1. Earliest ant mimics - life history and
evolution of Alienopteridae
Cihang LUO
Nanjing Institute of Geology and Palaeontology,
Chinese Academy of Sciences
University of Chinese Academy of Sciences
2. 1. Introduction
1.1 Structural features
Long, filiform
and
multisegmented
antennae
Biting mouthpart
Unequal claws
Raptorial
forelegs
Cehalo-pronotal
scissor-device
Large pan-
shape arolium
Large
compound
eyes
Triangular and
moveable
head
Short
sclerotized
forewings☆
Large hindwings
with distinct
venation and a
pterostigma
Wipfler et al., 2019
Palaeoentomology
3. 1.2 Main genera and geological deposits Mid-Cretaceous Kachin amber
Lower Cretaceous Crato Formation Middle Eocene Green River Formation
Alienopterus Caputoraptor Meilia Teyia Alienopterella
Vcelesvab
Apiblatta Grant Chimaeroblattina
1. Introduction
4. 1.3 Controversial points
Bai et al., 2018
Current Biology
Functions of the cephalo-
thoracic scissors?
Peter et al., 2018
Biologia
Wasp mimicry?
Hinkelman et al., 2019
Biologia
Ant mimicry?
Sendi et al., 2020
Biologia
Pollination?
1. Introduction
5. 2. Ant mimicry of alienopterid nymphs
Ant
treehopper
Alydidae bug
Ant
Mantis
cricket
Fly
Beetle
Ant
Spider Spider
Extant ant mimics
More than 2000 living species of
insects and spiders, representing 11
different arthropod orders, have an
altered morphology and behaviour to
resemble ants. This phenomenon
called myrmecomorphy (ant mimicry)
has evolved at least 30 times among
arthropods.
6. Wunderlich, 2000
Eocene Baltic amber
Ant-mimicking Corinnidae
Grimaldi and Engel, 2005
Miocene Dominican amber
Ant-mimicking cerambycid beetle
with Camponotus ant
Fossil records of ant mimicry
2. Ant mimicry of alienopterid nymphs
7. sphecomyrmine
ants
Alienopterid
nymphs
the ratios of the antennae and
legs to the body length are
approximately the same as in
some sphecomyrmine ants
the strongly
constricted anterior
abdomen resulting in a
structure similar to a
petiole of ants
The body length and
overall morphology are
almost the same
2. Ant mimicry of alienopterid nymphs
extinct sphecomyrmine
ants lack geniculate
antennae, which are
the same as the
antennae of
alienopterids
Alienopterid
nymphs
sphecomyrmine
ants
8. We performed a quantitative analysis to evaluate the similarity among three alienopterid
nymphs (morphotypes I, II and III), sphecomyrmine ant workers, a “roachoid” (representing
extinct dictyopterans), four cockroaches and a mantis, and results shows that the shape of
specimens of morphotype I is closest to that of an ant.
2. Ant mimicry of alienopterid nymphs
9. Significance:
1. This is the earliest record of ant mimicry (myrmecomorphy), which is contemporary with the
earliest ants and their eusociality, and with the earliest evidence of myrmecophily, extending the
geological range of myrmecomorphy by approximately 50 million years.
2. Reflecting that ants had already played an important role in the tropical Cretaceous terrestrial
ecosystems.
3. Providing new evidence for life style of alienopterid nymphs.
2. Ant mimicry of alienopterid nymphs
10. 3. Pollination of alienopterid nymphs
This is the sixth (after Coleoptera,
Diptera, Mecoptera, Neuroptera,
Thysanoptera) insect pollinator
lineage with Mesozoic gymnosperm
associations and the only known
fossil example from the
Polyneoptera, one of the major
lineages of winged insects.
Dictyopterans (including cockroaches,
mantis and termites) were more
diverse and abundant in the late
Paleozoic and early Mesozoic. Our
discovery suggests that some ancient
dictyopterans may be among the
earliest pollinators and perhaps
played an important and hitherto
undervalued role in pollination
ecology.
11. The order Aethiocarenodea was erected using an isolated specimen from mid-
Cretaceous Kachin amber in 2017, but its morphological characters indicate that it is
highly likely a nymph of Alienopteridae, belonging to the genus Teyia Vršanský,
Mlynskýet Wang, 2018.
Order Aethiocarenodea
Poinar & Brown, 2017
4. Systematic position of Aethiocarenodea
Triangular
head
Constricted
abdomen
Elongate
body
Alienopterid
nymphs
Teyia Vršanský,
Mlynskýet Wang, 2018
12. 5. Life style of alienopterid adults
Visually
oriented
predators
Reducing weight,
protect the
metathoracic wing
articulation
Dipterism enabled them to
move efficiently in narrow
spaces between twigs and
leaves of dense vegetation
Specialized tarsal
and pretarsal
attachment devices
enabled species of
Alienopteridae to
move efficiently on
plant surfaces
13. 1. The constricted abdomen, body size and shape, and wing morphology of Teyia is
similar to some large aculeate Hymenoptera from the same deposit, especially ampulicid
wasps, indicating the presence of wasp mimicry.
2. The nymphs and adults of the Early Cretaceous alienopterid genus Teyia imitate
entirely different hymenopteran models, and therefore probably provide the first fossil
record of transformational mimicry.
3. Wasp mimicry of other alienopterid adults should be treated carefully (lacking key
character or lacking potential models).
Teyia
Ampulicidae
Scoliidae
Angaros-phecidae
Vcelesvab
Apiblatta Grant Chimaeroblattina
Mid-Cretaceous
Kachin amber
Lower Cretaceous
Crato Formation
Middle Eocene
Green River Formation
5. Life style of alienopterid adults
14. 1. The constricted abdomen, body size and shape, and wing morphology of Teyia is
similar to some large aculeate Hymenoptera from the same deposit, especially ampulicid
wasps, indicating the presence of wasp mimicry.
2. The nymphs and adults of the Early Cretaceous alienopterid genus Teyia imitate
entirely different hymenopteran models, and therefore probably provide the first fossil
record of transformational mimicry.
3. Wasp mimicry of other alienopterid adults should be treated carefully (lacking key
character or lacking potential models).
Teyia
Ampulicidae
Scoliidae
Angaros-phecidae
Vcelesvab
Apiblatta Grant Chimaeroblattina
Mid-Cretaceous
Kachin amber
Lower Cretaceous
Crato Formation
Middle Eocene
Green River Formation
5. Life style of alienopterid adults
15. 1. The constricted abdomen, body size and shape, and wing morphology of Teyia is
similar to some large aculeate Hymenoptera from the same deposit, especially ampulicid
wasps, indicating the presence of wasp mimicry.
2. The nymphs and adults of the Early Cretaceous alienopterid genus Teyia imitate
entirely different hymenopteran models, and therefore probably provide the first fossil
record of transformational mimicry.
3. Wasp mimicry of other alienopterid adults should be treated carefully (lacking key
character or lacking potential models).
Teyia
Ampulicidae
Scoliidae
Angaros-phecidae
Vcelesvab
Apiblatta Grant Chimaeroblattina
Mid-Cretaceous
Kachin amber
Lower Cretaceous
Crato Formation
Middle Eocene
Green River Formation
5. Life style of alienopterid adults
16. 1. The constricted abdomen, body size and shape, and wing morphology of Teyia is
similar to some large aculeate Hymenoptera from the same deposit, especially ampulicid
wasps, indicating the presence of wasp mimicry.
2. The nymphs and adults of the Early Cretaceous alienopterid genus Teyia imitate
entirely different hymenopteran models, and therefore probably provide the first fossil
record of transformational mimicry.
3. Wasp mimicry of other alienopterid adults should be treated carefully (lacking key
character or lacking potential models).
Teyia
Ampulicidae
Scoliidae
Angaros-phecidae
Vcelesvab
Apiblatta Grant Chimaeroblattina
Mid-Cretaceous
Kachin amber
Lower Cretaceous
Crato Formation
Middle Eocene
Green River Formation
5. Life style of alienopterid adults
17. 6. Phylogenetic relationship, distribution and
evolutionary history of Alienopteridae
1. Alienopteridae and Umenocoleidae
are sister groups, and they should be
combined in a clade Alienoptera, and
placed as sister to crown-Mantodea.
2. Our phylogenetic analyses and its
global distribution indicate that
Alienoptera probably arose prior to the
J/K boundary and their common
ancestor may have occurred in the
Late Jurassic.
3. Alienoptera diversified in the Early Cretaceous, but
rapidly declined after this epoch, which might be the
results of the Angiosperm Revolution.
4. Alienoptera is an unsuccessful evolutionary
experiment of Dictyoptera.