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Cretaceous Achilidae
from Myanmar
- a strange medley of basal
and crown tribe-level groups
Alicja Magdalena Brysz
Laboratory of Evolutionary Entomology and
Museum of Amber Inclusions
University of Gdańsk, Poland
Achilidae definition and ecology
Achilidae is a small (~160 genera) planthopper (Hemiptera, Fulgoromorpha) family
distributed worldwide in all terrestrial habitats and climates of the Earth. Nymphs
are supposedly closely tied to wood decomposing fungi and cryptic, while adults
feed on phloem of many plant families, both angio- and gymnosperms.
Unfortunately not much more can be said about their general ecology, as it is a
rarely researched group.
Achilidae morphology is a much better known subject, although, as with many
other ’basal’ planthoppers, some of its diagnostic characters can appear in other,
closely related families (i.e. Derbidae, Kinnaridae, Cixiidae).
Nonetheless achilids are defined by (with omission of genital details):
- overlapping apical end of tegmina (first pair of wings)
- open clavus (venation character)
- second tarsomere (foot segment) with row of spines
- oval pedicel (second antennal segment)
Achilidae presence on Earth by Bourgoin 2018
Achilidae definition and ecology
Plectoderes scapularis
Achilus flammeus by David MulesApateson albomaculatum by myrockytop
Cixidia lapponica
Ptychoptilum sp.
Inter-family systematics
Internal Achilidae systematics is quite complicated for a group of this size. The whole family is
divided into 3 supertribes: Achilites, Apatesonites and Myconites, with some taxa (including 2
tribes) placed as incertae sedis within the family.
In total there are 13 tribes, of which two are completely extinct, with some more under
description, including one from Burmese (Myanmar) amber, here (as needed) named trib. nov.
One of the most often present in fossil record tribes – Achilini, is further divided into:
Achilina, Cixidiina and Elidipterina subtribes and is the exception in this matter.
Achilites Stål, 1866 Achilini Stål, 1866
Achilina Stål, 1866
Cixidiina Emeljanov, 1992
Elidipterina Fennah, 1950.
Achillini Cockerell, 1922
Apatesonites Metcalf, 1938 Apatesonini Metcalf, 1938
Ilvini Emeljanov, 1991
Seviini Emeljanov, 1991
Tropiphlepsiini Emeljanov, 1991
Myconites Fennah, 1950 Amphignomini Emeljanov, 1991
Mycarini Emeljanov, 1991
Myconini Fennah, 1950
Plectoderini Fennah, 1950
Rhotalini Fennah, 1950
Ptychoptilini† Emeljanov, 1990
Waghildini† Szwedo, 2006
Inter-family systematics
Tribe Number of genera / species recent extinct
Achilini 31 / 95 26 / 88 5 / 7
Achillini 3 / 5 3 / 5 -
Apatesonini 1 / 1 1 / 1 *
Ilvini + Seviini 1 / 1 + 1 / 7 1 / 1 + 1 / 7 -
Tropiphlepsiini 1 / 1 1 / 1 -
Amphignomini 1 / 1 1 / 1 *
Mycarini 5 / 6 5 / 6 -
Myconini 5 / 8 5 / 8 -
Plectoderini 99 / 348 99 / 348 *
Rhotalini 5 / 23 5 / 23 -
Ptychoptilini 2 / 3 - 2 / 3
Waghildini 1 / 1 - 1 / 1
Achilidae incertae sedis 6 / 16 - 4 / 5
*not described Myanmar fossils
Proposed Achilidae tribe relations
During decades of Achilidae research, to this day there was
only one proposed tree of relations in-between the tribes
created in 1990’s by Emeljanov. Unfortunately it does not
include any kind of fossil data.
Our own research suggest that the title of oldest Achilidae
tribe should belong to Plectoderini, while Rhotalini, which to
this day show no sign of any kind of fossil data are most
probably one of the youngest tribes, thus questioning its
placement in the definitely oldest Myconites supertribe.
Placement of other tribes seems to be generally accurate.
After Emeljanov 1991, 1992; Emeljanov & Fletcher 2004
Achilidae fossil trail starts in Lower Cretaceous , with appearance of Acixiites. It is the oldest
known achilid genus and, although its description remains dubious in some matters, its
ascription to this family is as sure as its is possible.
It is supposed that in late Lower Cretaceous occurred a sudden ’explosion’ of diversity of
Achilidae., which resulted in many already established tribes recorded in Upper Cretaceous,
the time of accumulation of Myanmar amber, such as: Plectoderini, Apatesonini,
Amphignomini and (probably) Ptychoptilini along with Waghildini.
Next such ’explosion’ occurred probably somewhere around the beginning of
the Eocene, which is reflected by even greater diversity present among inclusions of Baltic
amber (deposited in the middle of the Eocene).
Achilidae fossil data
Diagram covers only described
fossils with the exception of those
recognized from Myanmar amber
(marked red).
Scheme including taxa from other
sources (i.e. Baltic amber) can be
found in Brysz & Szwedo (2018).
Note that fossil data concerning
most of the closest relatives of
Achilidae are not currently known.
Achilidae fossil data
Plectoderini case
Plectoderini is the largest Achilidae tribe containing 62% of
all its genera and 68% of all species. Such proportions are
very unlikely, especially with high diversity rate of achilids.
Therefore we surmise that Plectoderini should be placed
under revision and most probably divided into further
tribes, and perhaps also subtribes.
Their morphology is in general very conservative, not having
changed much since the Cretaceous. It also seems that fossil
data from Myanmar amber will help in previously
mentioned revision, planned for the oncoming years.
Myanmar Plectoderini
One of the reasons that Myanmar Plectoderini are not
easy to work with is that for some reason those
inclusions are often embedded in dark and heavily
polluted resin, which in combination with small size of
those insects makes their analysis very time-consuming.
Fortunately it seems that most of their diagnostic
features are very close to those of modern Plectoderini.
‘Crown’ Achilidae groups
Despite Emeljanov’s opinion approved for a long time, it seems that groups placed in lower position in
his tree such as tribes Amphignomini, Mycarini and Rhotalini are probably placed correctly, but
branched from the other, elder groups; on the other hand, those tribes placed high on the same tree,
such as e.g. Apatesonini, having no fossil record at time, may be more ancient. Therefore finding in
Myanmar amber representatives carrying characteristics that without any doubt identified them as
Amphignoma relatives was such a shock. This suggests that even without fossil data Emeljanov had been
correct in at least some (if not most) matters concerning tribes placement in correlation with each
other.
Myanmar amber Apatesonini occurrence was not as big surprise, as there have been observed some
representatives of this tribe in Baltic amber (Eocene), nonetheless to this founding we did not expect for
this tribe to be as old as it seems to be. In combination with Cretaceous appearance of other
Apatesonites supertribe taxa (Amphignomini, Plectoderini) it seems that its position as oldest Achilidae
supertribes should be set.
Myanmar Amphignomini
Recent part of Amphignomini tribe
(and the only described) is composed
from one monotypic genus.
Morphological similarity of both of
those genera (as they cannot be
considered the same with current
views on the definition of genera or
species) is astounding and sets many
questions how such an old and small
taxa survived many difficult and
tumultuous climate changes without
any significant changes.
New Myanmar higher-level taxa
Apart from the representatives of already known tribes, Myanmar amber brought some more
surprises, that may have a huge impact on phylogeny and inner-family tribes relations.
One of those surprises is a new tribe belonging probably to Myconites, which corresponds
nicely with Cretaceous appearance of other tribes from this group. This tribe carries an
intriguing set of characteristics that place it very low on Myconites tribes relations tree, most
probably next to the oldest one - Plectoderini.
The second surprise comes in form of a new Fulgoromorpha family, very closely related to
Achilidae, which fills some gaps in phylogeny of the oldest portion of ’basal’ plenthoppers.
Both of these taxa will soon be announced in published form.
New extinct tribe from Myanmar amber
One could think that a new tribe from such an
old material (~100 Ma) will be sparsely
represented, but this tribe to this day is
recognised as comprising at least three genera,
each with more than one species.
Such abundance is not unheard of in one the
richest fossil data source that is Myanmar
amber, but still it is a nice change, not to be
force to work on only one specimen, as I am
sure all palaeontologists will agree.
Works on new Achilidae definition
All discoveries concerning Achilidae fossils in recent years only strengthened the necessity
for clear boundaries of the whole family and between other ’basal’ planthopper families,
which has always been a problem, as specialist can easily say which is which, but are unable to
name those differences in proper terms.
W hope that after finishing our work on fossil achilids a revised, if not new, definition of the
family Achilidae will emerge, and that it will set in motions similar changes in definitions for
other, more or less closely related groups.
I would like to thank the organisers of the conference for their work and all of You for the attention.
I also invite you to follow our work and to check some of the references:
• Bourgoin T. 2018. FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to
Fulgoromorpha. Version 8, updated 2018-11-20. http://hemiptera-databases.org/flow/
• Brysz A.M., Szwedo J. 2018. The fossil record of the planthopper family Achilidae, with particular reference to
those in Baltic amber (Hemiptera: Fulgoromorpha). Earth and Environmental Science Transactions of the Royal
Society of Edinburgh 107, 279-288. doi:10.1017/S175569101700041X
• Emeljanov A.F. 1991. To the problem of the limits and subdivisions of the family Achilidae (Homoptera, Cicadina).
Entomologicheskoe Obozrenie 70(2), 373-393.
• Emeljanov A.F. 1992. Description of tribes of the subfamily Achilinae (Homoptera: Achilidae) and revision of their
composition. Entomologicheskoe Obozrenie 71(3), 574-594.
• Emeljanov A.F., Fletcher M.J. 2004. Hemielissum evansi, a new genus and species of Breddiniolini (Hemiptera:
Fulgoromorpha), being the first Australian record of the tribe, with a discussion of the taxonomic position of the
Breddiniolini. Australian Journal of Entomology 43, 38-42.

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Brysz A. Cretaceous Achilidae from Myanmar paleovc

  • 1. Cretaceous Achilidae from Myanmar - a strange medley of basal and crown tribe-level groups Alicja Magdalena Brysz Laboratory of Evolutionary Entomology and Museum of Amber Inclusions University of Gdańsk, Poland
  • 2. Achilidae definition and ecology Achilidae is a small (~160 genera) planthopper (Hemiptera, Fulgoromorpha) family distributed worldwide in all terrestrial habitats and climates of the Earth. Nymphs are supposedly closely tied to wood decomposing fungi and cryptic, while adults feed on phloem of many plant families, both angio- and gymnosperms. Unfortunately not much more can be said about their general ecology, as it is a rarely researched group. Achilidae morphology is a much better known subject, although, as with many other ’basal’ planthoppers, some of its diagnostic characters can appear in other, closely related families (i.e. Derbidae, Kinnaridae, Cixiidae). Nonetheless achilids are defined by (with omission of genital details): - overlapping apical end of tegmina (first pair of wings) - open clavus (venation character) - second tarsomere (foot segment) with row of spines - oval pedicel (second antennal segment) Achilidae presence on Earth by Bourgoin 2018
  • 3. Achilidae definition and ecology Plectoderes scapularis Achilus flammeus by David MulesApateson albomaculatum by myrockytop Cixidia lapponica Ptychoptilum sp.
  • 4. Inter-family systematics Internal Achilidae systematics is quite complicated for a group of this size. The whole family is divided into 3 supertribes: Achilites, Apatesonites and Myconites, with some taxa (including 2 tribes) placed as incertae sedis within the family. In total there are 13 tribes, of which two are completely extinct, with some more under description, including one from Burmese (Myanmar) amber, here (as needed) named trib. nov. One of the most often present in fossil record tribes – Achilini, is further divided into: Achilina, Cixidiina and Elidipterina subtribes and is the exception in this matter.
  • 5. Achilites Stål, 1866 Achilini Stål, 1866 Achilina Stål, 1866 Cixidiina Emeljanov, 1992 Elidipterina Fennah, 1950. Achillini Cockerell, 1922 Apatesonites Metcalf, 1938 Apatesonini Metcalf, 1938 Ilvini Emeljanov, 1991 Seviini Emeljanov, 1991 Tropiphlepsiini Emeljanov, 1991 Myconites Fennah, 1950 Amphignomini Emeljanov, 1991 Mycarini Emeljanov, 1991 Myconini Fennah, 1950 Plectoderini Fennah, 1950 Rhotalini Fennah, 1950 Ptychoptilini† Emeljanov, 1990 Waghildini† Szwedo, 2006 Inter-family systematics
  • 6. Tribe Number of genera / species recent extinct Achilini 31 / 95 26 / 88 5 / 7 Achillini 3 / 5 3 / 5 - Apatesonini 1 / 1 1 / 1 * Ilvini + Seviini 1 / 1 + 1 / 7 1 / 1 + 1 / 7 - Tropiphlepsiini 1 / 1 1 / 1 - Amphignomini 1 / 1 1 / 1 * Mycarini 5 / 6 5 / 6 - Myconini 5 / 8 5 / 8 - Plectoderini 99 / 348 99 / 348 * Rhotalini 5 / 23 5 / 23 - Ptychoptilini 2 / 3 - 2 / 3 Waghildini 1 / 1 - 1 / 1 Achilidae incertae sedis 6 / 16 - 4 / 5 *not described Myanmar fossils
  • 7. Proposed Achilidae tribe relations During decades of Achilidae research, to this day there was only one proposed tree of relations in-between the tribes created in 1990’s by Emeljanov. Unfortunately it does not include any kind of fossil data. Our own research suggest that the title of oldest Achilidae tribe should belong to Plectoderini, while Rhotalini, which to this day show no sign of any kind of fossil data are most probably one of the youngest tribes, thus questioning its placement in the definitely oldest Myconites supertribe. Placement of other tribes seems to be generally accurate. After Emeljanov 1991, 1992; Emeljanov & Fletcher 2004
  • 8. Achilidae fossil trail starts in Lower Cretaceous , with appearance of Acixiites. It is the oldest known achilid genus and, although its description remains dubious in some matters, its ascription to this family is as sure as its is possible. It is supposed that in late Lower Cretaceous occurred a sudden ’explosion’ of diversity of Achilidae., which resulted in many already established tribes recorded in Upper Cretaceous, the time of accumulation of Myanmar amber, such as: Plectoderini, Apatesonini, Amphignomini and (probably) Ptychoptilini along with Waghildini. Next such ’explosion’ occurred probably somewhere around the beginning of the Eocene, which is reflected by even greater diversity present among inclusions of Baltic amber (deposited in the middle of the Eocene). Achilidae fossil data
  • 9. Diagram covers only described fossils with the exception of those recognized from Myanmar amber (marked red). Scheme including taxa from other sources (i.e. Baltic amber) can be found in Brysz & Szwedo (2018). Note that fossil data concerning most of the closest relatives of Achilidae are not currently known. Achilidae fossil data
  • 10. Plectoderini case Plectoderini is the largest Achilidae tribe containing 62% of all its genera and 68% of all species. Such proportions are very unlikely, especially with high diversity rate of achilids. Therefore we surmise that Plectoderini should be placed under revision and most probably divided into further tribes, and perhaps also subtribes. Their morphology is in general very conservative, not having changed much since the Cretaceous. It also seems that fossil data from Myanmar amber will help in previously mentioned revision, planned for the oncoming years.
  • 11. Myanmar Plectoderini One of the reasons that Myanmar Plectoderini are not easy to work with is that for some reason those inclusions are often embedded in dark and heavily polluted resin, which in combination with small size of those insects makes their analysis very time-consuming. Fortunately it seems that most of their diagnostic features are very close to those of modern Plectoderini.
  • 12. ‘Crown’ Achilidae groups Despite Emeljanov’s opinion approved for a long time, it seems that groups placed in lower position in his tree such as tribes Amphignomini, Mycarini and Rhotalini are probably placed correctly, but branched from the other, elder groups; on the other hand, those tribes placed high on the same tree, such as e.g. Apatesonini, having no fossil record at time, may be more ancient. Therefore finding in Myanmar amber representatives carrying characteristics that without any doubt identified them as Amphignoma relatives was such a shock. This suggests that even without fossil data Emeljanov had been correct in at least some (if not most) matters concerning tribes placement in correlation with each other. Myanmar amber Apatesonini occurrence was not as big surprise, as there have been observed some representatives of this tribe in Baltic amber (Eocene), nonetheless to this founding we did not expect for this tribe to be as old as it seems to be. In combination with Cretaceous appearance of other Apatesonites supertribe taxa (Amphignomini, Plectoderini) it seems that its position as oldest Achilidae supertribes should be set.
  • 13. Myanmar Amphignomini Recent part of Amphignomini tribe (and the only described) is composed from one monotypic genus. Morphological similarity of both of those genera (as they cannot be considered the same with current views on the definition of genera or species) is astounding and sets many questions how such an old and small taxa survived many difficult and tumultuous climate changes without any significant changes.
  • 14. New Myanmar higher-level taxa Apart from the representatives of already known tribes, Myanmar amber brought some more surprises, that may have a huge impact on phylogeny and inner-family tribes relations. One of those surprises is a new tribe belonging probably to Myconites, which corresponds nicely with Cretaceous appearance of other tribes from this group. This tribe carries an intriguing set of characteristics that place it very low on Myconites tribes relations tree, most probably next to the oldest one - Plectoderini. The second surprise comes in form of a new Fulgoromorpha family, very closely related to Achilidae, which fills some gaps in phylogeny of the oldest portion of ’basal’ plenthoppers. Both of these taxa will soon be announced in published form.
  • 15. New extinct tribe from Myanmar amber One could think that a new tribe from such an old material (~100 Ma) will be sparsely represented, but this tribe to this day is recognised as comprising at least three genera, each with more than one species. Such abundance is not unheard of in one the richest fossil data source that is Myanmar amber, but still it is a nice change, not to be force to work on only one specimen, as I am sure all palaeontologists will agree.
  • 16. Works on new Achilidae definition All discoveries concerning Achilidae fossils in recent years only strengthened the necessity for clear boundaries of the whole family and between other ’basal’ planthopper families, which has always been a problem, as specialist can easily say which is which, but are unable to name those differences in proper terms. W hope that after finishing our work on fossil achilids a revised, if not new, definition of the family Achilidae will emerge, and that it will set in motions similar changes in definitions for other, more or less closely related groups.
  • 17. I would like to thank the organisers of the conference for their work and all of You for the attention. I also invite you to follow our work and to check some of the references: • Bourgoin T. 2018. FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to Fulgoromorpha. Version 8, updated 2018-11-20. http://hemiptera-databases.org/flow/ • Brysz A.M., Szwedo J. 2018. The fossil record of the planthopper family Achilidae, with particular reference to those in Baltic amber (Hemiptera: Fulgoromorpha). Earth and Environmental Science Transactions of the Royal Society of Edinburgh 107, 279-288. doi:10.1017/S175569101700041X • Emeljanov A.F. 1991. To the problem of the limits and subdivisions of the family Achilidae (Homoptera, Cicadina). Entomologicheskoe Obozrenie 70(2), 373-393. • Emeljanov A.F. 1992. Description of tribes of the subfamily Achilinae (Homoptera: Achilidae) and revision of their composition. Entomologicheskoe Obozrenie 71(3), 574-594. • Emeljanov A.F., Fletcher M.J. 2004. Hemielissum evansi, a new genus and species of Breddiniolini (Hemiptera: Fulgoromorpha), being the first Australian record of the tribe, with a discussion of the taxonomic position of the Breddiniolini. Australian Journal of Entomology 43, 38-42.