The Trachypachidae (sometimes known as false ground beetles) are a
family
Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of
beetle
Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
s that generally resemble small
ground beetle
Ground beetles are a large, cosmopolitan distribution, cosmopolitan family (biology), family of beetles, the Carabidae, with more than 40,000 species worldwide, around 2,000 of which are found in North America and 2,700 in Europe. As of 2015, it ...
s, but that are distinguished by the large
coxae of their rearmost legs. There are only six known extant species in the family, with four species of ''
Trachypachus'' found in northern
Eurasia
Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
and northern
North America
North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, and two species of ''
Systolosoma'' in
Chile
Chile, officially the Republic of Chile, is a country in western South America. It is the southernmost country in the world and the closest to Antarctica, stretching along a narrow strip of land between the Andes, Andes Mountains and the Paci ...
and Argentina. They were much more diverse in the past, with dozens of described species from the
Mesozoic
The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
.
Their habits are similar to those of the ground beetles; they are usually found in the
leaf litter
Plant litter (also leaf litter, tree litter, soil litter, litterfall, or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that has fallen to the ground. This detritus or dead organic material and its constituen ...
of
conifer
Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
forests, and are thought to be mostly predators and scavengers, active during the day (
diurnal).
According to entomologist
George Ball, "the most interesting thing about this small family is its uncertain phylogenetic relationships"; it has previously been classified as a subfamily of
Carabidae
Ground beetles are a large, cosmopolitan family of beetles, the Carabidae, with more than 40,000 species worldwide, around 2,000 of which are found in North America and 2,700 in Europe. As of 2015, it is one of the 10 most species-rich animal ...
, and as a family been placed in several different positions relative to other beetle families.
[ George E. Ball, "Trachypachidae", in Ross H. Arnett, Jr. and Michael C. Thomas, '']American Beetles
''American Beetles'' is a comprehensive description of the beetles of North America north of the tropical area of Mexico. It was started by Ross H. Arnett, Jr. as an update of his classic ''The Beetles of the United States''; along with Michael ...
'' (CRC Press, 2001), vol. 1 Recent molecular phylogenies have recovered as the sister group to the clade Carabidae+
Cicindelidae
Tiger beetles are a family of beetles, Cicindelidae, known for their aggressive predatory habits and running speed. The fastest known species of tiger beetle, '' Rivacindela hudsoni'', can run at a speed of , or about 125 body lengths per second. ...
as part of the
Adephaga
The Adephaga (from Greek ἀδηφάγος, ''adephagos'', "gluttonous") are a suborder of beetles, and with more than 40,000 recorded species in 10 families, the second-largest of the four beetle suborders. Members of this suborder are collecti ...
subgroup Geadephaga.
The earliest fossils known of this family are of the genera ''
Petrodromeus,'' ''
Permunda'' and ''
Apermunda'' from the
Permian
The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...
-
Triassic
The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
boundary of
Russia
Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
around 252 million years ago.
Fossil genera of the family are classified in the subfamily Eodromeinae, while the two modern genera are placed in the subfamily Trachypachinae. The two subfamilies are distinguished by the shape and position of the
sclerite
A sclerite (Greek language, Greek , ', meaning "hardness, hard") is a hardened body part. In various branches of biology the term is applied to various structures, but not as a rule to vertebrate anatomical features such as bones and teeth. Instea ...
s that are jointed to the mesocoxae (the section where the middle pair of legs joins the body).
Species
*Subfamily Trachypachinae
** ''
Systolosoma'' Chile, Argentina
*** ''
Systolosoma breve''
*** ''
Systolosoma lateritium''
** ''
Trachypachus''
Holarctic
The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical reg ...
*** ''
Trachypachus holmbergi''
*** ''
Trachypachus gibbsii''
*** ''
Trachypachus zetterstedtii''
*** ''
Trachypachus slevini''
*Subfamily †Eodromeinae
*After revision in Kirejtshuk et Ansorge, 2022.
**†''
Coreoeicos'' Bode, 1953
Posidonia Shale
The Posidonia Shale (, also called Schistes Bitumineux in Luxembourg) geologically known as the Sachrang Formation, is an Early Jurassic (Early to Late Toarcian) geological formation in Germany, northern Switzerland, northwestern Austria, souther ...
, Germany Early Jurassic (
Toarcian
The Toarcian is, in the International Commission on Stratigraphy, ICS' geologic timescale, an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic. It spans the time between 184.2 Megaannum, Ma (million ...
)
**†''
Denudirabus'' Ren, 1995 China, Early Cretaceous (Aptian)
**†''
Dolichorabus'' Ponomarenko, 1985
Ichetuy Formation, Russia, Late Jurassic
**†''
Dundorabus'' Ponomarenko, 1989
Ulaan-Ereg Formation Mongolia, Late Jurassic (
Tithonian
In the geological timescale, the Tithonian is the latest age (geology), age of the Late Jurassic Epoch and the uppermost stage (stratigraphy), stage of the Upper Jurassic Series. It spans the time between 149.2 ±0.7 annum, Ma and 143.1 ±0.6 (mi ...
)
**†''
Eodromeites'' Ponomarenko, 1985 (2 species) Russia, Upper Jurassic
**†''
Eodromeus'' Ponomarenko, 1977 (10+ species) China Russia Mongolia, Middle Jurassic-Early Cretaceous
**†''
Liassodites'' Kirejtshuk et Ansorge, 2022
Green Series, Germany Early Jurassic (Toarcian)
**†''
Evertus'' Ponomarenko, 1986 (= ''Fortiseode'' Jia, Liang, Chang et Ren, 2011) (3 species) Mongolia,
Yixian Formation
The Yixian Formation (; formerly Romanization of Chinese, transcribed as Yihsien Formation or Yixiang Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the Barremian stage of the Early Cretaceous. I ...
, China, Early Cretaceous (
Aptian
The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
)
**†''
Karadromeus'' Ponomarenko, 1977 (6+ species) Kazakhstan, Mongolia, Russia, China?, Middle Jurassic-Early Cretaceous
**†''
Karatoma'' Ponomarenko, 1977 (2 species)
Karabastau Formation Kazakhstan, Middle-Late Jurassic,
Glushkovo Formation, Russia, Upper Jurassic (Tithonian)
**†''
Necronectulus'' Ponomarenko, 1977
Tologoi Formation, Kazakhstan, Upper Triassic (
Carnian
The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227.3 ...
-
Norian
The Norian is a division of the Triassic geological period, Period. It has the rank of an age (geology), age (geochronology) or stage (stratigraphy), stage (chronostratigraphy). It lasted from ~227.3 to Mya (unit), million years ago. It was prec ...
)
Jinju Formation, South Korea Early Cretaceous (
Albian
The Albian is both an age (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...
)
**†''
Novunda'' Kirejtshuk et Ansorge, 2022 (= ''Unda'' Ponomarenko, 1977 preoccupied genus) (5 species)
Daohugou, China, Middle Jurassic, Glushkovo Formation, Russia, Upper Jurassic
**†''
Platycoxa'' Ponomarenko, 1977 Kyrgyzstan, Lower Jurassic
**†''
Prosynactus'' Bode, 1953 Posidonia Shale, Germany Early Jurassic (Toarcian)
**†''
Psacodromeus'' Ponomarenko, 1977 Kazakhstan, Mongolia, Middle/Late Jurassic-Early Cretaceous, Jinju Formation, South Korea Early Cretaceous (Albian)
**†''
Sinodromeus'' Wang, Zhang et Ponomarenko, 2012, Yixian Formation, China, Early Cretaceous (Aptian)
** †''
Sogdodromeus'' Ponomarenko, 1977
Madygen Formation, Kyrgyzstan, Upper Triassic (
Carnian
The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227.3 ...
)
*** †''Sogdodromeus altus''
**†''
Xishanocarabus'' Hong, 1984
Lushangfen Formation, China, Early Cretaceous (Aptian)
** †''
Petrodromeus'' Ponomarenko et Volkov, 2013 Kemerovo, Russia, Late Permian (Changhsingian)
*** †''Petrodromeus asiaticus'' (=''Ademosynoides asiaticus''; Martynov, 1936)
** †''
Apermunda'' Kirejtshuk et Ansorge, 2022, Kemerovo, Russia, Late Permian (Changhsingian)
*** †''Apermunda minor'' (=''Petrodromeus minor'')
** †''
Permunda'' Ponomarenko et Volkov, 2013 Kemerovo, Russia, Late Permian (Changhsingian)
*** †''Permunda nana''
References
{{Taxonbar, from=Q150582
Adephaga
Beetle families
Beetles of North America