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TABLE 1. List of the fossil and subfossil Anthicidae and Ischaliidae records.

Taxon References Fossil Type and/or Locality Epoch Age, Ma
Anthicidae Latreille, 1819 Kirejtshuk and Azar, 2013; Peris et al., 2016 Lebanese amber Cretaceous 130-125.5
Rasnitsyn and Ross, 2000 Burmese amber Cretaceous 99.7-94.3
Kirejtshuk and Nel, 2013 Oise amber: Oise Department (France) Eocene 55.8-48.6
Perkovsky et al., 2007, 2010 Rovno amber Eocene 37.2-33.9
Berendt, 1845; Menge, 1856: as “Notoxida”; Helm, 1886; Klebs, 1889, 1910; Handlirsch, 1907; Bachofen-Echt, 1949; Abdullah, 1964; Spahr, 1981; Hieke and Pietrzeniuk, 1984; Ritzkowski, 1990; Kulicka and Ślipińsk, 1996; Weitschat and Wichard, 2002 Baltic amber Eocene 37.2-33.9
Poinar, 1992 Dominican amber Miocene 20.4-13.6
Poinar, 1992 Mexican amber Miocene 20-15
Schawaller, 1986 lithified, carbonaceous lime mudstone; Randeck Maar formation (Germany) Miocene 16-13.7
Bidashko et al., 1995 unlithified lignite and unlithified claystone: Astrakhan (Russian Federation) Holocene 0.1-0.0
Anthicinae Latreille, 1819  Kirejtshuk et al., 2019  Isle of Wight Insect Limestone (U.K.) Late Eocene  37.8-33.9
Anthicini Latreille, 1819        
Amblyderus sp. Klebs, 1910; Bachofen-Echt, 1949; Abdullah, 1964; Larsson, 1978; Spahr, 1981; Poinar, 1992; Kirejtshuk, 2002-2013 Baltic amber Eocene 37.2-33.9
Anthicus neli Arillo et Ortuño, 1997 [MCNA] [(? = Omonadus formicarius formicarius (Goeze, 1777)]. None of features provided in the original description of this taxon are good enough to separate it from extant cosmopolitan Omonadus formicarius formicarius (Goeze, 1777). Considering the fact Hörnschemeyner et al. (2010) pointed on, in certain cases it is not possible to find any morphological features that distinguish the fossils from the extant species on base of morphological characters. We had not examined Anthicus neli holotype, therefore it is not impossible this taxon is conspecific with extant O. formicarius. Authors compared (Arillo and Ortuño, 1997: 295) their new species with Omonadus floralis (Linnaeus, 1758) which is not as similar to it as O. formicarius formicarius. Arillo and Ortuño 1997; Kirejtshuk 2002-2013 lacustrine claystone: Izarra (Alava, Spain) Miocene 23-16
Anthicus oustaleti Telnov, 1999 (= melancholicus Oustalet, 1874 nec LaFerté-Sénectère, 1849) [MHNM] Oustalet, 1874; Telnov, 1999; Kirejtshuk, 2002-2013 lacustrine shale: Aix-en-Provence (France) Eocene 28.4-23
Anthicus sepultulus Cockerell, 1926 [BMNH] Cockerell, 1926, 1936; Kirejtshuk, 2002-2013 lacustrine: Maíz Gordo Formation (Argentina) Palaeocene 66-55.8
Anthicus sp. Hope, 1836; Berendt, 1845; Giebel, 1852, 1856a, 1856b; Motschulsky, 1856; Menge, 1856; Scudder, 1885, 1886, 1891; Handlirsch, 1907; Klebs, 1910; Bachofen-Echt, 1949; Abdullah, 1964; Larsson, 1978; Spahr, 1981; Poinar, 1992; Kirejtshuk, 2002-2013 Baltic amber Eocene 37.2-33.9
Anthicus sp. Kiselev and Nazarov, 2009 Oler Formation (Russian Federation) Pleistocene 2.58-0.8
Anthicus sp. Green et al., 2006 lacustrine; Hackney (United Kingdom) Pleistocene 0.8 - 0.1
Anthicus sp. Coope et al., 1961 lacustrine; Worcestershire (United Kingdom) Holocene 0.1-0.0
Anthicus sp. Kiselev and Nazarov, 2009 lacustrine; Russian Federation Holocene 0.1 - 0.0
Cyclodinus efficax Kirejtshuk, 2019 in Kirejtshuk et al., 2019 Kirejtshuk et al., 2019 Isle of Wight Insect Limestone (U.K.) Late Eocene 37.8-33.9
Nitorus succinius sp. nov. [CCHH] present paper Baltic amber Eocene 37.2-33.9
Endomiini Bonadona, 1958        
Endomia sp. (= Ochthenomus Dejean, 1834) Klebs, 1910: as Ochthenomus; Bachofen-Echt, 1949; Abdullah, 1964; Larsson, 1978; Spahr, 1981; Poinar, 1992; Kirejtshuk, 2002-2013 Baltic amber Eocene 37.2-33.9
Eurygeniinae LeConte, 1862        
Eurygeniini LeConte, 1862        
Oisegenius Kirejtshuk et Nel, 2008 Kirejtshuk and Nel, 2008, 2013; Kirejtshuk, 2002-2013; Oise amber: Oise department (France) Eocene 55.8-48.6
Oisegenius antiquus Kirejtshuk et Nel, 2008 [MNHN] Kirejtshuk and Nel, 2008; Kirejtshuk, 2002-2013; Brasero , 2009; Nel and Brasero, 2010 Oise amber: Oise department (France) Eocene 55.8-48.6
Lemodinae Lawrence et Britton, 1991        
Lemodicarmenia olmedoae Molino-Olmedo, 2017 [CMOM]. This species has general appearance of Tomoderinae. We had not studied the type and therefore follow the original placement. Molino-Olmedo, 2017 Burmese amber Cretaceous 99.7-94.3
Macratriinae LeConte, 1862 present paper Burmese amber Cretaceous 99.7-94.3
Camelomorphini Kirejtshuk, Azar et Telnov in Kirejtshuk and Azar, 2008. Most of authors who quoted this taxon ignored the fact the tribe Camelomorphini, genus Camelomorpha and species Camelomorpha longicervix was authored by Kirejtshuk, Azar et Telnov in Kirejtshuk and Azar, 2008 and not by Kirejtshuk et Azar Kirejtshuk and Azar, 2008, 2013; Kirejtshuk, 2002-2013; Azar et al., 2010; Telnov, 2012b Lebanese amber Cretaceous 130-125.5
Camelomorpha Kirejtshuk, Azar et Telnov in Kirejtshuk and Azar, 2008 Kirejtshuk and Azar, 2008, 2013; Kirejtshuk, 2002-2013; Azar et al., 2010; Telnov, 2012b Lebanese amber Cretaceous 130-125.5
Camelomorpha longicervix Kirejtshuk, Azar et Telnov in Kirejtshuk and Azar, 2008 [MNHN] Kirejtshuk and Azar, 2008, 2013; Kirejtshuk, 2002-2013; Penney, 2010; Telnov, 2012b; Soriano et al., 2014; Peris et al., 2016 Lebanese amber Cretaceous 130-125.5
Macratriini LeConte, 1862        
Macratria Newman, 1838 (= Protomacratria Abdullah, 1964) Abdullah, 1964, 1969, Telnov, 2012b Baltic amber Eocene 37.2-33.9
Macratria alleni Telnov, 2012 [LDM] Telnov, 2012b; Alekseev, 2013 Baltic amber Eocene 37.2-33.9
Macratria appendiculata (Abdullah, 1964) [BMNH] Abdullah, 1964: as Protomacratria, 1975: as Protomacratria; Larsson, 1978: as Protomacratria; Spahr, 1981: as Protomacratria; Kubisz, 2000: as Protomacratria; Kirejtshuk, 2002-2013: as Protomacratria; Telnov, 2012b; Alekseev, 2013 Baltic amber Eocene 37.2-33.9
Macratria gigantea Wickham, 1910 [YPM] Wickham, 1910; Kirejtshuk, 2002-2013; Telnov, 2011 lacustrine shale: Florissant, Colorado (USA) Eocene 37.2-33.9
Macratria succinia Abdullah, 1965 [GPIH] Abdullah, 1965, 1975; Larsson, 1978; Spahr, 1981; Kirejtshuk, 2002-2013: as M. succinea [sic!]; Telnov, 2011, 2012b; Alekseev, 2013: as M. succinea [sic!] Baltic amber Eocene 37.2-33.9
Macratria tripunctata (Abdullah, 1964) [BMNH] Abdullah, 1964: as Protomacratria, 1975: Protomacratria; Larsson 1978, as Protomacratria; Spahr, 1981: as Protomacratria; Kirejtshuk, 2002-2013: as Protomacratria; Telnov, 2012b; Alekseev, 2013 Baltic amber Eocene 37.2-33.9
Macratria sp. Klebs, 1910: as Mactratria [sic!]; Bachofen-Echt, 1949; Abdullah, 1964; Larsson, 1978; Spahr, 1981; Poinar, 1992; Kirejtshuk, 2002-2013; Telnov, 2012b; Alekseev, 2017 Baltic amber Eocene 37.2-33.9
tribus incertus, genus sp. present paper Burmese amber Cretaceous 99.7-94.3
Notoxinae Stephens, 1829        
Notoxus sp. Hope, 1836; Helm, 1896; Handlirsch, 1907; Larsson, 1978; Spahr, 1981; Poinar, 1992; Kirejtshuk, 2002-2013 Baltic amber Eocene 37.2-33.9
Steropinae Jacquelin du Val, 1863        
Steropes eleticinoides sp. nov. [LDM] present paper Baltic amber Eocene 37.2-33.9
Steropes sp. Klebs, 1910; Abdullah, 1964; Larsson, 1978; Spahr, 1981; Poinar, 1992; Kirejtshuk, 2002-2013 Baltic amber Eocene 37.2-33.9
Tomoderinae Bonadona, 1961        
Tomoderus balticus Telnov, 2012 [LDM] Telnov, 2012a; Alekseev, 2013 Baltic amber Eocene 37.2-33.9
Tomoderus longelytratus Telnov, 2013 [LDM] Telnov, 2013 Baltic amber Eocene 37.2-33.9
Tomoderus saecularis sp. nov.

[GPIH]

present papet Baltic amber Eocene 37.2-33.9
Tomoderus sp. Klebs, 1910; Bachofen-Echt, 1949; Abdullah, 1964; Larsson, 1978; Spahr, 1981; Poinar, 1992; Kirejtshuk, 2002-2013; Telnov, 2012a, 2013; Alekseev, 2017 Baltic amber Eocene 37.2-33.9
Ischaliidae Blair, 1920        
Ischalia (Ischalia) youngi Alekseev et Telnov, 2016 [CCHH] Alekseev and Telnov, 2016 Baltic amber Eocene 37.2-33.9
Ischalia (Eupleurida) danieli Alekseev et Bukejs, 2017 [GPIH] Alekseev and Bukejs, 2017 Baltic amber Eocene 37.2-33.9
Ischalia subg. Eupleurida LeConte, 1862 Alekseev, 2017 Baltic amber Eocene 37.2-33.9
Ischalia subg. Telnovia Alekseev et Bukejs, 2017 Alekseev and Bukejs, 2017; Alekseev, 2017 Baltic amber Eocene 37.2-33.9
Ischalia (Telnovia) dohnaturris Alekseev et Bukejs, 2017 [KAM] Alekseev and Bukejs, 2017 Baltic amber Eocene 37.2-33.9
Ischalia sp. Alekseev, 2017 Baltic amber Eocene 37.2-33.9

 

 

TABLE 2. Fossil records of higher rank Anthicidae taxa.

Subfamily Tribe Fossil records (Y / N)
Anthicinae Anthicini Latreille, 1819 Y
  Endomiini Bonadona, 1958 Y
  Formicomini Bucciarelli, 1980 N
  Microhorini Bucciarelli, 1980 N
Copobaeninae - N
Eurygeniinae Eurygeniini LeConte, 1862 Y
  Ictistygnini Borchmann, 1937 N
  Mitraelabrini Abdullah, 1969 N
Lemodinae - Y
Macratriinae Camelomorphini Kirejtshuk, Azar et Telnov in Kirejtshuk and Azar, 2008 Y
  Macratriini LeConte, 1862 Y
Notoxinae - Y
Steropinae - Y
Tomoderinae - Y

 

 

 

TABLE 3. Matrix of geographical distribution of extant and fossil Anthicidae and Ischaliidae, except incertae sedis groups (according to Chandler 2010; Lawrence et al. 2010). Legends: WP – Western Palaearctic, EP – Eastern Palaearctic, OR – Oriental, AT – Afrotropical and Madagascan, AU – Australian and Papuan, NA – Nearctic, NT – Neotropical, 0 – extant and fossil absence, 1 – extant presence and fossil absence, 2 – fossil presence and extant absence, 3 – fossil and extant presence.

High rank taxa WP EP OR AT AU NA NT
Anthicidae Latreille, 1819 3 3 3 1 1 3 1
Anthicinae Anthicini Latreille, 1819 3 3 1 1 1 1 3
Anthicinae Endomiini Bonadona, 1958 3 1 1 1 1 0 0
Anthicinae Formicomini Bucciarelli, 1980 1 1 1 1 1 0 1
Anthicinae Microhorini Bucciarelli, 1980 1 1 1 1 0 0 0
Copobaeninae Abdullah, 1969 0 0 0 0 0 0 1
Eurygeniinae Eurygeniini LeConte, 1862 3 1 1 1 1 1 1
Eurygeniinae Ictistygnini Borchmann, 1937 0 0 0 1 1 0 1
Eurygeniinae Mitraelabrini Abdullah, 1969 0 0 0 0 0 0 1
Lemodinae Lawrence et Britton, 1991 0 0 2 0 1 0 1
Macratriinae Camelomorphini Kirejtshuk, Azar et Telnov in Kirejtshuk and Azar, 2008 2 0 0 0 0 0 0
Macratriinae Macratriini LeConte, 1862 3 1 1 1 1 1 1
Macratriinae tribus incertus 0 0 2 0 0 0 0
Notoxinae Stephens, 1829 3 1 1 1 1 1 1
Steropinae Jacquelin du Val, 1863 3 1 1 0 1 1 0
Tomoderinae Bonadona, 1961 3 1 1 1 1 1 1
Ischaliidae Blair, 1920 2 1 1 0 0 1 0

 

 

TABLE 4. Matrix of selected critical characters of the Anthicidae and their minimum age. Only those critical characters visible and observed in fossil specimens are listed.

Period Epoch Age, Ma Subfamily, tribe Selected critical characters which appear in fossil taxa
Neogene Pliocene 5.333-2.58 Anthicinae Anthicini - [see below]
Miocene 23.03-5.333 Anthicinae Anthicini - [see below]
Paleogene Oligocene 33.9-23.03 -  
  Eocene 56-33.9 Anthicinae head with frontoclypeal suture or an impression at place of it; eyes entire; mandible apice uni- or bidentate; antenna 11-segmented; terminal maxillary palpomere fusiform or triangular; neck narrow, 1/4 of head width or less; pronotum anteriorly with rim of equal width dorsally and ventrally; ventrite separated from mesepisterna by distinct suture; metacoxae moderately to widely separated by strongly divergent intercoxal projection; elytra with irregular punctures in most subgroups; tibial spurs small; tarsal claws simple; male genitalia with phallobase and tegmen distinct.
      Anthicinae Anthicini [same as above]
      Anthicinae Endomiini dorsal setae thickened, scale-like; antennal insertions partly covered by protruding anterolateral margin of frons.
      Eurygeniinae Eurygeniini head lacking frontoclypeal suture; eyes emarginate in certain subgroups;terminal antennomere not lengthened; terminal maxillary palpomere securiform, axeform or cultriform; neck broad, 1/3-1/2 of head width; pronotum anteriorly with dorsal rim strongly protruding (flanged); elytra irregularly punctate; tibial spurs distinct.
      Macratriinae Macratriini - [same as subfamily, see below]
      Notoxinae antenna 11-segmented; neck broad and short, 1/3-1/2 of head width; pronotum with prominent anterodorsal horn; pronotum with cervical notches lateral to neck; antebasal sulcus of pronotum distinct; elytra with irregular punctures; tibial spurs distinct.
      Steropinae transverse impression at place of frontoclypeal suture; eyes notched anteriorly; maxillary palpomeres 2-3 rounded apically; terminal maxillary palpomere scalene triangular; antenna 11-segmented; three terminal antennomeres strongly lengthened; neck broad, 1/3-1/2 of head width; pronotum dorsally without anterior rim; pronotum lacking an apical flange; antebasal sulcus of pronotum complete; lateral fovea present on pronotum, ends in a pit distantly from procoxae; lateral fovea and pits glabrous; antebasal sulcus of pronotum complete; procoxal cavities open externally (presumably also internally; not visible); procoxa elongate and truncate fusiform; metacoxae nearly contiguous, narrowly separated by acute intercoxal projection; elytra confusedly punctate; tibial spurs distinct; tibial spur formula 2-2-2; tarsal claws with long ventral spine.
      Tomoderinae frontoclypeal suture variably present; antennae with a weak club of 3-4 terminal antennomeres; neck narrow, 1/3 of head width or less; pronotum with thin anterior rim; antebasal sulcus only indicated laterally near hypomeron; pronotum strongly constricted laterally postmedium; mesoventrite short, semicircular; mesosternum and mesepisterna meet behind the anterior edge of mesothorax; mesoventrite short, semicircular; metacoxae widely separated; metacoxae widely separated; tibial spurs very short to lacking.
      Ischaliidae pronotal disc with median carina, sometimes extending beyond posterior edge, or with longitudinal groove; prosternum flat to moderately convex; prosternum in front of coxae shorter than shortest diameter of procoxal cavity; elytral disc always with more or less complete sutural, sublateral and epipleural costae and often with additional, shorter humeral costa.
  Palaeocene 66-56 -  
Cretaceous Late 100-66 Lemodinae antenna 11-segmented; neck broad, 1/3-1/2 of head width; elytra irregularly punctate.
  Early 145-100 Macratriinae frontoclypeal suture not present; mandible apice bidentate; antenna 11-segmented; 1-3 terminal antennomeres strongly lengthened or widened; maxillary palpomeres 2-3 with angular mesal lobes; terminal maxillary palpomere securiform; neck narrow, 1/4 or less of head width; pronotum with anterior collar of equal width dorsally and ventrally; antebasal sulcus distinct; lateral fovea of pronotum extends anteroventrally at lateral margin towards procoxae in a pit, which is situated just dorsal of procoxa; lateral fovea and pits obscured by setation; mesoventrite triangular, fused to mesepisterna; procoxa comparatively short and ovoid; procoxal rests contiguous when available; metacoxae narrowly separated by acute intercoxal projection; elytra at least with tracks of puncture rows; tibial spurs long and distinct; tibial spur formula 2-2-2; tarsal claws deeply cleft; ovipositor coxite 2-segmented.
      Macratriinae Camelomorphini characters not evaluated (no comparative extant taxa known)

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