Vladimir N. Makarkin

Tertiary giant lacewings (Neuroptera: Polystoechotidae): Revision and description of new taxa from western north America and Denmark

Synopsis A preliminary definition of the Polystoechotidae (Insecta, Neuroptera) based on wing characters is provided. Tertiary Polystoechotidae are revised; the genus Palaeopsychops Andersen (10 species) and the new collective genus Polystoechotites gen. n. (6 species) are recognised. Ten new species (eight named) are described from the earliest Early Eocene Fur Formation (Denmark) and the Early to early Middle Eocene Okanagan Highlands (Quilchena and Horsefly River, British Columbia, Canada and Republic, Washington, USA): Palaeopsychops quadratus sp. nov. (Fur Formation); Palaeopsychops setosus sp. nov. (Horsefly River); Palaeopsychops marringerae sp. nov., Palaeopsychops timmi sp. nov., Polystoechotites lewisi gen. et sp. nov., Polystoechotites barksdalae gen. et sp. nov., Polystoechotites falcatus gen. et sp. nov. (Republic); Palaeopsychops douglasae sp. nov., Polystoechotites sp. A, Polystoechotites sp. B (Quilchena). Polystoechotes piperatus Cockerell from Late Eocene of Florissant (Colorado, USA) is assigned to Polystoechotites. Neuroptera incertae sedis sp. A (Republic) is determined as either an aberrant polystoechotid, or belonging to a family of psychopsid‐like Neuroptera. Palaeopsychops setosus sp. nov. bears a dense cover of macrotrichia across a portion of the wing membrane, not previously known in the order. Extended floating time of polystoechotids increased negative taphonomic bias, in most depositional settings probably lowering their fossil assemblage representation. In some, taphonomic factors such as the presence of mat‐forming organisms may have enhanced their fossil representation to greater than their community representation. This scenario is proposed for the Fur Formation, consistent with lithological evidence and palaeogeographic reconstruction. Dispersal of Palaeopsychops between Denmark and the Okanagan Highlands (direction unknown) was probably by the North Atlantic route, not before the late Palaeocene, during periods of continuous land connection between North America and northern Europe. Polystoechotidae in the Eocene, as today, appear to have ranged in forested regions of microthermal to lower mesothermal climate.

A Remarkable New Family of Jurassic Insects (Neuroptera) with Primitive Wing Venation and Its Phylogenetic Position in Neuropterida

Background Lacewings (insect order Neuroptera), known in the fossil record since the Early Permian, were most diverse in the Mesozoic. A dramatic variety of forms ranged in that time from large butterfly-like Kalligrammatidae to minute two-winged Dipteromantispidae. Principal Findings We describe the intriguing new neuropteran family Parakseneuridae fam. nov. with three new genera and 15 new species from the Middle Jurassic of Daohugou (Inner Mongolia, China) and the Early/Middle Jurassic of Sai-Sagul (Kyrgyzstan): Parakseneura undula gen. et sp. nov., P. albomacula gen. et sp. nov., P. curvivenis gen. et sp. nov., P. nigromacula gen. et sp. nov., P. nigrolinea gen. et sp. nov., P. albadelta gen. et sp. nov., P. cavomaculata gen. et sp. nov., P. inflata gen. et sp. nov., P. metallica gen. et sp. nov., P. emarginata gen. et sp. nov., P. directa gen. et sp. nov., Pseudorapisma jurassicum gen. et sp. nov., P. angustipenne gen. et sp. nov., P. maculatum gen. et sp. nov. (Daohugou); Shuraboneura ovata gen. et sp. nov. (Sai-Sagul). The family comprises large neuropterans with most primitive wing venation in the order indicated by the presence of ScA and AA1+2, and the dichotomous branching of MP, CuA, CuP, AA3+4, AP1+2. The phylogenetic position of Parakseneuridae was investigated using a phylogenetic analysis of morphological scoring for 33 families of extinct and extant Neuropterida combined with DNA sequence data for representatives of all extant families. Parakseneuridae were recovered in a clade with Osmylopsychopidae, Prohemerobiidae, and Ithonidae. Conclusions/Significance The presence of the presumed AA1+2 in wings of Parakseneuridae is a unique plesiomorphic condition hitherto unknown in Neuropterida, the clade comprising Neuroptera, Megaloptera, Raphidioptera. The relative uncertainty of phylogenetic position of Parakseneuridae and the majority of other families of Neuroptera reflects deficient paleontological data, especially from critical important periods for the order, earliest Triassic and latest Triassic/earliest Jurassic.

Phylogeny of Moth Lacewings and Giant Lacewings (Neuroptera: Ithonidae, Polystoechotidae) Using DNA Sequence Data, Morphology, and Fossils

ABSTRACT A phylogeny of lacewing families Ithonidae and Polystoechotidae is presented based on three gene markers (16S and 18S ribosomal DNA and CAD) and 23 morphological characters. Living and fossil genera presently placed in Polystoechotidae (Fontecilla Navás, Platystoechotes Carpenter and Polystoechotes Burmeister) and Ithonidae (Adamsiana Penny, Allorapisma Makarkin & Archibald, Ithone Newman, Megalithone Riek, Oliarces Banks, PrincipialaMakarkin & Menon, Rapisma Walker and Varnia Walker) were included in phylogenetic analyses (parsimony and Bayesian) and compared with outgroups selected from various families of Neuroptera. The resulting phylogeny recovered a monophyletic clade comprising Ithonidae and Polystoechotidae as hypothesized previously. Rapismatidae as a separate family is not supported and Ithonidae are rendered paraphyletic with three extant genera previously placed in Ithonidae (Adamsiana, Oliarces, and Rapisma), recovered deep within Polystoechotidae. The fossil genera Allorapismaand Principialaformed a sister-group relationship with Rapisma, also within Polystoechotidae. Due to the lack of mutually exclusive synapomorphies for either Ithonidae or Polystoechotidae, a single more inclusive family Ithonidae is proposed, including all ithonid genera and all genera previously placed in Polystoechotidae. Synapomorphies characterizing the revised concept of Ithonidae s.l. are discussed.

Family Affinity of the Genus Palaeopsychops Andersen with Description of a New Species from the Early Eocene of British Columbia, Canada (Neuroptera: Polystoechotidae)

Abstract Palaeopsychops dodgeorum sp. n. from the Early Eocene Okanagan Highlands of Quilchena, British Columbia, Canada is described. The systematic position of the genus Palaeopsychops Andersen, 2001 is discussed, interpreting this as most closely associated with Polystoechotidae. Osmylites protogaea (Hagen 1862) is considered as nomen nudum and an objective synonym of Osmylites excelsa (Oppenheim, 1888), syn. n.

Two New Species of Kalligramma Walther (Neuroptera: Kalligrammatidae) From the Middle Jurassic of China

ABSTRACT Two new fossil species of Kalligramma Walther (Neuroptera: Kalligrammatidae) are described from the Middle Jurassic locality at Daohugou, Inner Mongolia, China: Kalligramma elegans n. sp. and Kalligramma albifasciatum n. sp. The subcosta anterior wing vein is found in both species. K. albifasciatum n. sp. is remarkable by its forewing color pattern in having the broad white transverse fascia through wing, hitherto unknown in the genus. Kalligramma turutanovae Martynova is transferred to Huiyingogramma Liu et al., n. sit. A new diagnosis of the genus Kalligramma is provided (amended from Panfilov 1968).

The presence of the recurrent veinlet in the Middle Jurassic Nymphidae (Neuroptera): a unique character condition in Myrmeleontoidea.

Abstract A well-developed recurrent veinlet is found in the forewing of two species of Nymphidae from the Middle Jurassic locality of Daohugou (Inner Mongolia, China), Liminympha makarkini Ren & Engel and Daonymphes bisulca gen. et sp. n. This is the first record of this trait in the clade comprised of the superfamilies Myrmeleontoidea and Chrysopoidea. We interpret the recurrent veinlet in these species as a remnant of the condition present more basally in the psychopsoid + ithonoid + chrysopoid + myrmeleontoid clade (i.e., as a plesiomorphy). Other venational character states of Daonymphes bisulca of interest include the configuration of subcosta anterior (ScA), which is very similar to that of extant Nymphidae. We consider the short ScA terminating on ScP to be an autapomorphy of Neuroptera.

Two New Species of Kalligrammatidae (Neuroptera) from the Jurassic of China, with Comments on Venational Homologies

ABSTRACT Two new fossil species of Kalligrammatidae (Neuroptera) are described from the Middle Jurassic locality at Daohugou, Inner Mongolia, China: Limnogramma hani n. sp. and Limnogramma mongolicum n. sp. Some venational homologies in Neuroptera are discussed briefly.

A Diverse New Assemblage of Green Lacewings (Insecta, Neuroptera, Chrysopidae) from the Early Eocene Okanagan Highlands, Western North America

Abstract The early Eocene green lacewings (Neuroptera, Chrysopidae) of the Okanagan Highlands deposits of McAbee, and Driftwood Canyon, British Columbia (Canada) and Republic, Washington (U.S.A.) are treated in detail for the first time. At least six genera are present, one unnamed, three new, with at least 10 new species, six named: Protochrysa fuscobasalis n. sp. (McAbee) (Limaiinae, the youngest known record of the subfamily), Okanaganochrysa coltsunae n. gen. n. sp. (McAbee), Adamsochrysa aspera n. gen. n. sp. (McAbee), A. wilsoni n. gen. n. sp. (Republic), Archaeochrysa profracta n. sp. (McAbee), and Pseudochrysopa harveyi n. gen. n. sp. (Driftwood Canyon) (all Nothochrysinae, the latter provisionally). The four unnamed species include one assigned to Pseudochrysopa, two likely belonging to Adamsochrysa, and one of an unknown nothochrysine genus. Microtholi are detected on the abdominal sclerites of Adamsochrysa wilsoni, and the spermatheca and spermathecal duct in the abdomen of Pseudochrysopa harveyi, the first reported occurrences of these preserved in fossil Chrysopidae. Structures were detected on the apical wing margins of some species that appear similar to trichosors, which are unknown in Chrysopidae, but are present in some other neuropteran families. This is the richest described assemblage of the family anywhere in the fossil record. Okanagan chrysopids were also morphologically and presumably ecologically diverse, including large species with rich venation and well as those with simplified venation and the smallest known fossil species. This is the oldest reported occurrence of the family in North America.

New psychopsoid Neuroptera from the Early Cretaceous of Baissa, Transbaikalia

Abstract Two new genera and fi ve new species of the psychopsoid Neuroptera (one of which is unnamed) are described from the Early Cretaceous locality at Baissa, Transbaikalia, Russia: Epipsychopsis fusca n. gen., n. sp., E. variegata n. sp. (Psychopsidae), Pseudopsychopsis gradata n. gen., n. sp., P. baissensis n. sp. (Brongniartiellidae), and Sophogramma sp. (Kalligrammatidae).

Two new species of Sinosmylites Hong (Neuroptera, Berothidae) from the Middle Jurassic of China, with notes on Mesoberothidae

Abstract Two new species of the genus Sinosmylites Hong are described from the Middle Jurassic locality at Daohugou (Inner Mongolia, China): Sinosmylites fumosus sp. n. and Sinosmylites rasnitsyni sp. n. This is the oldest known occurrence of the family Berothidae. The berothid affinity of this genus is confirmed by examination of the hind wing venation characteristic of the family. The Late Triassic family Mesoberothidae may represent an early group of Berothidae.