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Jordanian–Lebanese–Syrian cockroaches s.s. from Lower Cretaceous amber – Monograph

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Abstract

The earliest amber with numerous important paleobiological samples, and the only significant Lower Cretaceous one containing cockroaches, comes from the Lebanese regions Bouarij, Falougha, Hammana-Mdeirij, Jezzine, Mechmech, Qanat Bakish, Bqaatouta and Rihane (and 21 unstudied outcrops), and from Jordanian amber of Wadi Zerka and Syrian amber of Bloudane. Complex data (RTFI spectroscopy, palynology and entomofauna with diverse and abundant Nematocera) indicated that the amber originated in a subtropical, warm and humid gymnosperm-dominated forest in close proximity to the sea. 79 specimens of cockroaches s. s. (excluding Isoptera, Mantodea and possibly Chresmoda) were housed in 18 species (and three species incertae sedis), including Pravdupovediac neklam, P. maaloufi gen. et spp. n; Elisama globosa, Cratovitisma spinosa and Neoblattella nechapetomu spp. n. The former three belong to the dominant family Blattulidae Vishniakova, 1982 (n = 14), while Umenocoleidae Chen et Tan, 1973 were codominant (n = 10) in the present fossil record. Diversity and disparity of the preserved forms was moderately high and comparable to the Middle Jurassic and Lower Cretaceous sedimentary sites – i.e., higher than in Early Jurassic sites and lower than in Upper Jurassic Karabastau sedimentary rocks and Late Cretaceous North Myanmar amber. Scarce syninclusions in small amber pieces represent a coccid, psychodid, ceratopogonid, possibly a pathogenic fungus and plant trichomes. Palaeogeographically Lepidopterix Sendi, 2020; Cryptoblatta Sendi, 2019; Miniblattina Sendi, 2021; Pravdupovediac gen. n. and possibly Nymphoblatta Vršanský et Grimaldi, 2004 (Lebanese amber); Compunctiotypus Kaddumi, 2005 and Grandocularis Kaddumi, 2005 (Jordanian amber); and Anenev Vršanský, Oružinský, Sendi, Choufani, El-Halabi et Azar, 2019 (Syrian amber) are reported only in the herein studied amber, while the Jantaropterix/ Pseudojantaropterix-complex; Cratovitisma Bechly, 2007; Nymphoblatta Vršanský et Grimaldi, 2004 aff. Perlucipecta Wei et Ren, 2013 and aff. Sivis Vršanský, 2009; Pseudomantina Sendi et Vršanský, 2021; Elisama Giebel, 1856; Ocelloblattula Anisyutkin et Gorokhov, 2008, ?Rhipidoblatta Vishniakova, 1968; Balatronis Šmídová et Lei, 2017 and Neoblatella Shelford, 1911 (nowadays restricted to the Americas) were cosmopolitan with occurrences in both Laurasia and Gondwana. The low percentage (0.4% compared with 2.5–5% in North Myanmar amber) of cockroaches preserved in Lebanese amber could be due to lower viscosity of the amber-forming resin, and/or due to different paleoenvironments, suggested also for Taimyr amber with a moderate climate (constituting 0.3% cockroaches of all samples). Phylogenetically, several still living families, such as Ectobiidae Brunner von Wattenwyl, 1865 and Blattidae Latreille, 1810 have their oldest representatives within Lebanese amber, suggesting that the amber is advanced in evolutionary aspect among all Mesozoic ecosystems (all other families went extinct near Cretaceous–Paleogene boundary, K/Pg). Furthermore, Neoblattella is the earliest recorded still living genus. However, respective species from the present amber complex do not reveal any specific phylogenetical signal and weren’t measurably more basal when compared with their younger North Myanmar amber counterparts. A wide range of morphotypes with various ecological adaptations are documented, suggesting already established, sophisticated trophic relationships, similar as in today’s ecosystems. Nevertheless, given the abundance of blattulids and mesoblattids, most had probably a standard detritivorous feeding habit - the more advanced morphotypes known from Myanmar amber were absent. To summarise, the herein studied amber complex reveals origination of a moderately high diversity and disparity of forms (corresponding to the beginning of the Cretaceous Terrestrial Revolution), similar to Upper Jurassic assemblages, with a number of advanced forms, but lower than in burmite, which might be caused due to collection bias, shorter deposition time, milder tropical climate, or due to a different ecosystem (e.g. with less diversified angiosperms). The present amber reveals the earliest record of already triggered Mesozoic-to-Cenozoic ecosystem change within global forests.

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Data Availability

The data supporting the findings of this study are available within the article and in supplementary Online Resources.

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Acknowledgements

We thank Mounir Maalouf for collecting two samples and Júlia Káčerová for making detailed illustrations of two specimens (Figs. 26, 28, 32, and 34) and an artistic paleoenvironmental reconstruction (Fig. 1). We also thank two anonymous opponents for keen revisions. This work was supported by the Slovak Research and Development Agency under the contracts No. APVV-0436-12; UNESCO- Amba/ MVTS supporting grant of Presidium of the Slovak Academy of Sciences and VEGA 2/0113/22. Part of the National Amber Project of Slovakia.

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Author notes

  1. Hemen Sendi, Peter Vršanský, Dany Azar contributed equally to this work.

Authors and Affiliations

  1. Institute of Zoology, Slovak Academy of Sciences, Dúbravská Cesta 9, 845 06, Bratislava, Slovakia

    Hemen Sendi & Peter Vršanský

  2. Faculty of Natural Sciences, Illkovicova 6, Bratislava, 84104, Slovakia

    Hemen Sendi

  3. Earth Science Institute, Slovak Academy of Sciences, Dúbravská Cesta 9, P.O. BOX 106, 840 05, Bratislava, Slovakia

    Peter Vršanský

  4. Faculty of Science II, Fanar, Natural Sciences Department, Lebanese University, PO Box 26110217, Fanar - Matn, Lebanon

    Dany Azar

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  1. Hemen Sendi
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Contributions

H.S. and P.V. provided photodocumentation, illustrations, descriptions, phylogenetic network analysis and wrote the text; H.S. performed the main systematic research. P.V. (PI) designed research and classified specimens. DA collected specimens and provided geological and geographical information on the amber outcrops in Lebanon.

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Correspondence to Peter Vršanský.

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On behalf of all authors, the corresponding author states that there is no potential (financial, political, religious) conflict of interest. Multiple affiliation of P.V. is within one governmental academic body (Slovak Academy of Sciences).

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The monograph is registered in Zoobank under: urn:lsid:zoobank.org:pub:20C8FD8E-CE8E-4C8E-88F4-1A81BDEA2C5D, together with one new genus and five new species (see above).

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Sendi, H., Vršanský, P. & Azar, D. Jordanian–Lebanese–Syrian cockroaches s.s. from Lower Cretaceous amber – Monograph. Biologia 78, 1447–1541 (2023). https://doi.org/10.1007/s11756-023-01357-y

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