The family Cretapsychidae (Insecta, Trichoptera) from mid-Cretaceous Burmese amber, with descriptions of two new species

. The Cretaceous family Cretapsychidae Wichard, 2021 belongs to the superfamily Sericostomatoidea Stephens, 1836 and is characterized by the five-segmented maxillary palps, with the second segment being the longest and the following three segments becoming successively smaller, and the tibial spur formula 2/2/4. In addition, the wing venation is characterized by apical forks II, III, and V in the forewings and forks I, III, and V in the hindwings. The two newly described species ( C. kachini sp. nov. and C. myanmari sp. nov.) extend the genus Cretapsyche Wichard, Neumann, Müller & Wang, 2018 to six extinct species, all of which were found in mid-Cretaceous Burmese amber from Kachin State, Myanmar. ings Forewings broader than hindwings, each with rounded apex. In forewings, forks II, III, and V present; cubitus vein Cu2 ending in Cu1b or connected to Cu1b by crossvein. Discoidal, median, and thyridial cells present or absent. Nygmata absent. Anal vein reaching wing margin before fork V or before crossvein reaching wing margin. Hindwings with forks I, III, and V present. M ale geniTalia . Inferior appendages two-segmented, consisting of basal coxopodite and harpago simply attached to apex of coxopodite. Coxopodite arcuate, sub-triangular or scaly, broad, long and curved; harpago finger- or needle-shaped, often straight at base and curved mesad at apex. Two parallel appendages located centrally in genitalia interpreted as probable mesodorsal pair of lobes of tergum X. Preanal appendages small, elongate or finger-like curved.


Material and methods
The specimens are from an amber mine likely located near Noije Bum Village, Tanaing Township, Myanmar (Kania et al. 2015), but the exact locality is unknown. The age given by U-Pb dating of zircons from the volcanoclastic matrix of the amber is early Cenomanian (98.8 ± 0.6 Ma) (Shi et al. 2012), but the geological age of Burmese amber can be expected to be slightly older than the zircon date.
The two fossil specimens are embedded in two small ambers cut from larger pieces of Burmese amber. They are almost completely intact and visible in ventral and dorsal views. The male genitalia are distinctly flattened and distorted, covered dorsally and laterally by the wings and therefore visible only in ventral or ventral-lateral view. Significant characters are often limited to the ventrally located inferior appendages. Hindwings are partially covered by forewings. Head, thorax, abdomen, and antennae are often complete or partially complete. The general wing pattern of Cretapsyche can be found in Fig. 1.   Fig. 1. Cretapsyche Wichard, Neumann, Müller & Wang, 2018, wing venation in forewing with forks II, III, and V present, discoidal and thyridial cells open or closed, nygmata absent, in hindwing with forks I, III, and V usually present.
European Journal of Taxonomy 833: 1-11 (2022) Photos were taken using a Leica M 420 Apozoom stereo microscope in combination with a Canon EOS 80D, EOS utility software and the Zerene Stacker software. All images and figures were prepared with CorelDraw X4 and Adobe Photoshop CS4.

Abbreviations
The wing venation terminology in general follows Holzenthal et al. (2007)

Revised description
Head. Male adult embedded in amber, light brown coloration. Large complex eyes on sides of head, slightly longer in diameter than head length. Ocelli absent. Antennae as long as, or slightly longer than forewings, each with scapus about as long as head, often bearing brush of dark setae; flagellomeres uniformly bar-shaped. Maxillary palps five-segmented; second segment longest, following three segments successively smaller and shorter; fifth segment shortest (Figs 2-4).
MesoscuTelluM. With one domed setal wart or pair of domed setal warts fused medially. European Journal of Taxonomy 833: 1-11 (2022) Wings ( Fig. 1). Forewings broader than hindwings, each with rounded apex. In forewings, forks II, III, and V present; cubitus vein Cu2 ending in Cu1b or connected to Cu1b by crossvein. Discoidal, median, and thyridial cells present or absent. Nygmata absent. Anal vein reaching wing margin before fork V or before crossvein reaching wing margin. Hindwings with forks I, III, and V present.
Male geniTalia. Inferior appendages two-segmented, consisting of basal coxopodite and harpago simply attached to apex of coxopodite. Coxopodite arcuate, sub-triangular or scaly, broad, long and curved; harpago finger-or needle-shaped, often straight at base and curved mesad at apex. Two parallel appendages located centrally in genitalia interpreted as probable mesodorsal pair of lobes of tergum X. Preanal appendages small, elongate or finger-like curved.  Ocelli absent. Antennae probably about as long as forewings (holotype antennae incomplete). Scapus about as long as head. Maxillary palps 5-segmented, 2 nd segment longest, following segments decreasing in length, 3 rd segment basally barrel-shaped and extended laterad to form pointed spur corresponding in length to 4 th and 5 th segments. Body and wings light brown. Forewing with length 2.8 mm longer than small hindwing. In forewings, apical forks II, III and V present, discoidal cell absent, thyridial cell present, Cu2 running continuously to Cu1b. In hindwings, apical forks I, III and V present (difficult to see, holotype forewing hides hindwings in dorsal view). Tibial spur formula 2/2/4. In male genitalia, two-segmented inferior appendages characterized by basally broad coxopodite, tapering and curving mesad. Harpago elongate and tapering continuously towards apex, two-thirds of coxopodite length.

Male
Ocelli absent. Antennae little longer than forewings. Scapus about as long as head, smaller pedicellus, followed by more than 30 cylindrical flagellomeres gradually shortening in length apically. Maxillary palps 5-segmented, 2 nd segment longest, following three segments successively smaller and shorter. Body and wings light brown. Forewing with length about 2.8 mm. In forewings, apical forks II, III and V present, discoidal and thyridial cells absent, Cu2 running continuously to Cu1b. In hindwings, apical forks I, III and V present (holotype hindwings partially spread laterally and visible). Tibial spur formula 2/2/4. In male genitalia, two-segmented inferior appendages characterized by finger-shaped, curved coxopodite and by short and cone-shaped harpago. Two appendages, each with two dark spines, located centrally in genitalia interpreted as probable mesodorsal pair of lobes of tergum X (Wichard 2021).

Discussion
The extinct family Cretapsychidae belongs to the superfamily Sericostomatoidea following Johanson et al. (2017), characterized by the absence of ocelli (except for Anomalopsychidae Flint, 1981), by the terminal maxillary and labial segments both not being flexible or annulate, and by the tibial spur formula being 2/2/4, the latter stated to be a synapomorphy for the superfamily by Ross (1967). The family Cretapsychidae is distinguished from all other families of Sericostomatoidea by its wing venation (the absence of forks I and IV in the forewings and of forks II and IV in the hindwings) and by its maxillary palps (second segment longest, the following three segments successively smaller and shorter; fifth segment shortest).
In the presence of forks II, III, and V and the absence of forks I and IV in the forewings, the extinct Cretapsychidae agree with the extinct Burmapsychidae Wichard, 2021 and the extant Helicophidae Mosely, 1953. However, the Burmapsychidae differ from the Cretapsychidae in the latter's threesegmented maxillary palps in males, in the scapus being longer than the head, and in the modified flagellomeres of the antennae (Wichard 2021).
The extant Helicophidae and the extinct Cretapsychidae differ significantly in their geographic distributions and paleogeological history. The family Helicophidae is distributed in the southern hemisphere in the Neotropical (Chile, Patagonia) and Australasian (South Australia, New Zealand, New Caledonia) regions. The family Cretapsychidae is found so far only in Southeast Asia in Burmese amber from the Middle Cretaceous. Evidence of the extinct family dating back nearly 100 million years (early Cenomanian) does not suggest a shared history with the Gondwanan family Helicophidae, especially since the Australian continent did not drift into the southwest Pacific and thus into the geographic proximity of what is now Southeast Asia until the late Eocene (Hall 2011