faults. Scale bar = 1 km. B- General aerial view of Ain Zhalta (Ain
Azimeh) outcrop. Scale bar = 50 m. C- General view of Ain Zhalta outcrop. |
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Figure 10: A- Geological map: Sarhmoul
(red arrow) (modified
from Dubertret, 1949c, 1950,
1951b). J6 = uppermost Jurassic; C1 = Salima
Formation (lower Valanginian); C2a = lower Barremian "Grès du Liban"
sandstone; C2b = Barremian clay and oolitic deposition of the upper part of the
"Grès du Liban" and oolitic deposition of the lower part of the
Jezzinian;
C3 = micritic part of the Jezzinian (uppermost Barremian-lowermost Aptian); C4
= Albian; C5 = Cenomanian; Q = Quaternary scree; βJ6 = Kimmeridgian
volcanic deposition. Thick lines represent faults. Scale bar = 1 km. B- General view of
Sarhmoul outcrop. |
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|
Figure 11: A- Geological
map: Jdeidet Bkassine (blue star) and Sniyya (red star) dysodile outcrops and of
the Roum - Aazour - Homsiyeh (yellow
arrow), Bkassine (Jouar Es-Souss) (red arrow), Wadi Jezzine (purple arrow),
Maknouniyeh (orange arrow) amber outcrops (modified from Dubertret,
1950).
J6 = uppermost Jurassic; C1 = Salima Formation (lower Valanginian); C2a = lower
Barremian "Grès du Liban" sandstone; C2b = Barremian clay and oolitic
deposition of the upper part of the "Grès du Liban" and oolitic deposition
of the lower part of the Jezzinian; C3 = micritic part of the Jezzinian
(uppermost Barremian-lowermost Aptian); C4 = Albian; C5 = Cenomanian; Q =
Quaternary scree; βC2 = Barremian volcanic deposition. Thick lines
represent faults. Scale bar = 1 km. B- General view of Wadi
Jezzine outcrop. C- General view of Roum
- Aazour - Homsiyeh outcrop. D- General
view of Bkassine (Jouar Es-Souss) outcrop. E- General
view of Maknouniyeh outcrop. |
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|
Figure 12: A- Geological
map: Rihane (red arrow) amber outcrop (modified
from Dubertret, 1951b). J6 = uppermost Jurassic; C1 = Salima Formation
(lower Valanginian); C2 = lower Barremian "Grès du Liban" sandstone; C3 =
micritic part of the Jezzinian (uppermost Barremian-lowermost Aptian); C4 =
Albian; C5 = Cenomanian; Q = Quaternary scree; βC2 = Barremian volcanic
deposition. Thick lines represent faults. Scale bar = 1 km. B- General view of
Rihane outcrop. |
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Figure 13: A- Geological
map: Esh-Sheaybeh (red arrow) amber outcrop (modified
from Dubertret, 1949b). J4 = Middle Jurassic; J5 = Oxfordian; J6 =
uppermost Jurassic, Kimmeridgian; C1 = Salima Formation (lower Valanginian); C2 =
lower Barremian "Grès du Liban" sandstone; C3 = micritic part of the
Jezzinian (uppermost Barremian-lowermost Aptian); C4 = Albian; C5 = Cenomanian; C4-5 = Albian -
Cenomanian. Thick lines represent faults. Scale
bar = 1 km. B- General view of Esh-Sheaybeh outcrop. |
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Figure 14: A- Geological
map: Aita El-Foukhar (red arrow) amber
outcrop (modified
from Dubertret, 1960). J6 = uppermost Jurassic; C2 = lower Barremian
"Grès
du Liban" sandstone; C3 = micritic part of the Jezzinian (uppermost
Barremian-lowermost Aptian); C4 = Albian; C5 = Cenomanian. Thick lines
represent faults. Scale bar = 1 km. Thick lines represent faults. Dotted
line represents the boundary between Lebanon and Syria. Scale bar = 1 km. B- General view of
Aita El-Foukhar outcrop. |
The clay-shale
layers of one of the amber outcrops, in Qahmez (Kesserouan District, Central
Lebanon, Káčerová & D. Azar,
in press) (Figs.
6.A ,
15.A ) produced some fossil fragments of insects originally floating with
other organic and vegetal remains, deposited in an abandoned reach of a fluvial
system. The outcrop today is a sand quarry (a vulnerable site that almost
disappeared with the growth of the quarry); the sand layers are intercalated
with centimetric-scale layers of grey silty clay-shale rich in floated organic
remains. In one of these layers (the thickest, ca. 50 cm thick)
corresponding to a lens of ca. 6 metres width, formed by a palaeo-fluvial
channel, a cockroach forewing (belonging to Mesoblattinidae) was recovered (Pl. 4 , fig.
A) (Káčerová & D. Azar,
in press).
Dysodiles
Lower Barremian
dysodiles crop out in five localities across Lebanon (Fig. 1 ) at the base of the
"Grès du Liban" unit; in Qrayn (North of Lebanon, Sir Ed-Danniyeh
District), Tarchich (Central Lebanon, Baabda District), Jdeidet Bkassine, Sniyya
and Zhalta (South of Lebanon, Jezzine District) (El Hajj et al.,
2021a).
Though the five
outcrops all present a great potential for fossil insects, nevertheless to date
only two of them, viz. Jdeidet Bkassine (Figs. 11.A ,
15.B ) and Sniyya
(Figs. 11.A ,
15.C ), produced some insect samples
(Pl. 4 , figs. B-E, Q) among which a mayfly
larva, Libanoephemera inopinatabranchia
D. Azar et al., 2019b, and an adult cockroach belonging to
Blattulidae (Káčerová
& D. Azar, in press) were described. These
paper shales (ca. 95-100 cm thick) were formed in small, isolated
palaeolakes and/or swamps, within the "Grès du Liban" unit.
Albian fossil insect outcrop
Cinerite associated with dysodiles
In
Qnat (Bcharreh District, North of Lebanon), Albian cinerite and a dysodile
associated with volcanism yielded fishes, vegetal remains, turtles, and insects
(Figs. 5.A ,
15.D ). To date, a complete articulated Mantodea imago, bearing several
plesiomorphic structures, has been recovered in the cinerite from this outcrop (Pl. 4 , fig. F).
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Figure 15: A-
Clay-shale layers in Qahmez presenting fossil insects’ fragments. B- General
view of the lower Barremian dysodile outcrop in Jdeidet Bkassine. C- General
view of the lower Barremian dysodile outcrop in Sniyya. D- General view of Qnat
outcrop. |
Mid-Cenomanian limestone with fossil insects
Marine fossil fish deposits
Late Cretaceous marine fish deposits in Sahel Alma, Haqel, Nammoura and Hjoula are famously known. Recently the two latter outcrops yielded unexpectedly complete and undisarticulated fossil insects (A. Nel et al., 2004; Vršanský & Makhoul, 2013; D. Azar et al., 2019b; Maksoud & D. Azar, 2021), indicating an exceptional depositional marine palaeoenvironment, close to a shoreline during the mid-Cenomanian.
The
Nammoura outcrop is located in El Ghabour, a valley at the North end of the
village of Nammoura, in the Kesserouan District (Vršanský & Makhoul,
2013), in a quarry dedicated to extracting limestone slabs (Fig.
16.A-C ). To
date, some specimens of the archaeorthopteran Chresmoda libanica A. Nel et al., 2004
(Chresmodidae),
and one specimen of a mesoblattinid cockroach, Mieroblattina pacis Vršanský & Makhoul,
2013 (Pl. 4 , figs. G-H), have been recovered from this locality.
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Figure 16: A- Geological map: Nammoura Lagerstätte (red star) (modified from Dubertret,
1945).
J6 = uppermost Jurassic; C2a = lower Barremian "Grès du Liban" sandstone;
C2b = Barremian clay and oolitic deposition of the upper part of the "Grès du
Liban" and oolitic deposition of the lower part of the Jezzinian; C3 =
micritic part of the Jezzinian (uppermost Barremian-lowermost Aptian); C4 =
Albian; C5 = Cenomanian; C5-6 = Cenomanian - Turonian; βJ6 = Kimmeridgian
volcanic deposition. Thick lines represent faults. Scale bar = 1 km. B- General view of Nammoura
Lagerstätte.
C- Detail of the geological layers in Nammoura outcrop. |
The
Hjoula outcrop is situated in Hjoula (Jbeil District, Northern part of Central
Lebanon), in the valley beneath the mosque of this village (Fig.
17.A-B ). Hitherto,
ten complete and articulated fossil insects were recovered from this outcrop (Maksoud
& D. Azar, 2021): six Odonata (belonging to three or four different
taxa), including one liupanshaniid, Libanoliupanshania
mimi D. Azar et al., 2019b; a libanocorduliid, Libanocordulia
debiei D. Azar et al., 2019b, and two undetermined Anisoptera; a
scarabaeoid coleopteran; a cicadellid hemipteran, a saucrosmylid neuropteran (Lebanosmylus
leae D. Azar & A. Nel,
2022); and an unidentified insect (Pl. 4 ,
figs.
I-P).
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Figure 17: A- Geological map: Hjoula Lagerstätte (red star) (modified from Dubertret, 1945; Dubertret & Wetzel,
1945). J6 = uppermost Jurassic; C2a = lower Barremian
"Grès
du Liban" sandstone; C2b = Barremian clay and oolitic deposition of the upper
part of the "Grès du Liban" and oolitic deposition of the lower part of the
Jezzinian; C3 = micritic part of the Jezzinian (uppermost Barremian-lowermost
Aptian); C4 = Albian; C5 = Cenomanian; Q = Quaternary scree; βJ6 =
Kimmeridgian volcanic deposition; βC3 = Jezzinian volcanic deposition. Thick lines represent faults. Scale bar = 1 km.
B- General view of Hjoula
Lagerstätte. |
3.1. Lower Barremian fossil insect outcrops
Amber
Lower Cretaceous Lebanese amber occurs in silts and dark shales with lignite and plant debris (sometimes including leaves from the resin-producing tree) in three intervals of the upper part of the "Grès du Liban". These deposits correspond to siliciclastic coastland estuarine environments based on the co-occurrence of bioturbation, echinoids and bivalves in the transgressive marine layers and amber and lignite in the regressive layers. The entomofaunal associations of the amber inclusions indicates a thick resin-producing forest under a warm tropical climate, which is also corroborated by the palynological data (D. Azar et al., 2011a).
Lower Cretaceous Lebanese amber found in the lowest interval of the upper part of the "Grès du Liban" is buried in a primary deposit, whilst it has been re-deposited in the mid and upper intervals. Until now, about 8,500 biological inclusions (mainly insects) have been found in the 29 Lebanese amber outcrops.
For a long time, the Lower Cretaceous amber outcrops were dated as Neocomian-Aptian (Zumoffen, 1926) or Neocomian-early Aptian (Dubertret & Vautrin, 1937), or even Valanginian-Hauterivian in age (Schlee & Dietrich, 1970), i.e., various ranges across the whole Early Cretaceous Subperiod, except Albian, relegated to the late Aptian. The age of the overlying limestone unit, i.e., the "Falaise de Blanche" [Blanche Cliff], was also poorly constrained until a recent detailed holostratigraphic study by Maksoud et al. (2014). These authors merged the whitish micritic limestones of the "Falaise de Blanche" with the immediate underlying yellowish grainy limestones in a single lithostratigraphic unit. This Unconformity Bounded Unit, now dubbed the Jezzinian regional stage, is dated as late Barremian-earliest Aptian (= early Bedoulian).
In Lebanon, the
Lower Cretaceous fossiliferous amber outcrops are mainly found in three
intervals in the upper part of the "Grès du Liban" (Fig.
18 ):
• the upper interval is located between the Jezzinian above and the "Banc de Mrejatt" below. Amber outcrops belonging to this interval are those of the waterfall at Jezzine (South Lebanon); Hammana, Kfar Selouan (Khallet Douaiq), Ain Zhalta (Ain Azimeh) and Bouarij (Central Lebanon);
• the middle interval is located between the "Banc de Mrejatt" above and a pisolitic interval below. Amber outcrops belonging to this interval are those of Wadi Jezzine (South Lebanon); Ain Dara and Kfar Selouan (Central Lebanon);
• the lower interval falls below the pisolitic interval. Amber outcrops belonging to this interval are those of Rihane, Maknouniyeh, Roum-Aazour-Homsiyeh and Jouar Es-Souss in Bkassine (South Lebanon); Ain Zhalta, Baskinta, Bqatouta and Mazraat Kfardibiane (Central Lebanon).
Stratigraphic details of these three intervals are given in Maksoud et al. (2017). All three fossiliferous amber-bearing intervals cited above are very rich in biological inclusions, mostly terrestrial arthropod remains such as insects, spiders and mites (D. Azar, 1997a, 1997b, 2012; D. Azar et al., 2010b) but also plants and vertebrate remains. Nevertheless, their entomofaunal associations are very similar, which suggests that they have very close, if not the same age (D. Azar et al., 2003; D. Azar, 2012; Veltz et al., 2013). Because it is commonly assumed that insects—with sometimes more than two generations per year—have a rapid rate of evolution, the entomofaunal similarity of these three intervals could imply that the age of the amber should be the same, i.e., that of the older/lower interval. As a matter of fact, the amber pieces found in the middle and upper intervals are rounded and commonly bored by martesiine pholadid bivalves, suggesting that the resin was already hardened and then transported to a transitional environment (i.e. deltaic zone) where it could be bored by bivalves before being deposited and buried. Therefore, these pebbles have possibly been reworked from the lower interval. In addition, palynomorphs related to the biorecord Superret-Nobarg, which is equivalent to Stellatopollis doylei Ibrahim, 2002, were found as inclusions in the amber (of Hammana), though this taxon does not exist in the palynological assemblage of the embedding sediments (D. Azar et al., 2011a). This too could point to a possible reworking of amber in younger sediments. These interpretations would support those of Veltz et al. (2013), who stated that the amber deposits found in the facies attributed to the obsolete so-called "Abeih Formation" (the middle and higher intervals), have been most likely reworked from older deposits. It is noteworthy to mention that the first use of fossil insects entombed in amber in biostratigraphy and relative dating is from Lebanese amber (D. Azar et al., 2003).
The age of the amber from the Lower Cretaceous or "Grès du Liban" is currently attributed to the early Barremian (Granier et al., 2016; Maksoud et al., 2017; Maksoud & D. Azar, 2020).
Rock impression/compression (adpressions)
The clay-shale layers at Qahmez (Kesserouan District, Central Lebanon, Káčerová & D. Azar, in press) (Pl. 4, fig. A) in the "Grès du Liban" unit are dated as early Barremian by Granier et al. 2015. The sand layers are fluvial and include deltaic deposits not far from the sea with tidal influence, witnessed by the presence of dinoflagellates in the sediment. The sand layers are intercalated with centimetric-thick layers of grey silty clay shale rich in floated organic fragments, especially plant remains, and occasionally insect fragments. The fragmentary state of the organic matter witnesses a transport system before deposition or even flotation after decomposition. One of the silty clay shale rich layers is thick (about 50 cm) and corresponds to a lens of about 6 metres wide, formed in a fluvial channel. The organic remains were originally transported in the fluvial system and deposited in a cut-off arm of water or an oxbow lake.
Dysodile
Lower Barremian dysodiles in South Lebanon (in the three localities of Jdeidet Bkassine, Sniyya and Zhalta) overlie altered volcanic claystone deposits and nodular carbonates. It is noteworthy that these southern outcrops are not distant from each other, suggesting that during deposition they could have been interconnected under certain conditions as demonstrated by the geochemical analyses (El Hajj et al., 2019).
During the lower Barremian, Lebanon was located in the northeastern Gondwana supercontinent, under a warm tropical climate on a continental border. Veltz (2008) highlighted that the deposition of the sandstone occurred in a tectonically unstable continental context. During the lower Barremian, Lebanon also witnessed many volcanic episodes related to fault reactivation (Brew et al., 2001), and faults might have promoted the development of small lakes and/or swamps. The dysodiles are associated with volcanic ash deposits; the volcanic activity occurring both prior to (Dubertret, 1955) and during the dysodile deposition since ash could also be identified within the dysodile layers. In addition, the claystone associated with dysodiles could be the result of an alteration of the basalt. Therefore, we envisage a close relationship between volcanism and dysodile formation.
The palaeogeographic situation of Lebanon, mostly influenced by the tectonic events and the warm tropical climate, may have enhanced the primary productivity and diversity of these lake systems. Volcanic activity would have enriched the lakes and/or swamps with nutrients favouring the proliferation of organisms (Veltz, 2008). However, volcanic activity could have also rendered lake waters toxic, leading to mass mortality (Veltz, 2008).
The excellent preservation of the organic matter and fossils could have been facilitated by calm, anoxic bottom conditions, evidenced by the microscopic fine parallel lamination and absence of bioturbation, as well as by the high content in total organic carbon or TOC (El Hajj et al., 2019), with a Pristane/Phytane ratio lower than 1 (Peters et al., 2005; El Hajj et al., 2019), and the absence of biodegradation. Rapid burial in these aquatic ecosystems could also have contributed to excellent preservation of fossil insects by inhibiting the degradation of organic matter.
3.2. Albian fossil insect outcrop
Cinerite associated with dysodiles
Qnat (Bcharreh District, North of Lebanon) Albian volcanism allowed formation of palaeo-relief that enabled the development of a warm and mineral rich palaeo-lake. This allowed the development of successively repeated bacterial and algal films that were deposited as a dysodile of nearly 30 cm thickness followed by nearly 35 m of cinerites showing successive alternation of layers extremely rich with ostracods and others almost only with cinerites, witnessing a succession of several phases of ostracod blooming that occurred in the nutrient rich and warm water, with each of those phases followed by a generalised asphyxia caused by the continuous deposition of cinerite due to volcanic activity. These cinerites buried rapidly the plants, insects, fishes and turtles.
3.3. Mid-Cenomanian limestone with fossil insects
Marine fossil fish deposits
The Hjoula and Nammoura Konservat-Lagerstätten are located respectively at the villages of Hjoula (Jbeil District, northern part of Central Lebanon) and Nammoura (Kesserouan District, Central Lebanon). They have historically been given several ages depending on different authors. Botta (1833) attributed the sediments in the co-eval Haqel outcrop to the Cretaceous. Lewis (1878) and Fraas (1878) postulated a Turonian age. Later Patterson (1967) and Hückel (1970) recognized a Cenomanian age. Patterson (1967) estimated a mid-Cenomanian age based on the fish fauna, whereas Hückel (1970) considered the beds at Haqel as lower Cenomanian based on the presence of the foraminiferan Orbitolina concava (Lamarck, 1816) and the ammonite Mantelliceras mantelli (Sowerby, 1814) already reported by Zumoffen (1926). Biostratigraphical studies of Dubertret (1959, 1966) and Saint-Marc (1974) determined the sequences at Hjoula to be lower Cenomanian. Hemleben (1977), dated them as late Cenomanian based on a planktonic foraminiferal assemblage, including Praeglobotruncana stephani (Gandolfi, 1942), Rotalipora cushmani (Morrow, 1934) and Rotalipora greenhornensis (Morrow, 1934). Walley (1997) assigned the outcrop to the Sannine Formation. Recently, Wippich and Lehmann (2004) confirmed a late Cenomanian age for Hjoula based on the presence of the ammonite Allocrioceras cf. annulatum (Shumard, 1860), which is a member of the lower upper Cenomanian Sciponoceras gracile Zone in the Western Interior of the USA and the Metoicoceras geslinianum Zone of the international standard. Fuchs et al. (2009) confirmed a late Cenomanian age for Hjoula as well.
During the mid-Cenomanian, Lebanon was mostly submerged and positioned on a shallow carbonate platform (where there were some small islands present) on the North-East of the Arabo-African palaeocontinent at nearly 8° latitude (Philip et al., 1993; Barrier & Vrielynck, 2008). Although the outcrop of Hjoula is clearly marine, continental fossils indicate deposition close to a shoreline during the mid-Cenomanian (more precisely early late Cenomanian), viz. continental plants (pteridophytes, gymnosperms, and angiosperms), invertebrates, and complete skeletons and isolated bone of pterosaurs (Kellner et al., 2019), turtles and fish, in addition to nine newly discovered complete and not disarticulated fossil insects.
According to Krassilov & Bacchia (2000), this mid-Cenomanian floral assemblage differs markedly from both the Lower Cretaceous and the Turonian plant assemblages of the Middle East and represents a distinct stage of the regional floristic evolution. These authors concluded that the phytogeographic affinities of the fossil fish outcrops of the mid-Cenomanian of Lebanon are in like contemporaneous floras of North America, Central Europe, and the Crimea; thus, a combination of features like xeromorphism, the prevalence of compound leaves, and the presence of deciduous angiosperms and gymnosperms may indicate climatic conditions similar to those of the present-day Mediterranean area (Krassilov & Bacchia, 2000).
The above cited outcrops yielding fossil insects are all from the Cretaceous Period, most of them from the Lower Cretaceous (lower Barremian) whilst remainder from the 'mid-Cretaceous' (Albian and Cenomanian). The Cretaceous is one of the most interesting and important geological periods in the history of the Earth. It is when the origin and radiation of the angiosperms took place, and most of the extant insect families first appeared (Grimaldi & Engel, 2005).
The Lebanese amber with its fossil insects is considered by most scientists as one of the most important deposits as it belongs to the Lower Cretaceous. For the first time, a large area with a significant amount of amber appears in the fossil record during the appearance and radiation of the flowering plants (angiosperms) -which is a major event in terrestrial evolution, as angiosperms constitute today more than ¾ths of the world flora. While the relationship between insects and plants is of considerable biological interest, it is important that, if we want to understand the origin of all recent ecosystems, we must go back to the subperiod of their starting point, which is in the Early Cretaceous. Moreover, this subperiod has not an important insect record, except Lebanese amber, which increases considerably the importance of this material. Insects can be found in Lebanese amber every 25 to 30 pieces in fossiliferous amber outcrops. To date, this amber contains no less than about 23 entomological (hexapod) orders, including Archaeognatha, Blattodea, Coleoptera, Collembola, Dermaptera, Dictyoptera (Blattodea, Isoptera, Mantodea), Diptera, Ephemeroptera, Hemiptera, Hymenoptera, Lepidoptera, Mecoptera, Neuroptera, Odonata, Orthoptera, Plecoptera, Psocodea, Strepsiptera, Thysanoptera, Thysanura, and Trichoptera.
A single piece of amber can contain one or several inclusions. Some of those that are found alone may provide important indirect hints of the presence of other organisms or reflect a specific habitat or palaeogeograph extension. Regarding the specimens where several inclusions (or syninclusions) are found together, some of them are present only in a chance manner, but a number of associations correspond to ecological associations such as mating or parasitism, etc. Some insect inclusions can provide indirect evidences of specific habitats, or climate. The information provided by well-preserved inclusions corroborates the data from palynology: the palaeoenvironment of the resin deposits was a tropical dense, warm, and humid forest with a very complex fluvial system (lots of channels), altogether close to the sea. In addition, most of the fauna entombed in the Lebanese amber is the one living on the lower to mid parts of trees. This could be explained by the fact that this type of fauna has more chance to be trapped, since normally all the resin drops falling down from the tree pass inevitably and more frequently through these zones. The study of the different inclusions has allowed the reconstruction of the palaeoenvironment.
To date, 247 fossil insect species (mainly in amber) in 191 genera and 113 families have been described from Lebanon (see Appendix); others are still waiting identification. The discovery of outcrops yielding fossil insects preserved as compression/impression (adpressions) in sedimentary layers (silts, dysodiles, cinerites) of the same early Barremian age, augments our knowledge of the palaeobiodiversity in amber, as it is known that resin preserves insects in a selective way. Several factors affect the selectivity of the amber inclusions, like size, attraction/repulsion of the odour of the resin itself, proximity of resin producing trees to the habitat of the inclusion, and ecological behaviour of the insect.
The discovery of fossil insects in the Albian dysodiles/cinerites and in marine Cenomanian lithographic limestones of Hjoula and Nammoura adds knowledge of the previously unknown entomological biodiversity of the North-East of the Arabo-African palaeocontinent.
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Figure 18: Simplified
stratigraphic section showing the different intervals where the fossiliferous
amber crops out in the Lower Cretaceous sandstone of Lebanon. |
With its 35 Cretaceous outcrops yielding fossil insects, either in amber or as rock impressions-compressions (adpressions), Lebanon has contributed significantly to the advance of palaeoentomology and in our understanding of entomological evolution and palaeobiodiversity. Lebanon is among the countries with the highest densities of outcrops with insects. Amber is a material that has fascinated people so much and forever will not cease doing it. It constitutes a wonderful 'natural time capsule' as termed by Ross (1998, 2010), and an original material that not only preserves superb biological inclusions in their pristine three-dimensional detail, but also aspects of their ways of life and ecology. Preservation of life forms in amber increases significantly our knowledge of palaeobiodiversity, the palaeoecology of their inclusions, and diverse aspects of the palaeoenvironment, and gives the amber its attribute of exceptional 'window to the past' (Grimaldi, 2003a).
Lebanese amber contains a lot of extinct insect families (some of them are known only from Lebanon) and the records of the oldest representatives of many modern families of terrestrial arthropods. Lebanese inclusions constitute most of the time including "missing links" between the ancient Jurassic fauna and the modern one. The study of the Lebanese amber inclusions of insects is to date the only one that gives a clue to determining North-East Gondwanan biodiversity and environment in the extremely significant Early Cretaceous Subperiod. The recent discoveries of new and very diverse outcrops of fossiliferous amber in Lebanon help to meet the challenge of considerably enriching our knowledge of the Past. Efforts are under way to categorize this natural treasure from the Lower Cretaceous in the list of Heritage of Humanity. The different Lebanese outcrops are not yet officially protected against vandalism. Their destruction or pillaging would be a great loss to Human Heritage and to scientific knowledge.
We thank Professors André A. Nel (MNHN, Paris, France), Xavier Delclòs (University of Barcelona, Spain) and Jacek Szwedo (University of Gdansk, Poland) for their valuable comments on an earlier version of this work. We thank Prof. Ed Jarzembowski for his linguistic help. We thank Prof. Jacek Szwedo for providing illustration of the Hadath El-Joubbeh amber outcrop. This paper is a contribution to the activity of the Laboratory entitled "Advanced Micropalaeontology, Biodiversity and Evolutionary Research" (AMBER) led by DA at the Lebanese University. DA wants to thank the Chinese Academy of Sciences for financial support under the President's International Fellowship Initiative (PIFI).
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Whalley P.E.S. (1978).- New taxa of fossil and recent Micropterigidae with a discussion of their evolution and a comment on the evolution of Lepidoptera (Insecta).- Annals of the Transvaal Museum, Pretoria, vol. 31, p. 71-86.
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Zumoffen G. (1926).- Géologie du Liban. Carte géologique au 200.000ième.- Barrière, Paris. 165 p.
Plate
1:
Fossil insects in amber. A- Diptera: Dolichopodidae: Microphorites
sp., female, specimen number MCH-1B (Mechmech, Ain El-Khyar). B- Blattaria:
Caloblattinidae: Rhipidoblatta ?, specimen number MCH-1D (Mechmech, Ain
El-Khyar). C- Diptera: Dolichopodidae: female, Microphorites
sp., male, specimen number MCH-1A (Mechmech, Ain El-Khyar). D- Hymenoptera:
Platygastroidea: Scelionidae, specimen number MCH-1C (Mechmech, Ain El-Khyar).
E- Hemiptera: Sternorrhyncha: Coccoidea: Monophlebidae, female,
specimen number DAB-13 (Nimrin, El-Dabsheh). F- Neuroptera:
Rhachiberothidae: Raptorapax terribilissima, specimen number NBS-1A (Brissa).
G- Hymenoptera, specimen number BKK-1A (Beqaa Kafra). H-
Hemiptera: Enicocephalidae: Enicocephalinus acragrimaldi, male,
specimen number HDJ-1A (Hadath El Joubbeh). I- Diptera: Psychodoidea: Xenopsychoda
harbi, female, holotype, specimen number T-1 (Tannourine). J-
Diptera, Ceratopogonidae, female specimen number MKD-1 (Mazraat Kfardibiane).
Scale bars in F = 1 mm, in all remaining figures = 0.5 mm. |
|
Plate
2:
Fossil insects in amber. A- Diptera: Ceratopogonidae, female, parasited by an
Acari, specimen number OTJ-1BC (Ouata El-Jaouz). B- Ephemeroptera wing, specimen
number OTJ-1A (Ouata El-Jaouz). C- Diptera: Chironomidae, male (Bqaatouta, El
Shqif). D- Diptera: Ceratopogonidae, female (Bqaatouta, El Shqif). E-
Diptera: Chironomidae, female (Bqaatouta, El Shqif). F- Hymenoptera (Baskinta, Qanat
Bakish). G- Several inclusions including a Trichoptera, an Ephemeroptera, two
Diptera: Ceratopogonidae, a Diptera (head): Brachycera (Baskinta, Qanat Bakish).
H-
Hemiptera: Progonocimicidae: Ilahulgabalus endaidus, male,
holotype, specimen number DAY–1C (Daychouniyyeh). I- Hymenoptera, specimen
number K (Kfar Selouan). J- Six Diptera: Brachycera, specimen number J- K23 A-M
(Kfar Selouan). K- Diptera: Chironomidae: Libanopelopia cretacica,
female (Kfar Selouan, Khallet Douaiq). L- Hemiptera: Sternorrhyncha: Coccoidea:
Steingeliidae: Palaeosteingelia sp., male (Kfar Selouan, Khallet Douaiq).
M-
Thysanoptera: Tubulifera (Kfar Selouan, Khallet Douaiq). Scale bars
in G-H = 1 mm, in all remaining figures = 0.5 mm. |
|
Plate
3:
Fossil insects in amber. A- Isoptera: Lebanotermes veltzae, holotype,
specimen number 341C (Mdeyrij-Hammana). B- Hemiptera: Enicocephalidae: Enicocephalinus
acragrimaldi, male, paratype, specimen number 14B (Mdeyrij-Hammana). C-
Psocodea: Sphaeropsocidae: Asphaeropsocites neli, female, holotype,
specimen number 1513 (Mdeyrij-Hammana). D- Dermaptera: Rhadinolabis
phoenicica, female, holotype, specimen number 1013 (Mdeyrij-Hammana). E-
Hymenoptera: Scolebythidae: Uliobythus terpsichore, holotype, specimen
number 157A (Mdeyrij-Hammana). F- Psocodea: Pachytrocidae: Libaneuphoris
jantopi, holotype, specimen number FAL-11A (Falougha). G- Diptera:
Ceratopogonidae, female, trapped on a cobweb, specimen number AZH-1ABC (Ain
Zhalta). H- Diptera: Psychodidae: Protopsychoda leoi, female, holotype,
specimen number AD-65 (Ain Dara). I- Neuroptera: Chrysopoidea: Tragichrysa
ovoruptora, neonate larvae and associated egg remains, specimen number
S-7A-F (Sarhmoul). J- Diptera: Chironomidae: Libanopelopia cretacica,
male, holotype, specimen number HAR-2 (Roum - Aazour - Homsiyeh). K-
Diptera: Tanyderidae: Nannotanyderus ansorgei, male, holotype, specimen number
JG.
385/2B (Bkassine, Jouar Es-Souss). L- Blattaria, larva (Wadi Jezzine). M-
Hymenoptera: Maimetshidae: Ahiromaimetsha najlae, female, holotype,
specimen number MKN-1A (Maknouniyeh). N- Diptera: Psychodidae, male, specimen
number RIH-1C (Maknouniyeh). O- Diptera: Psychodidae: Palopsychoda jacquelinea,
male, specimen number C-5C (Esh-Sheaybeh). P- Diptera: Nematocera, male,
specimen number TAR-1B (Bouarij). Scale bars in A and M = 2 mm, in all remaining
figures = 0.5 mm. |
|
Plate
4:
Fossil insects preserved as compressions/impressions from Lebanon. A-
Blattaria wing, Mesoblattina libanensis, holotype, specimen number
INS-26367/1 (Qahmez). B- Ephemeroptera: Libanoephemera inopinatabranchia,
holotype, specimen number INS-63124/1 (Jdeidet Bkassine). C- Coleoptera: specimen number INS-63124/2 (Jdeidet Bkassine). D- Diptera,
specimen number INS-63124/4 (Jdeidet Bkassine). E- Coleoptera:
Staphylinoidea, specimen number INS-63124/3 (Jdeidet Bkassine). F-
Mantodea (Qnat). G- Blattaria: Mesoblattinidae: Mieroblattina pacis,
female, holotype, specimen number NI-5B (Nammoura). H- Orthoptera:
Chresmodidae: Chresmoda libanica, female, holotype, specimen number
NI-3A (Nammoura). I- Odonata: Anisoptera: Liupanshaniidae: Libanoliupanshania mimi,
holotype, specimen number F63 (Hjoula). J- Odonata: Anisoptera:
Libanocorduliidae: Libanocordulia debiei, holotype, specimen number F64
(Hjoula). K- Odonata: Anisoptera: Libanocorduliidae: Libanocordulia debiei,
paratype, specimen number F65 (Hjoula). L- Odonata: Anisoptera:
Libanocorduliidae: Libanocordulia debiei
(Hjoula). M-N- Odonata (Hjoula). O- Coleoptera: Scarabaeoidea (Hjoula).
P- Hemiptera: Cicadellidae (Hjoula). Q- Coleoptera (Sniyya).
Scale bars = 5 mm in A, F-H and Q, 3 mm in B-C and P, 1 mm in D-E, 1 cm
in I-N, 2 cm in O. |
|
List of fossil insects from Lebanon. Fossil insects preserved as impression-compression in rocks are preceded by an asterisk in the table below; all remaining taxa are preserved in amber.
| Class | Order | Family | Taxa | Site name |
| Insecta | Archaeognatha | Meinertellidae | Cretaceomachilis libanensis Sturm & Poinar, 1998 | Jouar Es-Souss |
| Glaesimeinertellus intermedius Sánchez-García et al., 2019 | Mdeyrij-Hammana | |||
| Macropsontus azari Sánchez-García et al., 2019 | Rihane | |||
| Macropsontus bachae Sánchez-García et al., 2019 | Mdeyrij-Hammana | |||
| Blattodea | Liberiblattinidae | Cryptoblatta aquatica Sendi & D. Azar in Vršanský et al., 2019 | Mdeyrij-Hammana | |
| Pseudomantina occisor Sendi in Vršanský et al., 2021 | Mdeyrij-Hammana | |||
| Blattellidae | Ocelloblattula ponomarenkoi Anisyutin & Gorochov, 2008 | Mdeyrij-Hammana | ||
| * ?Ocelloblattula striatus Káčerová & D. Azar, in press | Jdeidet Bkassine | |||
| Blattidae | Anenev asrev Vršanský et al., 2019 | Bloudane | ||
| Balatronis libanensis Sendi & D. Azar, 2017 | Ain Dara | |||
| Mesoblattinidae | * Mieroblattina pacis Vršanský & Makhoul, 2013 | Nammoura | ||
| * Mesoblattina libanensis Káčerová & D. Azar, in press | Qahmez | |||
| Nymphoblatta azari Vršanský & Grimaldi, 2004 | Bcharreh Mountains | |||
| Umenocoleidae | Cratovitisma cortexi Sendi in Podstrelená & Sendi, 2018 | Mdeyrij-Hammana | ||
| Pseudojantaropterix lebani (Vršanský & Grimaldi, 2003) | Jouar Es-Souss | |||
| Coleoptera | Anthicidae | Camelomorphasa longicervix Kirejtshuk & D. Azar, 2008 | Mdeyrij-Hammana | |
| Cerophytidae | Lebanophytum excellens Kirejtshuk & D. Azar, 2008 | Mdeyrij-Hammana | ||
| Chelonariidae | Eochelonarium belle Kirejtshuk & D. Azar, 2013 | Kfar Selouane | ||
| Clambidae | Eoclambus rugidorsum Kirejtshuk & D. Azar, 2008 | Mdeyrij-Hammana | ||
| Curculionidae | Cylindrobrotus pectinatus Kirejtshuk et al., 2009 | Mdeyrij-Hammana | ||
| Dermestidae | Cretonodes antounazari Kirejtshuk & D. Azar, 2009 | Mdeyrij-Hammana | ||
| Elodophthalmidae | Elodophthalmus gracilis Kirejtshuk & D. Azar, 2008 | Mdeyrij-Hammana | ||
| Elodophthalmus harmonicus Kirejtshuk & D. Azar, 2008 | Mdeyrij-Hammana | |||
| Hybosoridae | Libanochrus calvus Kirejtshuk et al., 2011 | Bouarij | ||
| Kateretidae | Lebanoretes andelmani Kirejtshuk & D. Azar, 2008 | Mdeyrij-Hammana | ||
| Latridiidae | Archelatrius marinae Kirejtshuk & D. Azar, 2009 | Mdeyrij-Hammana | ||
| Atetrameropsis subglobosa Kirejtshuk, 2013 | Mdeyrij-Hammana | |||
| Tetrameropsis mesozoica Kirejtshuk & D. Azar, 2008 | Mdeyrij-Hammana | |||
| Lebanophytidae | Lebanophytum excellens Kirejtshuk & D. Azar, 2008 | Mdeyrij-Hammana | ||
| Micromalthidae | Cretomalthus acracrowsonorum Kirejtshuk & D. Azar, 2008 | Jouar Es-Souss | ||
| Monotomidae | Rhizophtoma elateroides Kirejtshuk & D. Azar, 2009 | Mdeyrij-Hammana | ||
| Rhizophtoma synchrotronica Kirejtshuk & D. Azar, 2013 | Mdeyrij-Hammana | |||
| Rhizobactron marinae Kirejtshuk & D. Azar, 2013 | Nabaa Es-Sukkar | |||
| Nemonychidae | Libanorhinus succinus Kuschel & Poinar, 1993 | Jouar Es-Souss | ||
| Oropsis marinae Legalov & Kirejtshuk, 2017 | Bouarij | |||
| Ptismidae | Ptisma zasukhae Kirejtshuk & D. Azar, 2016a, 2016b | Nabaa Es-Sukkar | ||
| Throcidae | Potergosoma gratiosa Kovalev & Kirejtshuk, 2013 | Mdeyrij-Hammana | ||
| Rhomboaspis laticollis Kovalev & Kirejtshuk, 2013 | Bouarij | |||
| Sphindidae | Libanopsis impexa Kirejtshuk, 2015 | Mdeyrij-Hammana | ||
| Libanopsis limosa Kirejtshuk, 2015 | Jouar Es-Souss | |||
| Libanopsis poinari Kirejtshuk, 2015 | Mdeyrij-Hammana | |||
| Libanopsis slipinskii Kirejtshuk, 2015 | Nabaa Es-Sukkar | |||
| Libanopsis straminea Kirejtshuk, 2015 | Bouarij | |||
| Staphylinidae | Libanoeuaesthetus pentatarsus Lefèbvre et al., 2005 | Mdeyrij-Hammana | ||
| Dermaptera | incertae sedis | Rhadinolabis phoenicica Engel et al., 2011a | Mdeyrij-Hammana | |
| Diptera | incertae sedis | Xenopsychoda harbi D. Azar & Ziadé, 2005 | Tannourine | |
| Archizelmeridae | Zelmiarcha lebanensis Grimaldi et al., 2003 | Mdeyrij-Hammana | ||
| Chaoboridae | Libanoborus lukashevici D. Azar et al., 2009 | Mdeyrij-Hammana | ||
| Chimeromyiidae | Chimeromyia acuta Grimaldi & Cumming, 1999 | Bcharreh Mountains | ||
| Chimeromyia intriguea Grimaldi & Cumming, 1999 | Bcharreh Mountains | |||
| Chimeromyia mediobscura Grimaldi & Cumming, 2009 | Mdeyrij-Hammana | |||
| Chimeromyia pilitibia Grimaldi & Cumming, 2009 | Mdeyrij-Hammana | |||
| Chimeromyia reducta Grimaldi & Cumming, 1999 | Jouar Es-Souss | |||
| Chironomidae | Cretadiamesa arieli Veltz et al., 2007 | Mdeyrij-Hammana | ||
| Cretaenne kobeyssii D. Azar et al., 2008 | Mdeyrij-Hammana | |||
| Cretaenne inexpectata D. Azar et al., 2008 | Mdeyrij-Hammana | |||
| Cretapelopia salomea Veltz et al., 2007 | Mdeyrij-Hammana | |||
| Lebanodiamesa deploegi Veltz et al., 2007 | Mdeyrij-Hammana | |||
| Lebanorthocladius furcatus Veltz et al., 2007 | Mdeyrij-Hammana | |||
| Libanochlites neocomicus Brundin, 1976 | Jouar Es-Souss | |||
| Libanopelopia cretacica Veltz et al., 2007 | Roum-Aazour-Homsiyeh | |||
| Haematotanypus libanicus D. Azar et al., 2008 | Jouar Es-Souss | |||
| Paicheleria magnifica D. Azar & A. Nel, 2010b | Mdeyrij-Hammana | |||
| Wadelius libanicus Veltz et al., 2007 | Mdeyrij-Hammana | |||
| Ziadeus kamili D. Azar & A. Nel, 2010b | Mdeyrij-Hammana | |||
| Ceccidomyiidae | Cretadicerura salimi D. Azar & A. Nel, 2020 | Bouarij | ||
| Libanoclinorrhytis jaschhofi D. Azar & A. Nel, 2020 | Mdeyrij-Hammana | |||
| Libanohilversidia doryi D. Azar & A. Nel, 2020 | Tannourine | |||
| Libanowinnertzia perrichoti D. Azar & A. Nel, 2020 | Mdeyrij-Hammana | |||
| Ceratopogonidae | Archiaustroconops annae Choufani et al., 2014 | Mdeyrij-Hammana | ||
| Archiaustroconops bocaparvus Borkent, 2000 | Jouar Es-Souss | |||
| Archiaustroconops ceratoformis Szadziewski, 1996 | Jouar Es-Souss | |||
| Archiaustroconops cretaceous (Szadziewski, 1996) | Jouar Es-Souss | |||
| Archiaustroconops dominiakae Choufani et al., 2014 | Mdeyrij-Hammana | |||
| Archiaustroconops hammanaensis Choufani et al., 2014 | Mdeyrij-Hammana | |||
| Archiaustroconops hamus Borkent, 2000 | Jouar Es-Souss | |||
| Archiaustroconops szadziewskii Borkent, 2000 | Bcharreh Mountains | |||
| Austroconops fossilis Szadziewski, 1996 | Jouar Es-Souss | |||
| Austroconops gladius Borkent, 2000 | Jouar Es-Souss | |||
| Austroconops gondwanicus Szadziewski, 1996 | Jouar Es-Souss | |||
| Austroconops megaspinus Borkent, 2000 | Jouar Es-Souss | |||
| Fossileptoconops lebanicus Szadziewski, 1996 | Jouar Es-Souss | |||
| Lebanoculicoides bloudani Choufani et al., 2015 | Bloudan | |||
| Lebanoculicoides daheri Choufani et al., 2014 | Mdeyrij-Hammana | |||
| Lebanoculicoides mesozoicus Szadziewski, 1996 | Jouar Es-Souss | |||
| Leptoconops amplificatus Borkent, 2001 | Bcharreh Mountains | |||
| Leptoconops antiquus Borkent, 2001 | Bcharreh Mountains | |||
| Minyohelea bacula Borkent, 2001 | Jouar Es-Souss | |||
| Minyohelea falcate Borkent, 2001 | Bcharreh Mountains | |||
| Minyohelea lebanica (Szadziewski, 1996) | Jouar Es-Souss | |||
| Minyohelea minuta (Szadziewski, 1996) | Jouar Es-Souss | |||
| Minyohelea nexuosa Pielowska-Ceranowska et al., 2022 | Mdeyrij-Hammana | |||
| Minyohelea schleei Szadziewski, 1996 | Jouar Es-Souss | |||
| Minyohelea wirthi (Szadziewski, 1996) | Jouar Es-Souss | |||
| Protoculicoides acraorum Borkent, 2001 | Jouar Es-Souss | |||
| Protoculicoides krzeminskii Choufani et al., 2014 | Mdeyrij-Hammana | |||
| Protoculicoides punctus Borkent, 2001 | Bcharreh Mountains | |||
| Protoculicoides schleei (Szadziewski, 1996) | Jouar Es-Souss | |||
| Protoculicoides succineus Szadziewski, 1996 | Jouar Es-Souss | |||
| Protoculicoides unus Borkent, 2001 | Jouar Es-Souss | |||
| Corethrellidae | Corethrella cretacea Szadziewski, 1995 | Jouar Es-Souss | ||
| Dolichopodidae | Microphorites extinctus Hennig, 1971 | Jouar Es-Souss | ||
| Microphorites oculeus Grimaldi & Cumming, 1999 | Jouar Es-Souss | |||
| Microphorites similis Grimaldi & Cumming, 1999 | Jouar Es-Souss | |||
| Sympycnites primaevus Grimaldi & Cumming, 1999 | Bcharreh Mountains | |||
| Empididae | Atelestites senectus Grimaldi & Cumming, 1999 | Bcharreh Mountains | ||
| Avenaphora hispida Grimaldi & Cumming, 1999 | Bcharreh Mountains | |||
| Phaetempis lebanensis Grimaldi & Cumming, 1999 | Bcharreh Mountains | |||
| Trichinites cretaceous Hennig, 1970 | Jouar Es-Souss | |||
| Hilarimorphidae | Cretahilarimorpha lebanensis Myskowiak et al., 2016 | Mdeyrij-Hammana | ||
| Ironomyiidae | Lebambromyia acrai Grimaldi & Cumming, 1999 | Jouar Es-Souss | ||
| Limoniidae | Lebania levantia Podenas & Poinar, 2001 | Jouar Es-Souss | ||
| Lebania longaeva Podenas & Poinar, 2001 | Jouar Es-Souss | |||
| Helius lebanensis Kania et al., 2013 | Bouarij | |||
| Helius ewa Krzemiński et al., 2014 | Mdeyrij-Hammana | |||
| Lonchopteroidea incertae sedis family | Alonchoptera lebanica Grimaldi, 2018 | Bcharreh Mountains | ||
| Lonchopteridae | Lonchopterites prisca Grimaldi & Cumming, 1999 | Bcharreh Mountains | ||
| Lonchopteromorpha asetocella Grimaldi & Cumming, 1999 | Bcharreh Mountains | |||
| Lygistorrhinidae | Lebanognoriste prima Blagoderov & Grimaldi, 2004 | Jouar Es-Souss | ||
| Platypezidae | Lebanopeza azari Grimaldi, 2018 | Mdeyrij-Hammana | ||
| Psychodidae | Cretapsychoda inexpectata D. Azar et al., 1999 | Mdeyrij-Hammana | ||
| Eophlebotomus gezei D. Azar et al., 2003 | Mdeyrij-Hammana | |||
| Libanophlebotomites ramyii D. Azar et al., 2022a | Qanat Bakish (Baskinta) | |||
| Libanophlebotomus lutfallahi D. Azar et al., 1999 | Mdeyrij-Hammana | |||
| Libanopsychoda abillamai D. Azar et al., 1999 | Mdeyrij-Hammana | |||
| Libanosycorax dimyi D. Azar et al., 2018 | Mdeyrij-Hammana | |||
| Mesophlebotomites hennigi D. Azar et al., 1999 | Mdeyrij-Hammana | |||
| Paleopsychoda inexpectata D. Azar & A. Nel, 2002 | Mdeyrij-Hammana | |||
| Paleopsychoda jacquelinae D. Azar et al., 1999 | Mdeyrij-Hammana | |||
| Paleopsychoda jarzembowskii D. Azar & Maksoud, 2022 | Mdeyrij-Hammana | |||
| Paleopsychoda solignaci D. Azar et al., 1999 | Mdeyrij-Hammana | |||
| Paralibanopsychoda agnieszkae D. Azar & A. Nel, 2002 | Mdeyrij-Hammana | |||
| Phlebotomites brevifilis Hennig, 1972 | Jouar Es-Souss | |||
| Phlebotomites longifilis Hennig, 1972 | Jouar Es-Souss | |||
| Protopsychoda hammanaensis D. Azar et al., 1999 | Mdeyrij-Hammana | |||
| Protopsychoda leoi D. Azar & Maksoud, 2020 | Ain Dara | |||
| Protopsychoda nadiae D. Azar et al., 1999 | Mdeyrij-Hammana | |||
| Ptychopteridae | Leptychoptera dimkina Lukashevich & D. Azar, 2003 | Mdeyrij-Hammana | ||
| Leptychoptera vovkina Lukashevich & D. Azar, 2003 | Bcharreh Mountains | |||
| Rhagionidae | Lebanoleptis huangi Angelini et al., 2016 | Bouarij | ||
| Mesobolbomyia acari Grimaldi & Cumming, 1999 | Jouar Es-Souss | |||
| Paleochrysopilus hirsutus Grimaldi & Cumming, 1999 | Bcharreh Mountains | |||
| Sciadoceridae | Archisciada lebanensis Grimaldi & Cumming, 1999 | Bcharreh Mountains | ||
| Tanyderidae | Nannotanyderus ansorgi Krzemiński et al., 2013 | Jouar Es-Souss | ||
| Trichoceridae | Ewaurista pusilla Shcherbakov & D. Azar, 2019 | Mdeyrij-Hammana | ||
| Xylomyidae | Cretoxyla azari Grimaldi & Cumming, 2011 | Mdeyrij-Hammana | ||
| Ephemeroptera | incertae sedis | * Libanoephemera inopinatabranchia D. Azar et al., 2019a | Jdeidet Bkassine | |
| Leptophlebiidae | Conovirilus poinari McCafferti, 1997 | Jouar Es-Souss | ||
| Hemiptera | incertae sedis | Xiphos vani Vea & Grimaldi, 2015 | Mdeyrij-Hammana | |
| Aleyrodidae | Aretsaya therina Drohojowska & Szwedo, 2015 | Ain Dara | ||
| Baetylus kahramanus Drohojowska & Szwedo, 2011 | Mdeyrij-Hammana | |||
| Bernaea neocomica Schlee, 1970 | Jouar Es-Souss | |||
| Gapenus rhinariatus Drohojowska & Szwedo, 2013 | Rihane | |||
| Heidea cretacica Schlee, 1970 | Jouar Es-Souss | |||
| Milqartis azari Drohojowska & Szwedo, 2015 | Mdeyrij-Hammana | |||
| Shapashe aithiopa Drohojowska & Szwedo, 2015 | Mdeyrij-Hammana | |||
| Yamis libanotos Drohojowska & Szwedo, 2015 | Mdeyrij-Hammana | |||
| Cixiidae | Karebodopoides aptianus (Fennah, 1987) | Jouar Es-Souss | ||
| Apticoccidae | Apticoccus minutus Koteja & D. Azar, 2008 | Mdeyrij-Hammana | ||
| Apticoccus fortis Vea & Grimaldi, 2015 | Mdeyrij-Hammana | |||
| Apticoccus longitenuis Vea & Grimaldi, 2015 | Ain Dara | |||
| Enicocephalidae | Enicocephalinus acragrimaldii D. Azar et al., 1999 | Mdeyrij-Hammana | ||
| Hammanococcidae | Hammanococcus setosus Koteja & D. Azar, 2008 | Mdeyrij-Hammana | ||
| Hodgsonicoccidae | Hodsonicoccus patefactus Vea & Grimaldi, 2015 | Bcharreh Mountains | ||
| Lebanococcidae | Lebanococcus longiventris Koteja & D. Azar, 2008 | Mdeyrij-Hammana | ||
| Liadopsyllidae | Liadopsylla apedetica Ouvrard et al., 2010 | Mdeyrij-Hammana | ||
| Neazoniidae | Neazonia immature Szwedo, 2007 | Mdeyrij-Hammana | ||
| Neazonia imprinta Szwedo, 2007 | Jouar Es-Souss | |||
| Neazonia tripleta Szwedo, 2007 | Mdeyrij-Hammana | |||
| Ortheziidae | Cretorhezia hammanaica Koteja & D. Azar, 2008 | Mdeyrij-Hammana | ||
| Perforissidae | Aafrita biladalshama Szwedo & D. Azar, 2013 | Mdeyrij-Hammana | ||
| Progonocimicidae | Ilahulgabalus endaidus Szwedo et al., 2011 | El-Dayshouniyyeh | ||
| Protopsyllidae | Talaya batraba Drohojowska et al., 2013 | Mdeyrij-Hammana | ||
| Pseudococcidae | Williamsicoccus megalops Vea & Grimaldi, 2015 | Mdeyrij-Hammana | ||
| Putoidae | Palaeotupo danieleae Koteja & D. Azar, 2008 | Mdeyrij-Hammana | ||
| Pennygullaniidae | Pennygullania electrina Koteja & D. Azar, 2008 | Mdeyrij-Hammana | ||
| Schizopteridae | Libanohypselosoma popovi D. Azar & A. Nel, 2010a | Mdeyrij-Hammana | ||
| Steingeliidae | Palaeosteingelia acrai Koteja & D. Azar, 2008 | Mdeyrij-Hammana | ||
| Palaeosteingelia caudate Koteja & D. Azar, 2008 | Mdeyrij-Hammana | |||
| Tajmyraphididae | Lebanaphis minor Heie, 2000 | Mdeyrij-Hammana | ||
| Megarostrum azari Heie, 2000 | Mdeyrij-Hammana | |||
| Thelaxidae | Gondvanoaphis estephani Węgierek & Grimaldi, 2010 | Bcharreh Mountains | ||
| Yuripopovinidae | Yuripopovina magnifica D. Azar et al., 2011b | Bouarij | ||
| Hymenoptera | Archaeoserphitidae | Archaeoserphites melqarti Engel, 2015 | Bcharreh Mountains | |
| Bethylidae | Lancepyris opertus Azevedo & D. Azar, 2012 | Mdeyrij-Hammana | ||
| Dryinidae | Archaeodryinus palaeophoenicius (Olmi, 2000) | Jouar Es-Souss | ||
| Evaniidae | Eovernevania cyrtocerca Deans, 2004 | Mdeyrij-Hammana | ||
| Lebanevania azari Basibuyuk & Rasnitsyn, 2002 | Jouar Es-Souss | |||
| Protoparevania lourothi Deans, 2004 | Mdeyrij-Hammana | |||
| Gallorommatidae | Cretaceomma libanensis Rasnitsyn & D. Azar in Rasnitsyn et al., 2022 | Mdeyrij-Hammana | ||
| Maimetshidae | Ahiromaimetsha najlae Perrichot et al., 2011 | Maknouniyyeh | ||
| Scelionidae | Cretaxenomerus jankotejai A. Nel & D. Azar, 2005 | Mdeyrij-Hammana | ||
| Proteroscelio gravatus Johnson et al., 2008 | Mdeyrij-Hammana | |||
| Sclerogibbidae | Sclerogibbodes embioleia Engel & Grimaldi, 2006 | Bcharreh Mountains | ||
| Scolebythidae | Libanobythus milkii Prentice & Poinar in Prentice et al., 1996 | Jouar Es-Souss | ||
| Uliobythus terpsichore Engel & Grimaldi, 2007 | Mdeyrij-Hammana | |||
| Zapenesia libanica Engel & Grimaldi, 2007 | Mdeyrij-Hammana | |||
| Serphitidae | Leptoserphites iriae Rasnitsyn & D. Azar in Rasnitsyn et al., 2022 | Qanat Bakish | ||
| Leptoserphites pabloi Rasnitsyn & D. Azar in Rasnitsyn et al., 2022 | Qanat Bakish | |||
| Microserphites libanensis Rasnitsyn & D. Azar in Rasnitsyn et al., 2022 | Mdeyrij-Hammana | |||
| Spathiopteridae | Mymaropsis baabdaensis Krogmann et al., 2016 | Mdeyrij-Hammana | ||
| Stigmaphronidae | Libanophron astarte Engel & Grimaldi, 2009 | Mdeyrij-Hammana | ||
| Isoptera | incertae sedis | Lebanotermes veltzae Engel et al., 2011a | Mdeyrij-Hammana | |
| Hodotermitidae | Melquartitermes myrrheus Engel et al., 2007 | Bcharreh Mountains | ||
| Lepidoptera | Micropterigidae | Parasabatinca aftimacrai Whalley, 1978 | Jouar Es-Souss | |
| Manthodea | Gryllomantidae | Gryllomantis lebanensis (Grimaldi, 2003b) | Bcharreh Mountains | |
| Neuroptera | incertae sedis Chrysopidea | Tragichrysa ovoruptora Pérez de la Fuente et al., 2018a | Sarhmoul | |
| Tyruschrysa melqart Pérez de la Fuente et al., 2018b | Bouarij | |||
| Berothidae | Banoberotha enigmatica Whalley, 1980 | Jouar Es-Souss | ||
| Sibelliberotha rihanensis D. Azar & A. Nel, 2013 | Rihane | |||
| Coniopterygidae | Libanoconis fadiacra (Whalley, 1980) | Jouar Es-Souss | ||
| Libanosemidalis hammanaensis D. Azar et al., 2000 | Mdeyrij-Hammana | |||
| Rhachiberothidae | Chimerhachiberotha acrasarii A. Nel et al., 2005 | Jouar Es-Souss | ||
| Paraberotha acra Whalley, 1980 | Jouar Es-Souss | |||
| Raptorapax terribilissima Petrolevičius et al., 2010 | Bouarij | |||
| Spinoberotha mickaelacrai A. Nel et al., 2005 | Mdeyrij-Hammana | |||
| Saucrosmylidae | * Lebanosmylus leae D. Azar & A. Nel, 2022 | Hjoula | ||
| Odonata | incertae sedis | Libanolestes flecki D. Azar et al., 2010c | Ain Dara | |
| Libanocorduliidae | * Libanocordulia debiei D. Azar et al., 2019b | Hjoula | ||
| Liupanshaniidae | * Libanoliupanshania mimi D. Azar et al., 2019b | Hjoula | ||
| Orthoptera | Chresmodidae | * Chresmoda libanica A. Nel et al., 2004 | Nammoura | |
| Haglotettigoniidae | ?Halotettigonia aenigmatosa Gorokhov, 2010 | Mdeyrij-Hammana | ||
| Rhaphidophoridae | Aenigmaraphidophora mouniri D. Azar et al., 2022b | Bqaatouta | ||
| Psocodea | ? Amphientomidae | Libanomphientomum nudus Choufani et al., 2011 | Mdeyrij-Hammana | |
| Paramesopsocidae | Paramesopsocus lu D. Azar et al., 2008 | Mdeyrij-Hammana | ||
| Prionoglarididae / Archaeatropidae | Bcharrehglaris amunobi D. Azar & A. Nel, 2004 | Bcharreh Mountains | ||
| Libanoglaris chehabi D. Azar & A. Nel, 2004 | Mdeyrij-Hammana | |||
| Libanoglaris mouawadi D. Azar et al., 2003 | Mdeyrij-Hammana | |||
| Libanoglaris randatae D. Azar & A. Nel, 2004 | Jouar Es-Souss | |||
| Setoglaris reemae D. Azar & A. Nel, 2004 | Mdeyrij-Hammana | |||
| Prionoglarididae | Palaeosiamoglaris hammanaensis Hakim et al., 2022 | Mdeyrij-Hammana | ||
| Pachytrocidae | Libaneuphoris jantopi D. Azar et al., 2015 | Falougha | ||
| Psyllipsocidae | Libanopsyllipsocus alexanderasnitsyni D. Azar & A. Nel, 2011 | Mdeyrij-Hammana | ||
| Sphaeropsocidae | Sphaeropsocites lebanensis Grimaldi & Engel, 2006 | Jouar Es-Souss | ||
| Asphaeropsocites neli D. Azar et al., 2010a | Mdeyrij-Hammana | |||
| Raphidioptera | Mesoraphidiidae | Lebanoraphidia nana Bechly & Wolf-Schwenninger, 2011 | Jouar Es-Souss | |
| Thysanoptera | Adiheterothripidae | Neocomothrips hennigianus Strassen, 1973 | Jouar Es-Souss | |
| Progonothrips horridus Strassen, 1973 | Jouar Es-Souss | |||
| Rhetinothrips elegans Strassen, 1973 | Jouar Es-Souss | |||
| Scaphothrips antennatus Strassen, 1973 | Jouar Es-Souss | |||
| Exitelothrips mesozoicus Strassen, 1973 | Jouar Es-Souss | |||
| Jezzinothrips cretacicus Strassen, 1973 | Jouar Es-Souss | |||
| Moundithripidae | Moundthrips beatificus P. Nel et al., 2007 | Jouar Es-Souss | ||
| Phlaeothripidae | Rohrthrips libanicus P. Nel et al., 2010 | Mdeyrij-Hammana | ||
| Thripidae | Tethysthrips libanicus P. Nel et al., 2010 | Mdeyrij-Hammana | ||
| Trichoptera | Dipseudopsidae | Phylocentropus succinolibanensis Wichard & D. Azar, 2018 | Mdeyrij-Hammana | |
| Ecnomidae | Ecnomus cretacia Wichard & D. Azar, 2018 | Mdeyrij-Hammana |
