Ozymandipteryx campana sp. nov.

Named after Ozymandias, a poem by Percy Bysshe Shelley (1818) about the loss of greatness and forgetting of glory by the passing of time. It refers to the state of absent/very reduced metatarsi in the genus, which can be found in the modern-day Tridactylidae genera Ellipes Scudder, 1902 and Xya Latreille, 1809, but is not present in any extant Ripipterygidae.

Absence or almost complete reduction of the metatarsus. Protibiae without dactyls. Mesotibiae not inflated. Cerci one-segmented, cylindrical, with long hairs. Forewing and hindwing present. Metafemur with two small apical protrusions. Dorsal metatibial ridge distally with slight serration. Apical spurs of metatibia much longer (4×) than subapical spurs.

The species’ name is Latin for “bell”. It is meant to refer to the “Great Bell of Dhammazedi”, a treasure lost in time.

The specimen was included in amber found in Hkamti, Sagaing Division, Myanmar or Tanai, Kachin State Burma, Myanmar, two nearby amber mining locations. The amber from Hkamti is ca. 110 My and the amber from Tanai is ca. 99 My old.

Sex unknown. Specimen part of the LIB-Hamburg collection; collection number GPIH07209 (ex collection Martin Husemann MH0082).

As for genus (monotypic).

Preservation of complete specimen, unfortunately with some parts obscured by structural damage inside the amber.

Measurements: Some measurements could not be precisely made, because the corresponding feature was somewhat obscured. Such measurements are indicated with a “ca.”. Body length ca. 1.72–1.76 mm (head to abdominal apex). Forewing length ca. 0.72 mm. Hindwing length ca. 1.26 mm. Head height ca. 0.5 mm. Mesofemur length 0.89 mm. Mesotibia length 0.72 mm. Mesofemur width/mesotibia width 1.18 (both at max. width). Pronotum length (dorsal) 0.42 mm. Metafemur length 1.18 mm, width 0.54 mm. Metatibia length 1.17 mm, width 0.09 mm. Length of subapical spurs 0.08 mm and of apical spurs 0.32 mm. Cercus length ca. 0.24 mm.

Head: Antennae 10-segmented (including scape and pedicel). 10^th antennomere larger than previous and bean shaped. Compound eyes protruding from head. Ocelli not visible.

Thorax: Pronotum not prolonged over abdomen, with elevation from thorax. Both forewings and hindwings present. Hindwings not prolonged over abdomen.

Legs: Prothoracic leg: Tibia inflated. Dorsal margin with four evenly spaced hairs. No dactyls present. Tarsus very slender, with two claws.

Mesothoracic leg: Femur without hair or appendages. Tibia not inflated, with hairs on ventral margin along the entire length and also on dorsal margin towards apex. Tarsus two-segmented, bearing two claws.

Metathoracic leg: Femur greatly inflated along its entire length. Two small protrusions apically. Dorsal tibial ridge distally with slight serration proximal to subapical spurs. Two subapical and two apical spurs present. Apical spurs much longer than subapical spurs. Metatarsus not visible, either absent or too vestigial to be seen.

Abdomen: Only one cercus visible, one-segmented and cylindrical in shape. Setulose with long hair.

The new species can be assigned to the Ripipterygidae based on the following characters: 1) Cercus one-segmented and 2) mesotibiae not inflated. In this family, it differs from the other members found in Kachin amber by an absent or very vestigial metatarsus, a protibia without dactyls and two small protrusions on the apex of its metafemur. The protibia without dactyls is a character the new species shares with some members of fossil Tridactylidae: Burmadactylus grimaldii Heads, 2009, Paraxya huiCao et al., 2019 and Ellipes dominicana Poinar, 2020. Likewise, a vestigial metatarsus is also present in some extant Tridactylidae, namely the genera Ellipes Scudder, 1902 and Xya Latreille, 1809 (Günther 1977; Heads 2010).

Based on our current understanding of Tridactyloidea morphology, the presence of characters only known from Tridactylidae among the extant fauna in a fossil specimen that should be placed in Ripipterygidae, suggests four possible conclusions: 1) Our understanding of Tridactyloidea morphology is not complete. 2) A fully reduced metatarsus and protibia without dactyls were present for some time in the ancestors of modern Ripipterygidae, but this lineage was subsequently lost. 3) Ozymandipteryx does not actually belong to the Ripipterygidae but is instead a member of a stem-group Tridactyloidea occurring prior to the split of Tridactylidae and Ripipterygidae. 4) Ozymandipteryx is part of a separate lineage of Tridactyloidea not yet described. If one of the last two hypotheses is true, attribution of fossil Tridactyloidea from the Cretaceous may have to be reconsidered; however, this can only be validated by future findings.

Interestingly, O. campana is not the first fossil species with a fully reduced metatarsus. In the description of Magnidactylus robustus Xu et al., 2020, the apical spurs of the specimen were interpreted as the metatarsus. But in fact, the metatarsus appears to be absent in M. robustus just like in O. campana. This finding may suggest a closer phylogenetic relationship between the two species. However, M. robustus differs from O. campana by the presence of four dactyls on its protibia, no hairs on the cerci and no apical protrusions on the metafemur. Further, M. robustus is more than three times the size of O. campana (Xu et al., 2020). With this re-interpretation of the morphology of M. robustus, the species does not belong to the same genus as Magnidactylus mirusGu et al., 2022 and M. gracilisGu et al., 2022, which both possess a normal (for Tridactyloidea) metatarsus. We propose Yakkhapipteryx gen nov. as a new genus to house M. mirus and M. gracilis.

Yakkhapipteryx mirus comb. nov. (Gu et al., 2022)

Yakkhapipteryx mirus comb. nov., Y. gracilis comb. nov. (Gu et al., 2022)

Named after the Yakkha (Sanskrit: Yaksha), guardians of buried treasures (in reference to amber) from Burmese mythology.

As provided by Gu et al. (2022) for Magnidactylus, which the authors based on the specimens now assigned to Yakkhapipteryx.

Ciconipteryx bidactylus sp. nov.

The new genus is named after its long legs. Ciconi- is derived from the bird family Ciconiidae, the storks.

Very small body. Interocular distance larger than width of compound eye. Protibia with two dactyls. Mesothoracic leg longer than body (head to abdominal apex) and very slender along entire length (femur and tibia not much wider than tarsus). Metafemur as long as body. Forewings present, four to five veins visible. Cercus somewhat longer than paraproctal lobes. Both structures setulose; setae more prominent on paraproctal lobes. Valves of ovipositor visible, much shorter than paraproctal lobes.

The species’ name refers to the protibia with (only) two dactyls.

The specimen was included in amber found in Hkamti, Sagaing Division, Myanmar or Tanai, Kachin State Burma, Myanmar, two nearby amber mining locations. The amber from Hkamti is ca. 110 My and the amber from Tanai ca. 99 My old.

Female. Specimen part of the LIB-Hamburg collection, collection number GPIH07210 (ex collection Martin Husemann MH0093).

As for genus (monotypic).

The specimen is relatively well preserved, but unfortunately the apical region of the metatibia and the metatarsus are missing.

Measurements: Body length (head to abdominal apex) 1.4 mm. Head height 0.6 mm. Eye height 0.21 mm, width 0.11 mm. Interocular distance (mid-eye level) ca. 0.2 mm. Protibia 0.24 mm long. Longest dactyl 0.05 mm long. Mesofemur 0.67 mm long. Mesotibia 0.59 mm long. Mesotarsus 0.27 mm long. Metafemur 1.4 mm. Cercus 0.12 mm long. Paraproctal lobes 0.1 mm long.

Head: Antennae not preserved. Interocular distance wider than compound eye width. Eyes somewhat tear-shaped, laterally protruding from head. Ocelli not visible.

Thorax: Pronotum with posterior margin straight (not rounded or pointed), covering base of forewings. Pronotum without elevation. Forewings present with four or five simple veins visible. Hindwings present, shorter than abdomen.

Legs: Prothoracic leg: Femur and tibia setulose with long hairs. Tibia only slightly inflated with two relatively long dactyls. Tarsus with two claws.

Mesothoracic leg: Very long and slender along entire length. Femur inconspicuous. Tibia not inflated. With 8–11 fine spines on the ventral margin of the distal half and five hairs subapically on the dorsal side. Tarsus two-segmented with first segment much shorter than second. Bulbous apical lobe on first tarsal segment. Second tarsal segment with two claws.

Metathoracic leg: Femur long and inflated along its entire length. Only ca. proximal third of metatibia preserved.

Abdomen: Cercus one-segmented, cylindrical in shape with some hairs. Paraproctal lobes one-segmented, sligthly clavate and slightly shorter than cercus; setulose with long hair, especially apically. From a dorsal view apices of ovipositor valves just visible between paraproctal lobes.

Ciconipteryx bidactylus gen. et sp. nov. is assigned to Ripipterygidae based on the following characters: 1) Cercus one-segmented. 2) Mesotibiae not inflated. 3) Ovipositor visible. 4) Paraproctal lobes with distinct array of setae. It differs from previously described genera of Kachin amber Ripipterygidae by its two dactyls on the protibia (in other species from this locality and horizon there are either four dactyls or 0, as in Ozymandipteryx campana gen. et sp. nov.) and the markedly longer mesothoracic leg and metafemur which are longer or just as long as the body, respectively. The two dactyls on the protibia represent a character shared between Ciconipteryx and some species of the fossil as well as extant genus Mirhipipteryx Günther, 1969. While certain species of this genus, such as M. antillarum Heads, 2010, are reported to have three protibial dactyls (Heads 2010), some modern species such as M. pulicaria (Saussure, 1896) are said to have only two (Baena-Bejarano et al. 2018), as in C. bidactylus. However, Ciconipteryx can be differentiated from Mirhipipteryx by its proportionally longer legs (in M. pulicaria the mesothoracic leg length is only ca. 60% of the total body length) and its interocular distance between the compound eyes, which is wider than the compound eye width in Ciconipteryx, but significantly shorter in Mirhipipteryx (Heads 2010; Baena-Bejarano 2018). Species of the other extant Ripipterygidae genus, Ripipteryx Newman, 1834, are much larger than Ciconipteryx and Mirhipipteryx (Heads 2010).