Kalligrammatidae | |
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Kalligrammatid diversity | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Neuroptera |
Superfamily: | Psychopsoidea |
Family: | † Kalligrammatidae Handlirsch, 1906 |
Subfamilies | |
Kalligrammatidae, sometimes known as kalligrammatids or kalligrammatid lacewings, is a family of extinct insects in the order Neuroptera (lacewings) that contains twenty genera and a number of species. The family lived from the Middle Jurassic to the early Late Cretaceous before going extinct. Species of the family are known from Europe, Asia, and South America. The family has been occasionally described as "butterflies of the Jurassic" based on their resemblance to modern butterflies in morphology and ecological niche.
The known distribution of Kalligrammatidae is widespread both in time and in location. Fossils of the family have been recovered from sediments in Western Europe, the British Isles, Central Asia, and China. The majority of described species, thirty one, are from Jurassic and Cretaceous fossils found in China. Eight species are known from Kazakhstan, the second largest number of species for a single country, while only two species are represented by fossils found in Russia, though only one is described due to the incomplete nature of the other specimen. One species has been described from Mongolia. Species from Europe are much less common, with four species from Germany and one species from England. [1] Only one genus has been described from the Western Hemisphere, with two species found in Brazil. [2] Six of the genera have been found at two or more locations; Kalligrammula and Kalligramma are the most widely distributed genera. [3]
Jurassic species are found in both Asia and Europe. The oldest described species are from the early Toarcian Posidonia Shale in Germany, [4] with the next youngest being from the Callovian Haifanggou and Daohugou Formations in China. [3] [5] Cretaceous species are less common, but still found across Eurasia, the youngest being from China and Britain and the family lasting until the Aptian in Brazil (Crato Formation). [2] In 2018, it was recognised that the subfamily Cretanallachiinae from the Burmese amber, formerly assigned to Dilaridae, belonged to Kalligrammatidae, extending the range of the family into the early Late Cretaceous. [6]
Most species are known from compression-impression fossils preserved in layers of soft sedimentary rock. Many of the species are known only from isolated fore or hind wings, though full bodies are known for some species. Species are typified by bodies that are over 50 mm (2.0 in) long when known and covered in dense layers of setae. The antennae are generally not longer than the length of the fore wings and have a simple filiform structure. There is variation in the mouthparts, which commonly are 11–25 mm (0.43–0.98 in) long [3] [7] siphon shaped proboscis, but some basal species have more distinct mandibles. [8] The proboscis is formed from the same mouth parts as those of Nymphalidae butterflies and were used for probing and sucking. Species of at least one genus, Oregramma , have elongated lance shaped ovipositors. The wings are distinctly large, over 50 mm (2.0 in) long, often with centrally placed eye spots and the ovoid to triangular wings have numerous closely spaced branching veins. [1] Most of the species also have distinctly developed wing scales, a feature seen in Lepidopterans. Two types of scales are seen in the kalligrammatids, shorter scales with a broad base that taper to a tip, and longer narrow scales with a spatulate shape. [8] The species Makarkinia adamsi has the longest forewing of any neuropteran species, estimated at 160 mm (6.3 in). [2]
It is thought that their large body size and large wings would have made kalligrammatids weak fliers. [1] Wing color patterning on many species and the setae covering on Makarkinia are indications that the species were day-time fliers similar to butterflies. The prevalence of eye spots in many species, the presence of light striping along the wing margins of Sophogramma [1] were evolved to act as anti-predator adaptations against predation from pterosaurs and early birds. Given the structuring of the mouthparts, they are thought to have most likely been pollinators, feeding on pollen and plant juices, [1] [2] [7] possibly produced by Bennettitales and Cheirolepidiaceae. [3] The pollination behavior is nearly unique in the Neuroptera, most of which are predatory. The similarity of features and ecology between lepidopterans and kaligrammatids has led to the group occasionally being called "butterflies of the Jurassic". [1] The only modern neuropteran family that feeds on pollen is Nemopteridae, and kalligrammatids are the only neuropterans which have developed proboscises. [7] As flowering plants emerged and diversified, the host plants of kalligrammatids dwindled, possibly resulting in their extinction. [7]
There are currently six described subfamilies of kalligrammatid, with all but two genera placed into them. The remaining two genera Makarkinia and Palparites have been left incertae sedis . [1]
A phylogeny of the family was produced in 2014 by a team of researchers showing the family to have 4 distinct subfamily clades. A placement of Palparites was not made due to the incomplete nature of the only fossil known, and Makarkninia was not included in the paper, as the second more complete species was not described until 2016. [1]
Kalligrammatidae | |
The insect order Neuroptera, or net-winged insects, includes the lacewings, mantisflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera is grouped together with the Megaloptera and Raphidioptera (snakeflies) in the unranked taxon Neuropterida.
Mantispidae, known commonly as mantidflies, mantispids, mantid lacewings, mantisflies or mantis-flies, is a family of small to moderate-sized insects in the order Neuroptera. There are many genera with around 400 species worldwide, especially in the tropics and subtropics. Only five species of Mantispa occur in Europe. As their names suggest, members of the group possess raptorial forelimbs similar to those of the praying mantis, a case of convergent evolution.
Osmylidae are a small family of winged insects of the net-winged insect order Neuroptera. The osmylids, also called lance lacewings, stream lacewings or giant lacewings, are found all over the world except North and Central America. There are around 225 extant species.
Hemerobiidae is a family of Neuropteran insects commonly known as brown lacewings, comprising about 500 species in 28 genera. Most are yellow to dark brown, but some species are green. They are small; most have forewings 4–10 mm long. These insects differ from the somewhat similar Chrysopidae not only by the usual coloring but also by the wing venation: hemerobiids differ from chrysopids in having numerous long veins and forked costal cross veins. Some genera are widespread, but most are restricted to a single biogeographical realm. Some species have reduced wings to the degree that they are flightless. Imagines (adults) of subfamily Drepanepteryginae mimic dead leaves. Hemerobiid larvae are usually less hairy than chrysopid larvae.
The Berothidae are a family of winged insects of the order Neuroptera. They are known commonly as the beaded lacewings. The family was first named by Anton Handlirsch in 1906. The family consists of 24 genera and 110 living species distributed discontinuously worldwide, mostly in tropical and subtropical regions. Numerous extinct species have also been described. Their ecology is poorly known, but in the species where larval stages have been documented, the larvae are predators of termites.
The Nevrorthidae are a small family of lacewings in the order Neuroptera. There are 19 extant species in four genera, with a geographically disjunct distribution: Nevrorthus, comprising 5 species with scattered distributions around the Mediterranean; Austroneurorthus, with two species known from southeastern Australia; Nipponeurorthus, comprising 11 species known from China and Japan; and Sinoneurorthus, known from a single species described from Yunnan Province, China. They are traditionally placed in the Osmyloidea, alongside Osmylidae and the spongillaflies (Sisyridae), but some research has considered them to be the sister group to the rest of Neuroptera. The larvae have unique straight jaws that are curved at the tips, and live as unspecialised predators in the sandy bottom sediments of clear, fast flowing mountain rivers and streams. They pupate underwater on the underside of stones. The adults are likely predators or feed on honeydew and other sugar-rich fluids.
Nymphidae, sometimes called split-footed lacewings, are a family of winged insects of the order Neuroptera. There are 35 extant species native to Australia and New Guinea.
Psychopsidae is a family of winged insects of the order Neuroptera. They are commonly called silky lacewings.
Sisyridae, commonly known as spongeflies or spongillaflies, are a family of winged insects in the order Neuroptera. There are approximately 60 living species described, and several extinct species identified from the fossil record.
Microberotha is an extinct monotypic genus of "beaded lacewing" in the family Berothidae known from a fossil found in North America. When described the genus contained a single Ypresian-age species Microberotha macculloughi.
Ithonidae, commonly called moth lacewings and giant lacewings, is a small family of winged insects of the insect order Neuroptera. The family contains a total of ten living genera, and over a dozen extinct genera described from fossils. The modern Ithonids have a notably disjunct distribution, while the extinct genera had a more global range. The family is considered one of the most primitive living neuropteran families. The family has been expanded twice, first to include the genus Rapisma, formerly placed in the monotypic family Rapismatidae, and then in 2010 to include the genera that had been placed into the family Polystoechotidae. Both Rapismatidae and Polystoechotidae have been shown to nest into Ithonidae sensu lato. The larvae of ithonids are grub-like, subterranean and likely phytophagous.
Undulopsychopsis is an extinct genus of lacewing in the silky lacewing family Psychopsidae. The genus is solely known from a Cretaceous fossil found in China. Currently the genus is composed of a single species, Undulopsychopsis alexi.
Principiala is an extinct genus of lacewing in the moth lacewings family Ithonidae. The genus is known from Cretaceous fossils found in South America, Europe, and possibly Asia. The genus is composed of two species, the type species Principiala incerta, and Principiala rudgwickensis.
Cretomerobius is an extinct genus of lacewings in the neuropteran family Hemerobiidae known from fossils found in Asia. The genus currently contains a single species, the Aptian C. disjunctus.
Makarkinia is an extinct genus of lacewings in the family Kalligrammatidae described by Martins-Neto in 1997 from fossils found in the Crato Formation of the Araripe Basin in northeastern Brazil. The genus contains three species dating to the Late Aptian, Makarkinia adamsi, Makarkinia kerneri and Makarkinia irmae.
Rafaelnymphes is an extinct genus of lacewing in the family Nymphidae known from a fossil found in South America. The genus contains a single species, Rafaelnymphes cratoensis.
2015 in paleoentomology is a list of new fossil insect taxa that were described during the year 2016, as well as other significant discoveries and events related to paleoentomology that were scheduled to occur during the year.
Aneuretopsychidae is an extinct family of scorpionflies known from the Mesozoic. Fossils are known from the Jurassic (Callovian-Oxfordian) to the early Late Cretaceous (Cenomanian). It is part of Mesopsychoidea, a group of scorpionflies with siphonate proboscis. They are suggested to have been nectarivores, feeding off the liquid pollination drops of and acting as pollinators for now extinct insect pollinated gymnosperms such as Bennettitales.
Palaeopsychops is an extinct genus of lacewing in the moth lacewings family Ithonidae. The genus is known from Early Eocene fossils found in Europe, and North America and is composed of ten species. The ten species can be informally separated into two species groups based on veination of the forewings, the "European" and "North American" groups. When first described, the genus was placed in the family Psychopsidae, but later was moved to Polystoechotidae, which itself is now considered a subgroup of the moth lacewings.
Oregramma is a genus of "butterfly mimicking" lacewing in the extinct family Kalligrammatidae. Fossils of this genus have been found in the Early Cretaceous Yixian Formation of China. It likely fed on pollen and plant juices, unlike modern lacewings, most of which are carnivorous.