Conference: Identifying the next challenges in ichneumonid systematics and evolutionary ecology Basel, Switzerland, 24-28 June 2019
Patchy, challenging and enigmatic: the ichneumonid fossil record
Seraina Klopfstein
Naturhistorisches Museum Basel, BIO, Basel, Switzerland. [email protected]
Fossils not only inform us about character evolution, historical biogeography and the impact of mass extinctions, they are also the most important source of information for dating molecular trees. Because fossilization rates are highly uneven across both the temporal, spatial and phylogenetic scales, the fossil record is highly skewed and only gives us a very patchy picture of the past. The fossil record of ichneumonids furthermore suffers from severe neglect by the scientific community – only 275 fossil species have been described to date, 75% of which by just eight authors, and hundreds of fossil specimens are still awaiting detailed study in paleontological collections. Many of the described fossil species stem from the time before Townes published his seminal volumes that overturned ichneumonid classification. They are in need of revision in the light of new insights into character evolution and phylogenetics in this family. The position of most of the Mesozoic fossils is also in need of review, and special focus should be put on the interpretation of diagnostic characteristics of some supposedly extinct subfamilies and their distinction from extant groups. Experiments that aim to improve our understanding of the fossilization process of ichneumonids could include controlled rotting and squashing, which might shed light on past misinterpretations of the morphology of ichneumonid fossils. Current work on the Cenozoic fossil record tackles the very rich Messel fossil record in Germany (about 220 specimens, about 48 Mya) and the virtually unstudied Fur Formation in Denmark (about 80 specimens, about 56 Mya). Many more discoveries await and will provide a more complete picture of ichneumonid evolution through time.
PATCHY, CHALLENGING AND ENIGMATIC: THE ICHNEUMONID FOSSIL RECORD
Seraina Klopfstein Please do not reproduce or distribute any of the fossil photographs in this talk! (many of them are under copyright by the respective institutions…)
Disclaimer: no part of this document constitutes a nomenclatorial act in the sense of the international code of zoological nomenclature. oldest clear Pimplinae and Rhyssinae (48 Ma) oldest ichneumonid (120 Ma)
oldest Hymenoptera (235 Ma)
oldest insect (396 Ma) WHY CARE?
• fossils inform about character evolution
• impact of mass extinctions
• biogeography
• co-evolution
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 patchy – gaps in (our knowledge of) the fossil record
challenging – the difficulties of fossil interpretation
enigmatic – solving fossil riddles to understand ichneumonid evolution
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 PATCHY watching a movie, but just in glimpses of a few seconds at a time
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 PATCHY
• 275 species described (1% of extant fauna) • by just a few authors • early authors: Cockerell, Brues, Theobald • recent authors: Kasparyan, Khalaim, Kopylov, Spasojevic, Zhang • undescribed diversity is huge!
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 PATCHY Number of described fossil species of ~ichneumonids
Florissant (USA), Baltic amber Isle of Wight (UK) (EU) Khurlit (CN) Olskaya (RU) diverse (Aix, Yixan (CN) Rott, etc.) Green River (USA) Baissa (RU)
million years ago
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 MESOZOIC ICHNEUMONIDS recent subfamilies • 3 Tryphoninae (Townes 1973) --> moved to Labenopimplinae (Kopylov 2012) • 1 Labeninae (McKellar et al. 2013) --> placement uncertain (authors, own observ.) extinct subfamilies • Tanychorinae • Paleoichneumoninae • Labenopimplinae
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 † TANYCHORINAE Rasnitsyn
Tanychora petiolata braconid? Russia, Baissa, lower Cretaceous (>127 Ma) or ichneumonid stem lineage? or (braconid + ichneumonid) stem lineage?
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 † PALEOICHNEUMONINAE Kopylov
• notauli converging • areolet wide pentagonal • T1 short & wide • 2m-cu «almost always» with one bulla
Dischysma maculata Bon Tsagan, Khurilt seq., ~100 Ma Kopylov 2010 Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 † LABENOPIMPLINAE Kopylov
Labenopimpla kasparyani Russia, Olskaya formation, >93 Ma Kopylov 2010
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 † LABENOPIMPLINAE Kopylov
• highly diverse
• notauli absent or parallel • areolet quadrate or pentagonal, often wide • 2 bullae in 2m-cu • propodeum with 13-15 areas Labenopimpla kasparyani Russia, Olskaya formation, >93 Ma Kopylov 2010
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 PATCHY…
Cenozoic Ichneumonids we have barely scratched the surface… 6 emails 30 kg of rocks and > 300 photos of undescribed ichneumonids
more later…
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 CHALLENGING
«taphonomy» = science of how things fossilize • how do ichneumonids fossilize? apomorphy, plesiomorphy or homoplasy? • host-related character complexes -> often unreliable… • scarcity of good characters for many extant subfamilies crown group or stem group • how old are the recent subfamilies / genera? is it even possible that some fossils belong to recent groups?
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019
undescribed species, Messel pit, 48 Ma, ©Senkenberg
INTERPRETATION?
• wide areolet -> Labeninae or Labenopimplinae? or what else? • rather wide T1 -> Labenopimplinae • 2m-cu bowed outwards -> Labenopimplinae
• Labenopimplinae: • known only from Mesozoic • definition somewhat vague • «propodeum with 13-15 areas»
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 TAPHONOMY OF PROPODEUM
undescribed species, Messel pit, 48 Ma, ©Senkenberg Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 TAPHONOMY OF PROPODEUM
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 spiracle
metapleuron half of metasternum!
mesosternal scrobe REVIEW STATUS OF LABENOPIMPLINAE
• refine definition of the subfamily • diagnose genera individually from extant groups • relation to Paleoichneumoninae • phylogenetic placement
Labenopimpla kasparyani Russia, Olskaya formation, >93 Ma Kopylov 2010
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 CHALLENGING
«taphonomy» = science of how things fossilize • let them rot! --> ichneumonid taphonomy project? apomorphy, plesiomorphy or homoplasy? • morphological phylogenetics --> workshop! crown group or stem group • dated phylogeny of Ichneumonidae
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019
ENIGMATIC how can we lessen the gaps in the ichneumonid fossil record?
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 MESSEL FOSSILS
~48 Ma, crater lake 216 specimens how many species? cherry-picking? --> eventually summarize entire fauna
Spasojevic et al. 2018 PLoS ONE EXCEPTIONAL PRESERVATION
- habitus --> Pimplinae? - lobed claws
- oldest unequivocal representative of Pimplinae - genus: Scambus (s.l.) FUR FORMATION, DENMARK
• sea deposit (!) • ~56 Ma (Eocene – Paleocene) • two layers rich of insects – and devoid of fishes
• 80 specimens –> 25 species?
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 CONSPICUOUS COLOUR PATTERN, ETC…
--> Xanthopimpla?
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019
RHOMBIC AREOLET
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 † PHERHOMBINAE Kasparyan
• 1 genus, 3 species • all from Baltic amber (~44 Ma) • all male • elongate T1
Pherhombus antennalis Baltic amber, ~44 Ma Kasparyan 1988 RHOMBIC AREOLET
--> Pherhombus? --> subfamily 12 Myr older
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019
HOW TO COPE WITH UNCERTAINTY? most fossils can NOT be placed with high confidence • communicate uncertainty properly (open taxonomy) • report tentative placement with «?» • morphological phylogenetic analysis to identify alternative placements
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 «ROGUE PLOTS» TO ILLUSTRATE FOSSIL PLACEMENT
Polyhelictes bipolarus Spasojevic et al. 2018 - > Bayesian morphological phylogeny --> rogue plot
--> R package to produce RoguePlots on Github Klopfstein & Spasojevic 2018 PLoS ONE
HELP MAKE IT LESS PATCHY! revise fossil taxa along with extant ones do your own fossil study fossil ichneumonid email list • sign up to receive emails about new fossil species! • collaborate on interpreting fossils • potential placements in recent subfamilies & genera • discussion of character evidence
Seraina Klopfstein – ICHNEUMONID MEETING, Basel 2019 THANKS GO TO
Tamara Spasojevic
Sonja Wedmann
René Sylvestersen
& you for your Dmitry Kopylov attention!