Zootaxa

Insect Evolution at the Dawn of the Cretaceous Terrestrial Revolution (Title page)

DI-YING HUANG — 2024-12-31

Insect Evolution at the Dawn of the Cretaceous Terrestrial Revolution (Table of contents)

DI-YING HUANG — 2024-12-31

Lower Cretaceous terrestrial outcrops with fossil insects from Lebanon and China

CHEN-YANG CAI — 2024-12-31

The first aquatic insect fossil from the Cenomanian Konservat-Lagerstätte of Haqel (Lebanon) fills a gap in the fossil record of giant water bugs (Heteroptera: Belostomatidae)

MATHIEU BODERAU — 2024-12-31

Libanobelostoma calineae gen. et sp. nov., the first mid-Cretaceous giant water bug, is characterized, illustrated and described from the fossil fish Konservat-Lagerstätte of Haqel, Lebanon. This new discovery is important for both the fossil record of the Belostomatidae and for increasing our understanding of the paleoenvironment of Haqel. The new genus displays a combination of synapomorphies and symplesiomorphies for Belostomatidae, along with putative autapomorphies justifying its taxonomic treatment. Together with Libanoaeshna mikhaeli Azar et al., 2024 they represent the first insect discoveries in the fossil fish Konservat-Lagerstätte of Haqel, making it an additional palaeoentomological site to the numerous other sites in Lebanon.

A puzzling Cretaceous psyllid-type forewing from the South of China

MARINA HAKIM — 2024-12-31

The new taxon Fujianella paradoxa gen. et sp. nov. is described and illustrated from the Early Cretaceous Bantou Formation, in Yong’an County, Fujian Province, southeastern China. It is placed in the insect suborder Sternorrhyncha (Hemiptera) but left unassigned to any family at present due to its controversial features. This fossil record represents the second psyllid-looking taxon reported from the Cretaceous of China, found mainly in the Jurassic. This discovery further showcases the similarities in entomofauna between the Middle Jurassic Yanliao biota in the North of China and the Early Cretaceous in the South of China.

**Redescription of the ‘libelluloid’ Mesocordulia boreala (Odonata: Mesocorduliidae) from the Lower Cretaceous of China**

ANDRÉ NEL — 2024-12-31

On the basis of a very well-preserved new specimen, we redescribe the Early Cretaceous genus and species Mesocordulia boreala, currently attributed to the Cretaceous subfamily Mesocorduliinae of Araripelibellulidae. We refine key characteristics of wing venation, which allow us to exclude this species from the Araripelibellulidae and elevate the Mesocorduliinae to the family rank as Mesocorduliidae. The relationships of this fossil are clarified. As the genus name Mesocordulia is preoccupied, we propose a replacement name, Guocordulia.

The first and oldest record of Issidae from the Lower Cretaceous of Lebanon (Hemiptera, Fulgoromorpha)

DANY AZAR — 2024-12-31

Libanissus bkassinensis Azar, Maksoud & Nel, gen. et sp. nov. is illustrated and described from the Lower Cretaceous dysodile (oil papershales) of Bkassine, South Lebanon, and its taxonomic position discussed. Libanissus bkassinensis Azar, Maksoud & Nel, gen. et sp. nov. is characterized by its banded legs and body, a hind wing with two lobes, RA and RP very short, M with only two very short apical branches, CuA with two branches, and a small but distinct apical furcation of PCu close to the incision of the wing margin. Libanissus bkassinensis Azar, Maksoud & Nel, gen. et sp. nov. represents the earliest record (lower Barremian) of the Issidae. Prior to this discovery, the oldest known record was from the Paleocene of France.

The oldest false fairy wasp (Hymenoptera: Mymarommatidae) from Lower Cretaceous Lebanese amber

SERGIO ÁLVAREZ-PARRA — 2024-12-31

Mymarommatidae (Hymenoptera), also known as false fairy wasps, are mainly characterised by their minute size, the peculiar wings, and the bi-segmented petiole. They form part of the superfamily Mymarommatoidea together with the Cretaceous families †Alavarommatidae, †Dipterommatidae, and †Gallorommatidae. The clade Bipetiolarida embraces Mymarommatoidea and †Serphitoidea, the two proctotrupomorphan superfamilies whose members have bi-segmented petiole. The biology of the mymarommatids is poorly known and only one extant species is known to be an egg endoparasitoid of barklice (Psocodea). Here, we describe a new species of mymarommatid wasp, Archaeromma phoenicium sp. nov., from Lower Cretaceous (Barremian) amber of Lebanon. We compare it with the other species of the genus and discuss the validity of some characters for generic identification. This new species is the oldest representative of the genus Archaeromma Yoshimoto, 1975 and of the family Mymarommatidae. A checklist and a key to the species of Archaeromma is included. Furthermore, we discuss the palaeobiogeography of the Cretaceous mymarommatoids. The description of the new species increases the list of insect co-occurrences between Lebanese and Spanish ambers to 21 genera.

**A new species of Manlaya (Hymenoptera: Baissidae) from the Lower Cretaceous Shouchang Formation, Eastern China**

SERGIO ÁLVAREZ-PARRA — 2024-12-31

Baissid wasps (Hymenoptera: Baissidae) represent a family within the superfamily Evanioidea composed of 36 species in seven genera restricted to a period spanning from the Berriasian (earliest Cretaceous) to the Turonian (early Late Cretaceous). The status and rank of the family have been changing over time, and recent phylogenetic analyses indicate that Baissidae may be paraphyletic to Aulacidae + Gasteruptiidae. The genus Manlaya Rasnitsyn, 1980 is the most diverse and widespread in the family, including representatives from western and eastern Laurasia. Here, we describe Manlaya minima sp. nov. from the Lower Cretaceous Shouchang Formation (Zhejiang Province, China). The new species has elbowed second medial cell, similar to Electrobaissa Engel, 2013, although the rest of the characters match with Manlaya, indicating that this character might be related to interspecific variability rather than differences between genera. The co-occurrence of Manlaya in the contemporaneous Shouchang Formation and the Jehol biota may point to a connection between the two entomofaunas. A key to the species of the family Baissidae from China is provided.

**A new species of Elodophthalmus from Lower Cretaceous Lebanese amber unravels the systematic placement of Elodophthalmidae and reconciling phylogenetic conflicts within Tenebrionoidea**

CHENYANG CAI — 2024-12-31

Elodophthalmidae, represented by the sole genus Elodophthalmus Kirejtshuk & Azar, is an extinct beetle family described from Lower Cretaceous Lebanese amber. Elodophthalmidae has been hypothesized to share close affinities with the extant superfamily Scirtoidea, but their exact systematic position remains elusive. Here, we describe a new species of Elodophthalmus, Elodophthalmus maksoudae sp. nov., based on a well-preserved fossil from Early Cretaceous amber collected in Hammana/Mdeyrij, Central Lebanon, the same locality as other congeneric species. Using improved imaging technologies, we reveal detailed morphological characters of the fossil, providing critical evidence for a more robust systematic placement of Elodophthalmus. Our findings present compelling evidence supporting the placement of Elodophthalmidae within the superfamily Tenebrionoidea. Furthermore, we highlight that conflicts in the interfamilial relationships within Tenebrionoidea across various datasets can, at least partially, be mitigated by adopting the site-heterogeneous CAT-GTR+G model. We suggest that the traditional reliance on site-homogeneous models in Sanger sequencing-based studies may fail to capture the complexities of nucleotide substitution patterns. With the increasing recognition of the importance of modeling among-site compositional heterogeneity, our study points to the necessity for more rigorous model testing in the phylogenetic studies of ancient lineages such as Tenebrionoidea.

Quantitative morphology of fossil adephagan beetle larvae including a first record from the Jehol biota does not indicate major diversity losses over time

JOACHIM T. HAUG — 2024-12-31

Coleoptera is a hyper-diverse group of animals with about 400,000 formally described species. Also the morphological diversity of beetles is very high, not only in the adults, but also in the larvae. To understand the evolutionary origin of this enormous diversity, investigations of fossils are crucial, but especially for the larvae such fossil are still scarce. In this study, we present 15 new fossil larvae from different deposits of Cretaceous to Miocene age, which we interpret as larvae of the beetle ingroup Adephaga. Most of these are three-dimensionally preserved and either embedded in amber or silicified. One specimen is a compression fossil, which is very rare for beetle larvae, and represents the first report of an adephagan beetle larva from the Early Cretaceous Jehol biota of China. Of all previously known and new fossil adephagan larvae and of selected extant ones the shapes of the head capsules and mandibles were compared with quantitative morphology methods (elliptic Fourier analysis and principal component analysis). The shapes of the fossil larvae lie all within the morphospace of those of the extant larvae, which indicates that the highest diversity in this aspect is present in the modern fauna, hence no diversity loss occurred. Other lineages of Holometabola show similar patterns, with indications of larval diversifications already in the Cretaceous, further specialisation afterwards, and no major losses. This pattern may be a reason for the enormous species richness of certain holometabloan groups in the modern fauna.

Cretaceous beetles from Jinju Formation: Trachypachidae (Coleoptera: Adephaga)

SOO BIN LEE — 2024-12-31

Here we describe two new species, Psacodromeus recta sp. nov. and Necronectulus lazarus sp. nov., that belong to the family Trachypachidae. Only one species of the genus Psacodromeus, P. minor, was reported from the Gurvaneren Formation in Mongolia, which is estimated as Early Cretaceous. Other species of the genus Psacodromeus, including P. crassus, P. gutta, P. ovalis, and P. rugosus, were reported from the Karabastau Formation in Kazakhstan, which is dated to the Late Jurassic. Necronectulus avus, which is the only species of the genus Necronectulus, was reported from the Tologoi Formation in Kazakhstan, which is dated to the Late Triassic. Therefore, our study has expanded the fossil record of the genera Psacodromeus and Necronectulus from the Early Cretaceous.

New fossil Anaxyelidae (Insecta: Hymenoptera: Siricoidea) from the Lower Cretaceous Yixian Formation of northeastern China

YI LI — 2024-12-31

One new genus and two new species, Hemisyntexis lepida gen. et sp. nov., Eosyntexis conflata sp. nov., and new material of Parasyntexis khasurtensis Kopylov, 2019 are respectively described and illustrated based on three well-preserved compression fossils from the Lower Cretaceous Yixian Formation of northeastern China. Eosyntexis conflata sp. nov., placed in the genus Eosyntexis Rasnitsyn, together with H. lepida, belongs to the subfamily Syntexinae of Anaxyelidae. The new species provide important morphological characters to enhance our understanding of the subfamily Syntexinae, and to revise the diagnostic characters of Eosyntexis as well. Moreover, we carried out a comprehensive survey of all currently published extant and extinct species of Anaxyelidae to gain a better understanding of the trend of change for the forewing and body length. Furthermore, we formally documented that the Anaxyelidae had four different color patterns for the wings simultaneously in the Jehol Biota of the Lower Cretaceous.