Pterostichini

Notonomus sp., copyright Reiner Richter.

Belongs within: Harpalinae.

The Pterostichini are a group of mostly mid-sized, relatively stout ground beetles, with antennae that are setose from the fourth segment onwards. Most possess two pairs of supra-orbital setae though only one pair is present in Pterostichus honestus (Bell 1990). Members of the Pterostichini include Notonomus, a genus of over 100 species of flightless ground beetle from Australia, characterised by a mentum with oblique sides to the sinus, and a narrow border on the pronotum. The genus Pterostichus is widespread in the Holarctic region as well as in the northern parts of the Oriental and Neotropical regions (Ball & Bousquet 2001).

<==Pterostichini
    |--Myas Stürm 1826 [incl. Neomyas Allen 1980, Trigonognatha Motschulsky 1857]BB01
    |--Gastrellarius Casey 1918BB01
    |    `--G. honestus (Say 1823)BB01
    |--Abaris Dejean 1831BB01
    |    `--A. splendidula (LeConte 1863)BB01
    |--Ophryogaster Chaudoir 1878BB01
    |    `--O. flohri Bates 1882 [incl. Pterostichus arizonicus Schaeffer 1910]BB01
    |--Lophoglossus LeConte 1852BB01
    |    `--L. scrutator (LeConte 1848)BB01
    |--Piesmus LeConte 1848BB01
    |    |--P. monedulus [=Pterostichus (Piesmus) monedulus]Be90
    |    `--P. submarginatus (Say 1823)BB01
    |--Stereocerus Kirby 1837 [incl. Boreobia Tschitschérine 1896]BB01
    |    |--S. haematopus (Dejean 1831)BB01
    |    `--S. rubripes (Motschulsky 1860)BB01
    |--Abax Bonelli 1810BB01
    |    |--A. parallelepipedus (Piller & Mitterpacher 1783)BB01
    |    |--A. parallelusFFB88
    |    `--A. striolaG20 [=Carabus striolaL02, Harpalus (Abax) striolaG20]
    |--Stomis Clairville 1806BB01
    |    |--S. (Stomis) pumicatus (Panzer 1795)BB01
    |    `--S. (Neostomis Bousquet 1983)BB01
    |         `--S. (N.) termitiformis (Van Dyke 1925)BB01
    |--Cyclotrachelus Chaudoir 1838BB01
    |    |--C. (Cyclotrachelus) [incl. Ferestria Leng 1915, Fortax Motschulsky 1866]BB01
    |    `--C. (Evarthrus LeConte 1852) (see below for synonymy)BB01
    |         |--C. (E.) furtivusBB01
    |         |--‘Evarthrus’ gravesiBe90
    |         |--C. (E.) sigillatusBB01
    |         |--C. (E.) sodalis (LeConte 1848)BB01
    |         `--C. (E.) substriatus (LeConte 1848)BB01
    |--Poecilus Bonelli 1810 [incl. Feronia Latreille 1817]BB01
    |    |  i. s.: ‘Feronia’ barbaraBl90
    |    |         P. coerulescensC01
    |    |         P. conformisG89
    |    |         P. fortipesC01
    |    |         P. gebleriC01
    |    |         P. lepidusC01
    |    |         P. numidicus Lucas 1846E12
    |    |         P. punctulatusC01
    |    |         P. scitulusOM08
    |    |--P. (Poecilus) (see below for synonymy)BB01
    |    |    `--P. (P.) cupreus (Linnaeus 1758)G20, L58 (see below for synonymy)
    |    `--P. (Derus Motschulsky 1844) [incl. Derulus Tschitschérine 1896]BB01
    |         `--P. (D.) nearcticus Lindroth 1966BB01
    |--Pterostichus Bonelli 1810BB01
    |    |  i. s.: P. anthracinusC01
    |    |         P. burmeisteriVB21
    |    |         P. chalcites [=Poecilus chalcites]BB68
    |    |         P. cupreus (Linnaeus 1758)VPB11
    |    |         P. dilutipesC01
    |    |         P. eschscholtziiC01
    |    |         P. honestusBe90
    |    |         P. metallicusG20 [=Carabus metallicusL02, Harpalus (Pterosticus) metallicusG20]
    |    |         P. nigerC01
    |    |         P. nigrita (Paykull 1790)VPB11
    |    |         P. oblongopunctatusC01
    |    |         P. pumilioFFB88
    |    |         P. rufipesR26
    |    |         P. vitreusC01
    |    |--P. (Pterostichus) cristatusOT05
    |    |    |--P. c. cristatusOT05
    |    |    `--P. c. heydenianus Jacobson 1907OT07
    |    |--P. (Abacidus LeConte 1863) [incl. Peristethus LeConte 1863]BB01
    |    |--P. (Argutor Dejean 1821) [incl. Lagarus Chaudoir 1838, Pseudargutor Casey 1918, Pseudolagarus Lutshnik 1922]BB01
    |    |    `--P. (A.) commutabilis (Motschulsky 1866) [incl. P. leconteianus Lutshnik 1922]BB01
    |    |--P. (Bothriopterus Chaudoir 1835) [incl. Dysidius Chaudoir 1838, Parargutor Casey 1918]BB01
    |    |    |--P. (B.) adstrictus Eschscholtz 1823BB01
    |    |    `--P. (B.) tropicalis Bates 1882BB01
    |    |--P. (Cylindrocharis Casey 1918)BB01
    |    |--P. (Eosteropus Tschitschérine 1902) [incl. Refonia Casey 1918]BB01
    |    |    |--P. (E.) circulosus Lindroth 1966BB01
    |    |    |--P. (E.) moestus (Say 1823)BB01
    |    |    `--P. (E.) superciliosus (Say 1823)BB01
    |    |--P. (Euferonia Casey 1918)BB01
    |    |--P. (Feronina Casey 1918)BB01
    |    |    `--P. (F.) palmi Schaeffer 1910BB01
    |    |--P. (Gastrosticta Casey 1918)BB01
    |    |--P. (Hypherpes Chaudoir 1838) (see below for synonymy)BB01
    |    |    |--P. (H.) adoxus (Say 1823)BB01
    |    |    |--P. (H.) morionoides Chaudoir 1868BB01
    |    |    `--P. (H.) tristis (Dejean 1828)BB01
    |    |--P. (Lamenius Bousquet 1999)BB01
    |    |    `--P. (L.) caudicalis Say 1823BB01
    |    |--P. (Lenapterus Berlov 1996)BB01
    |    |    `--P. (L.) punctatissimus (Randall 1838)BB01
    |    |--P. (Melanius Bonelli 1810) (see below for synonymy)BB01
    |    |    `--P. (M.) castor Goulet & Bousquet 1983BB01
    |    |--P. (Metallophilus Chaudoir 1838) [incl. Lyperopherus Motschulsky 1844]BB01
    |    |    `--P. (M.) sublaevis Sahlberg 1880BB01
    |    |--P. (Monoferonia Casey 1918)BB01
    |    |    `--P. (M.) diligendus Chaudoir 1868BB01
    |    |--P. (Morphnosoma Lutshnik 1915) [incl. Omaseidius Jeannel 1942]BB01
    |    |    `--P. (M.) melanarius (Illiger 1798)BB01 [incl. P. vulgarisRD77]
    |    |--P. (Orsonjohnsonus Hatch 1933)BB01
    |    |    `--P. (O.) johnsoni Ulke 1889BB01
    |    |--P. (Paraferonia Casey 1918)BB01
    |    |    `--P. (P.) lubricus LeConte 1852BB01
    |    |--P. (Phonias des Gozis 1886) [incl. Americomaseus Csiki 1930, Micromaseus Casey 1918]BB01
    |    |    `--P. (P.) strenuus (Panzer 1797)BB01
    |    |--P. (Pseudoferonina Ball 1965) [incl. Melvilleus Ball 1965]BB01
    |    |--P. (Pseudomaseus Chaudoir 1838)BB01
    |    `--P. (Steropus Dejean 1821)BB01
    |         `--P. (*S.) madidus Fabricius 1775BB01
    |--Cryobius Chaudoir 1838BB01
    |    |--C. aralarensis (Mateu 1945)OT05
    |    `--C. pumilio (Dejean 1828)OT05
    `--Notonomus Chaudoir 1865LB91, LM87 (see below for synonymy)
         |--*N. triplogenioides (Chaudoir 1865) (see below for synonymy)LM87
         |--N. aequabilis Moore 1960LM87
         |--N. aequalis Sloane 1907LM87
         |--N. aeques (Castelnau 1867) [=Feronia aeques, N. eques (l. c.)]LM87
         |--N. amabilis (Castelnau 1867) [=Feronia amabilis]LM87
         |--N. angulosus Sloane 1913LM87
         |--N. angustibasis Sloane 1902LM87
         |--N. apicalis Sloane 1913LM87
         |--N. arthuri Sloane 1890LM87
         |--N. atrodermis Sloane 1903 [=N. atridermis Csiki 1930]LM87
         |    |--N. a. atrodermisLM87
         |    `--N. a. smithi Moore 1961LM87
         |--N. auricollis (Castelnau 1867) [=Feronia auricollis; incl. F. (Notonomus) parallelomorpha Chaudoir 1878]LM87
         |--N. australasiae (Dejean 1828) [=Feronia australasiae; incl. F. mitchelii Castelnau 1867]LM87
         |--N. australis (Castelnau 1840) (see below for synonymy)LM87
         |--N. bakewelli Sloane 1902 [incl. N. hamifer Moore 1960]LM87
         |--N. banksi Sloane 1910LM87
         |--N. besti Sloane 1902 [incl. N. besti aeneodorsis Sloane 1903]LM87
         |--N. bodeae Sloane 1913LM87
         |--N. carteri Sloane 1907LM87
         |--N. chalybaeus (Dejean 1828) [=Feronia chalybea]LM87
         |--N. colossus Sloane 1902 [incl. N. heroicus Lutshnik 1915]LM87
         |--N. crenulatus Sloane 1911 [=N. australis crenulatus]LM87
         |--N. croesus (Castelnau 1867) [=Feronia croesus; incl. N. frenchi Sloane 1902, F. plutus Castelnau 1867]LM87
         |--N. cupricolor Sloane 1903LM87
         |--N. curvicollis Sloane 1923LM87
         |--N. cylindricus Sloane 1913LM87
         |--N. darlingii (Castelnau 1867) [=Feronia darlingii]LM87
         |--N. dehiscens Sloane 1923LM87
         |--N. depressipennis (Chaudoir 1874) [=Feronia (Notonomus) depressipennis]LM87
         |--N. dimorphicus Darlington 1961LM87
         |--N. dives Sloane 1915LM87
         |--N. doddi Sloane 1913LM87
         |--N. dyscoloides (Motschulsky 1865) (see below for synonymy)LM87
         |--N. ellioti Darlington 1961LM87
         |--N. ellipticus Sloane 1923LM87
         |--N. excisipennis Sloane 1902 [incl. Feronia (Notonomus) kingii Chaudoir 1865 non Poecilus kingi Macleay 1826]LM87
         |--N. fergusoni Sloane 1910LM87
         |--N. flos Darlington 1961LM87
         |--N. froggatti Sloane 1902 [incl. N. atripennis Sloane 1903]LM87
         |--N. frontevirens Sloane 1916LM87
         |--N. gippsiensis (Castelnau 1867) [=Feronia gippsiensis]LM87
         |--N. gravis (Chaudoir 1865) [=Feronia (Notonomus) gravis; incl. F. montana Castelnau 1867]LM87
         |--N. hedleyi Sloane 1916LM87
         |--N. hopsoni Sloane 1923LM87
         |--N. ingratus (Chaudoir 1865) [=Feronia (Notonomus) ingrata]LM87
         |--N. johnstoni Sloane 1907 [incl. N. johnstoni parvula Sloane 1915]LM87
         |--N. kershawi Sloane 1902LM87
         |--N. kingi (Macleay 1826)LM87 (see below for synonymy)
         |--N. kosciuskianus Sloane 1902LM87
         |--N. lateralis Sloane 1890 [incl. N. sloanianum Lutshnik 1922]LM87
         |--N. leai Sloane 1902LM87
         |--N. lesueurii (Castelnau 1867)LM87 [=Feronia lesueuriiLM87, Sarticus lesueuriS89]
         |--N. longus Sloane 1913LM87
         |--N. macoyi Sloane 1902 [incl. N. howitti Sloane 1902]LM87
         |--N. marginatus (Castelnau 1840) (see below for synonymy)LM87
         |--N. masculinus Darlington 1953LM87
         |--N. mediosulcatus (Chaudoir 1865) (see below for synonymy)LM87
         |--N. melas Sloane 1903LM87
         |--N. metallicus Sloane 1913LM87
         |--N. miles (Castelnau 1867) [=Feronia miles]LM87
         |--N. minimus Sloane 1907LM87
         |--N. molestus (Chaudoir 1865) (see below for synonymy)LM87
         |--N. montellus Darlington 1961LM87
         |--N. montorum Darlington 1961LM87
         |    |--N. m. montorumLM87
         |    `--N. m. azul Darlington 1961LM87
         |--N. muelleri Sloane 1902LM87
         |--N. nitescens Sloane 1911 [incl. N. bellingeri Sloane 1911]LM87
         |--N. nitidicollis (Chaudoir 1865) (see below for synonymy)LM87
         |--N. obscurus Moore 1963LM87
         |--N. opacicollis (Chaudoir 1865) (see below for synonymy)LM87
         |--N. opulentus (Castelnau 1867) [=Feronia opulenta; incl. N. bassi Sloane 1902]LM87
         |    |--N. o. opulentusLM87
         |    `--N. o. gippslandicus (Castelnau 1867) [=Feronia gippslandica; incl. N. viridulus Straneo 1953]LM87
         |--N. peronii (Castelnau 1867) [=Feronia peronii]LM87
         |--N. philippi (Newman 1842) (see below for synonymy)LM87
         |--N. phillipsii (Castelnau 1867) [=Feronia phillipsii]LM87
         |--N. planipectus Sloane 1903 [incl. N. planipectus purpurata Sloane 1913]LM87
         |--N. pluripunctatus Sloane 1903LM87
         |--N. politulus (Chaudoir 1865) [=Feronia (Notonomus) politula; incl. F. tasmanica Castelnau 1867]LM87
         |--N. polli Sloane 1913LM87
         |--N. prominens Sloane 1913LM87
         |    |--N. p. prominensLM87
         |    `--N. p. illidgei Sloane 1913LM87
         |--N. queenslandicus Sloane 1902LM87
         |--N. rainbowi Sloane 1902 [=N. montanus Rainbow 1899 non Feronia montana Castelnau 1867]LM87
         |--N. resplendens (Castelnau 1867) [=Feronia resplendens; incl. N. rugitarsis Sloane 1913]LM87
         |--N. saepistriatus Sloane 1907 [incl. N. triplicatus Darlington 1953]LM87
         |--N. satrapa (Castelnau 1867) [=Feronia satrapa]LM87
         |--N. scotti Sloane 1907 (see below for synonymy)LM87
         |--N. semiplicatus (Castelnau 1840) [=Poecilus semiplicatus]LM87
         |--N. spenceri Sloane 1902LM87
         |--N. sphodroides (Dejean 1828) (see below for synonymy)LM87
         |--N. spurgeoni Darlington 1953LM87
         |--N. striatocollis (Castelnau 1867) [=Feronia striatocollis; incl. N. discorimosus Sloane 1902]LM87
         |--N. strzeleckianus Sloane 1902LM87
         |--N. subiridescens (Chaudoir 1865) [=Feronia (Notonomus) subiridescens; incl. N. liragerus Sloane 1902]LM87
         |--N. taylori Sloane 1903LM87
         |--N. tenuistriatus Sloane 1913 [incl. N. tenuistriatus reticollis Sloane 1913]LM87
         |--N. tessellatus Sloane 1913LM87
         |--N. tillyardi Sloane 1913LM87
         |--N. transitus Darlington 1961LM87
         |--N. truncatus Sloane 1916LM87
         |--N. tubericauda Bates 1878LM87
         |--N. variicollis (Chaudoir 1865) [=Feronia (Notonomus) variicollis; incl. F. comes Castelnau 1867]LM87
         |--N. victoriensis Sloane 1902LM87
         |--N. violaceus (Castelnau 1834) [=Trigonotoma violacea; incl. N. fletcheri Sloane 1902]LM87
         |--N. wentworthi Sloane 1913LM87
         `--‘Feronia’ wilcoxii Castelnau 1867 [incl. F. (Notonomus) subopacus Chaudoir 1874]LM87

Cyclotrachelus (Evarthrus LeConte 1852) [incl. Anaferonia Casey 1918, Eumolops Casey 1918, Evarthrops Casey 1920, Megasteropus Casey 1918]BB01

Notonomus Chaudoir 1865LB91, LM87 [incl. Adetipa Castelnau 1867LM87, Neuropates Motschulsky 1865LM87, Orbitus Motschulsky 1865LM87, Ternox Motschulsky 1865LM87]

Notonomus australis (Castelnau 1840) [=Trigonotoma australis; incl. F. (Notonomus) aeneomicans Chaudoir 1865, F. lapeyrousii Castelnau 1867]LM87

Notonomus dyscoloides (Motschulsky 1865) [=Neuropates dyscoloides; incl. Feronia semiviolacea Castelnau 1867, F. (Notonomus) simulans Chaudoir 1874, F. subvilis Castelnau 1867, F. victoriae Castelnau 1867]LM87

Notonomus kingi (Macleay 1826)LM87 [=Poecilus kingiLM87, Feronia (Poecilus) kingiiB35; incl. F. (Notonomus) incrassata Chaudoir 1874LM87]

Notonomus marginatus (Castelnau 1840) [=Poecilus marginatus; incl. Feronia (Notonomus) discodera Chaudoir 1865, F. hunteriensis Castelnau 1867, Notonomus marginatus luculentus Sloane 1902, N. sydneyensis Sloane 1902]LM87

Notonomus mediosulcatus (Chaudoir 1865) [=Feronia (Notonomus) mediosulcata; incl. F. occidentalis Castelnau 1867, *Adetipa punctata Castelnau 1865, F. satanas Castelnau 1867]LM87

Notonomus molestus (Chaudoir 1865) [=Feronia (Notonomus) molesta; incl. F. lacustis Castelnau 1867, *Ternox obsoletus Motschulsky 1865]LM87

Notonomus nitidicollis (Chaudoir 1865) [=Feronia (Notonomus) nitidicollis; incl. N. angustipennis Macleay 1871, N. cyaneocinctus Macleay 1871, Feronia impressicollis Castelnau 1867, N. latibasis Sloane 1902, N. nitidicollis obtusicollis Sloane 1913, N. purpureipennis Macleay 1871, N. violaceomarginatus Macleay 1871, N. viridicinctus Macleay 1871, Feronia viridilimbata Castelnau 1867, F. viridimarginata Castelnau 1867]LM87

Notonomus opacicollis (Chaudoir 1865) [=Feronia (Notonomus) opacicollis; incl. F. impressipennis Castelnau 1867, F. purpureolimbata Castelnau 1867, *Orbitus purpuripennis Motschulsky 1865, Notonomus rugosicollis Sloane 1902]LM87

Notonomus philippi (Newman 1842) [=Feronia philippi; incl. N. philippi arcuata Sloane 1913, F. bipunctatus Castelnau 1867, N. chalybeus otwayensis Sloane 1902]LM87

Notonomus scotti Sloane 1907 [=N. kingi Sloane 1902 nec Poecilus kingi Macleay 1826 nec Feronia (Notonomus) kingii Chaudoir 1865]LM87

Notonomus sphodroides (Dejean 1828) [=Feronia (Poecilus) sphodroides; incl. F. (Notonomus) accedens Chaudoir 1865, *Neuropates pristonychoides Motschulsky 1865]LM87

*Notonomus triplogenioides (Chaudoir 1865) [=Feronia (*Notonomus) triplogenioides; incl. F. ducalis Castelnau 1867, Notonomus jervisensis Sloane 1902]LM87

Poecilus (Poecilus) [incl. Americobius Lutshnik 1915, Leconteus Lutshnik 1915, Parapoecilus Jeannel 1942, Sogines Stephens 1828]BB01

Poecilus (Poecilus) cupreus (Linnaeus 1758)G20, L58 [=Carabus cupreusL02, Harpalus (P.) cupreusG20; incl. Poecilus cupreus ab. erythropusC01]

Pterostichus (Hypherpes Chaudoir 1838) [incl. Anilloferonia Van Dyke 1926, Brachystilus Chaudoir 1838, Gonoderus Motschulsky 1859, Hammatomerus Chaudoir 1868, Haplocoelus Chaudoir 1838, Holciophorus LeConte 1852, Leptoferonia Casey 1918, Pheryphes Casey 1920]BB01

Pterostichus (Melanius Bonelli 1810) [incl. Lyperus Chaudoir 1838, Metamelanius Tschitschérine 1900, Omaseus Dejean 1821]BB01

*Type species of generic name indicated

References

[BB01] Ball, G. E., & Y. Bousquet. 2001. Carabidae Latreille, 1810. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 32–132. CRC Press: Boca Raton.

[BB68] Basford, N. L., J. E. Butler, C. A. Leone & F. J. Rohlf. 1968. Immunologic comparisons of selected Coleoptera with analyses of relationships using numerical taxonomic methods. Systematic Zoology 17 (4): 388–406.

[Be90] Bell, R. T. 1990. Insecta: Coleoptera Carabidae adults and larvae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1053–1092. John Wiley & Sones: New York.

[Bl90] Blackburn, T. 1890. Notes on Australian Coleoptera, with descriptions of new species. Part IV. Proceedings of the Linnean Society of New South Wales, series 2, 4 (3): 707–746.

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d’Urville. Faune entomologique de l’océan Pacifique, avec l’illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

[C01] Csiki, E. 1901. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l’Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[FFB88] Franke, U., B. Friebe & L. Beck. 1988. Methodisches zur Ermittlung der Siedlungsdichte von Bodentieren aus Quadratproben und Barberfallen. Pedobiologia 32: 253–264.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso “Esploratore” dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 5–72.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[LB91] Lawrence, J. F., & E. B. Britton. 1991. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 543–683. Melbourne University Press: Carlton (Victoria).

[LM87] Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis revised 10th ed. vol. 1. Laurentii Salvii: Holmiae.

[OM08] Ober, K. A., & D. R. Maddison. 2008. Phylogenetic relationships of tribes within Harpalinae (Coleoptera: Carabidae) as inferred from 28S ribosomal DNA and the wingless gene. Journal of Insect Science 8: 63.

[OT05] Ortuño, V. M., & M. Toribio. 2005. Descripción de un nuevo Trechus Clairville, 1806 (Coleoptera, Carabidae, Trechini) de los montes cantábricos orientales (norte de España). Graellsia 61 (1): 115–121.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms’ General Textbook of Entomology 10th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[R26] Risso, A. 1826. Histoire naturelle des principales productions de l’Europe méridionale et particulièrement de celles des environs de Nice et des Alpes maritimes vol. 5. F.-G. Levrault: Paris.

[S89] Sloane, T. G. 1889. Studies in Australian entomology. No. I.—Review of the genus Sarticus (Carabidae). Proceedings of the Linnean Society of New South Wales, series 2, 4 (2): 501–512.

[VPB11] Varet, M., J. Pétillon & F. Burel. 2011. Comparative responses of spider and carabid beetle assemblages along an urban-rural boundary gradient. Journal of Arachnology 39 (2): 236–243.

[VB21] Vasilikopoulos, A., M. Balke, S. Kukowka, J. M. Pflug, S. Martin, K. Meusemann, L. Hendrich, C. Mayer, D. R. Maddison, O. Niehuis, R. G. Beutel & B. Misof. 2021. Phylogenomic analyses clarify the pattern of evolution of Adephaga (Coleoptera) and highlight phylogenetic artefacts due to model misspecification and excessive data trimming. Systematic Entomology 46: 991–1018.

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