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It Came From The Wastebasket #14: The Protorthoptera Puzzle

Protorthoptera was a group of fossil insects created in the early 20^th century to categorize “primitive” neopterans – some of the earliest insects to have evolved the ability to fold their wings down over their backs. Known mostly from just fossilized forewings, they first appeared around 320 million years ago in the late Carboniferous, and after heavy losses during the Great Dying mass extinction they eventually disappeared in the mid-Triassic about 240 million years ago.

And this group was a massive wastebasket taxon.

As early as the mid-20^th century the protorthopterans were recognized as being a general taxonomic dumping ground, containing a mixture of early members of multiple different “orthopteroid” insect lineages. But invertebrate paleontologists at the time considered this collection of “primitive” insects to lack enough distinctive features to confidently separate them out from each other, and so the highly paraphyletic grouping continued to be used well into the 1990s.

An illustration of Ctenoptilus elongatus, an extinct insect. It has long thin antennae, a small grasshopper-like head, six legs each ending in two small claws, a cylindrical abdomen, and two pairs of large wings folded over its back. It's colored red, tan, and dark brown, with striped markings on its wings. — *Ctenoptilus elongatus*

But in the early 2000s this situation finally changed. Proper cladistic analysis of protorthopteran fossils identified defining features of the wing vein patterns, and many species were reclassified into various lineages within the Archaeorthoptera – which includes modern grasshoppers, crickets, and locusts along with several closely related fossil groups like the titanopterans and caloneurodeans.

“Protorthoptera” is still sometimes used in a loose sense for fossil neopteran insects that still can’t be confidently classified anywhere else, so the wastebasket isn’t entirely cleared here.

And there are some alternate classification systems (mainly proposed by Russian paleontologists) that instead consider many protorthopterans to be notopterans closely related to modern ice-crawlers, and place others as part of other modern neopteran lineages such as webspinners and true bugs.

Hopefully better fossil discoveries and future studies will eventually help clear things up, and give us a better overall picture of the evolution of these insects.

Weird Heads Month #17: Trapjaw Ants From Hell

Ants first evolved sometime in the Late Jurassic or Early Cretaceous, but only really began to diversify about 100 million years ago in the Late Cretaceous after the rise of flowering plants.

One of their evolutionary experiments around that time was a group called the haidomyrmecinae – also known as the “hell ants”.

Known from Asia, Europe, and North America, hell ants had bizarre-looking heads, possessing huge upward-curving scythe-shaped mandibles and a horn-like projection between their antennae.

They were fast-moving arboreal predators that would have fed mainly on other invertebrates such as soft-bodied beetle larvae, and unlike most modern ants their workers were probably solitary hunters. They were capable of gaping their mandibles by almost 180°, and when they got close enough to their targets the long sensory hairs around their faces triggered their jaws to snap vertically upwards, impaling their prey against their horn in a unique trap-jaw mechanism.

Some species also reinforced the exoskeleton of their horns with metal particles, strengthening them against impacts from both struggling prey and their own powerful jaws.

Ceratomyrmex ellenbergeri was one of the oddest-looking of all known hell ant species. Known from a few specimens preserved in amber, with adult workers up to 6mm long (~0.25″), it lived during the Late Cretaceous of Myanmar about 100-94 million years ago.

It had an especially pronounced horn and very long mandibles, which may have been adaptations for tackling significantly larger prey items than other hell ants.

And due to this being a species known from Burmese amber, sadly we also have to address the controversy surrounding these sorts of specimens. This amber is currently mined in incredibly dangerous conditions, often using child labor, with sales of both jewellery and paleontological specimens directly funding the ongoing violent conflict in the region.

It’s the fossil equivalent of blood diamonds, and a huge ethical dilemma for the paleontology community.

Island Weirdness #52 — Labidura herculeana

Located in the South Atlantic Ocean, Saint Helena is a small volcanic island that formed several million years ago on the Mid-Atlantic Ridge. It’s one of the most remote islands in the world, and with no native land mammals and no large predators it developed an ecosystem with several unique flightless birds, and hundreds of endemic invertebrates and plants.

The Saint Helena giant earwig (Labidura herculeana) was, as its name suggests, an enormous species of earwig. Growing as large as 8.4cm long (3.3″) it was the biggest of its kind in the world, and was completely flightless with no hind wings.

It lived in deep burrows in the arid plain and gumwood forest regions of the island, and only came out at night during the summer rains. Since it was probably descended from the shore earwig (Labidura riparia) it was likely a similar sort of opportunistic carnivore, eating smaller invertebrates and carrion.

Humans didn’t reach Saint Helena until the early 1500s, and it wasn’t until 1798 that the earwig was noticed by naturalists and given its scientific name. Then it was more or less forgotten about, and scientific interest in it didn’t start to resume until the 1960s.

But by then it was just very slightly too late.

Extensive habitat destruction and predation by invasive cats, rodents, and centipedes had taken a huge toll on the earwig, and it had become incredibly rare. The last sighting of a live individual was in 1967, and attempts to locate more for potential captive breeding programs in the 1980s and 1990s failed to find any at all.

The last traces of the species were some isolated subfossil pincers found in the mid 1990s, and the giant earwig was officially declared extinct in 2014.

Island Weirdness #43 — Flightless Flies

One of the most defining features of the true flies is a pair of wings, but various different lineages have actually become flightless.

Flightlessness is very rare in the long-legged fly family (Dolichopodidae), however, with only about 12 out of over 5000 species known to have lost functional wings — and eight of those are endemic to the Hawaiian islands.

The Koʻolau spurwing long-legged fly (Emperoptera mirabilis, sometimes classified as Campsicnemus mirabilis) was found only on Mount Tantalus in the southern Koʻolau Range of Oʻahu, close to Honolulu. About 2mm long (>0.1″), its wings were reduced to thin stiff spines, and it moved around by walking and hopping in leaf litter in the moist cool forest at elevations of about 300m (~1000ft).

Like most other long-legged flies it would have been predatory, hunting other tiny invertebrates.

The Koʻolau spurwing was actually still common on Tantalus as recently as the early 1900s, but multiple searches since the 1980s have failed to find any more of them at all. The species is most likely completely extinct, probably due to a combination of predation from invasive ants and habitat destruction from feral wild boar rooting up the forest floor.

Of the other flightless Hawaiian long-legged flies several other species are now possibly extinct — only one out of the five known Emperoptera species still definitely survives on the highest slopes of Mount Kaʻala, and one of the three Campsicnemus is either very rare or also extinct.

The Hawaiian islands also have three endemic species of flightless crane fly in the genus Dicranomyia, all of which are incredibly rare.

Kalligrammatids

Did you know butterflies weren’t the first insects to look like butterflies?

Lepidopterans (the group of insects containing moths and butterflies) have been around since the Late Triassic – but it wasn’t until the diversification of flowering plants during the Cretaceous that recognizable moths would have evolved, and true butterflies didn’t actually appear until the early Cenozoic.

Before then, back in the mid-Jurassic about 165 million years ago, a completely different group of insects convergently evolved remarkably butterfly-like features such as large colorful scaled wings and long sucking proboscises.

Known as the kalligrammatids, these insects were giant members of the lacewing group, related to modern forms like antlions and owlflies. But unlike their predatory relatives the kalligrammatids were specialized pollinators, possibly having a mutualistic relationship with the flower-like cones of bennettitales or the pollination drops of some types of conifers. They seem to have originated in China and were found across Asia and Europe by the Late Jurassic, but a few fossils from South America suggest they were even more widespread and may just have a poor fossil record.

They reached wingspans of up to 16cm (~6″), comparable to some of the largest modern butterflies, and often sported conspicuous anti-predator markings on their wings such as stripes and eyespots – so it’s not surprising that they’re often nicknamed the “butterflies of the Jurassic”.

Rather ironically, the extinction of the kalligrammatids was probably linked to the rise of the flowering plants that the true butterflies would later be so dependent on. As flowers diversified and plants like the bennettitales declined, the kalligrammatids dwindled and disappeared, with the last known fossil record coming from the mid-Cretaceous of Brazil about 113 million years ago.

But while they were around, I do wonder if they also exhibited some similar behaviors – such as mud-puddling for extra nutrients, and specifically the habit of drinking the tears of larger animals that we see in some species. Perhaps some non-avian dinosaurs like this Dilong occasionally put up with kalligrammatids sitting on their faces!

Linguamyrmex

Linguamyrmex vladi, an ant from the Late Cretaceous of Myanmar (~99 mya). Part of an extinct group known as the Haidomyrmecini, or “hell ants”, it measured about 5mm long (0.2″) and is known from several individuals in amber.

It had huge scythe-shaped mandibles and a horn-like appendage on its head which together formed a powerful trap-jaw mechanism, snapping vertically shut when a pair of long sensitive trigger hairs touched against a target. One specimen was preserved close to a large soft-bodied beetle larva, which may have been an intended prey item.

When closed, the mandibles formed a tube-like channel to Linguamyrmex’s mouth, allowing it to suck out the “blood” from its impaled victims – and inspiring its species name, referencing Vlad Dracula.

The horn was also reinforced with metal particles in the chitinous exoskeleton, strengthening it against the impact of its closing jaws.