
Belongs within: Harpalinae.
Contains: Galeritina, Gigadema, Planetina, Helluomorphoides, Dryptini.
The Zuphiitae are a group of carabid ground beetles whose members possess spination on the first stylomere of the ovipositor (Ober & Maddison 2008). They have been divided between several tribes; however, Ober & Maddison’s (2008) phylogenetic analysis did not support monophyly of the tribes Zuphiini, Galeritini and Helluonini.
Members of the Anthiini and Helluonina are large beetles with an enlarged labrum covering most of the mandibles in dorsal view. Species of the genus Anthia are capable of ejecting a spray of defensive chemicals from glands in the abdomen up to a distance of a metre. Members of the Zuphiini are characterised by truncate elytra and uniformly setose antennae with the first antennomere longer than the following two together. The genus Zuphium, with the head markedly constricted behind the eyes to form a ‘neck’, has a cosmopolitan distribution concentrated in the tropics. The Indo-Australian genus Pogonoglossus has a markedly depressed and elongate body form, a large head with elongate mandibles, a wide, cordiform prothorax, and parallel-sided elytra (Baehr 2014a). Pseudaptinus is a Neotropical genus with the pronotum truncate at the base and the third antennomere shorter than the fourth (Ball & Bousquet 2001).
Ground beetles for today
Published 1 February 2011

The subject of today’s post is a group of ground beetles (Carabidae) that has been treated in the past as the subfamily Zuphiinae, but seems to now be more commonly treated as a supertribe Zuphiitae within the Harpalinae. Whatever their appropriate formal name, the zuphiites are distinguished by a relatively long and thick scape (the first major segment of the antennae) and spination on the first stylomere of the female’s ovipositor (Ball 1985; Ober & Maddison 2008); the clade is also supported by molecular data (Ober & Maddison 2008). Many zuphiites also have a relatively narrow pronotum (noticeably longer than wide) and truncate elytra. Species of Zuphiitae are found around the world, primarily in the tropics, with known centres of diversity in Australia and the Neotropics (Baehr 1985). Diversity in Asia and Africa is supposedly lower but, as most of the Australian taxa appear to be ultimately derived through immigrations from Asia (Baehr 1985), it would not be at all surprising if this distribution was biased by higher levels of study in the former continents*.
*Though ‘higher level of study’ is definitely a relative term: the Australian fauna, for instance, seems to owe its recognised diversity overwhelmingly to the work of Martin Baehr alone.

Within the Zuphiitae, current classifications recognise the tribes Zuphiini, Anthiini, Galeritini, Dryptini, Helluonini and Physocrotaphini, though the monophyly of some of these tribes is not firmly established (Ober & Maddison 2008). The Zuphiini are distinguished by a particularly long scape whereas the Galeritini possess asymmetrically dilated tarsi in the males (Baehr 1985).
Systematics of Zuphiitae
<==Zuphiitae [Anthiitae, Dryptinae, Dryptitae]OM08
|--Colasidia [Leleupidiina, Leleupidiini]B86
| `--C. monteithi Baehr 1988Ba14b
|--AnthiiniOM08
| |--Cypholoba [Cypholobina]OM08
| `--Anthia [Anthiina]OM08
| |--A. cinctipennisB55
| | |--A. c. cinctipennisB55
| | `--A. c. actaeonB55
| |--A. circumscriptaB55
| | |--A. c. circumscriptaB55
| | `--A. c. omostigma [incl. A. niveicincta]B55
| |--A. maxillosaBo14
| |--A. sexguttata [=Graphipterus (Anthia) sexguttatus]G20
| `--A. thoracica (Thunberg 1784)Bo14
`--Zuphiini (see below for synonymy)OM08
| i. s.: ProgaleritaB86
| EunostusB86
| Meladroma lugubreB55
| TriaenogeniusB55
| |--T. anthioides (Chaud. 1872) [=Acanthogenius anthioides]G89
| |--T. arabicus Gestro 1889G89
| |--T. carinulatus (Fairm. 1887) [=Acanthogenius carinulatus]G89
| |--T. corpulentus Chaud. 1877G89
| |--T. ferox (Erichs. 1843) [=Helluo ferox]G89
| |--T. helluonoides (Ancey 1881) [=Anthia helluonoides]G89
| `--T. sculpturatus (Gerst. 1866) [=Acanthogenius sculpturatus]G89
| Acanthogenius trimaculatus Chaud. 1872B89
|--+--GaleritinaOM08
| `--+--+--OmphraOM08
| | `--Macrocheilus Kirby 1838OM08, BR05
| | |--M. bimaculatusB55
| | `--M. biplagiatusB55
| `--+--Acrogenys Macleay 1864OM08, Ba14b [Dicrodontina]
| | | i. s.: A. centralisGWW10
| | |--A. (Acrogenys)LM87
| | | |--*A. (A.) hirsuta Macleay 1864LM87
| | | |--A. (A.) demarzi Baehr 1984LM87
| | | `--A. (A.) laticollis Baehr 1984LM87
| | `--A. (Paracrogenys Baehr 1984)LM87
| | `--A. (*P.) longicollis Gestro 1875LM87
| `--HelluoninaOM08
| | i. s.: Helluo Bonelli 1813LM87
| | |--*H. costatus Bonelli 1813 [incl. H. carinatus Chaudoir 1848]LM87
| | `--H. insignis Sloane 1890LM87
| |--Dicranoglossus Chaudoir 1872OM08, LM87
| | `--*D. resplendens (Castelnau 1867) (see below for synonymy)LM87
| `--+--GigademaOM08
| `--Aenigma Newman 1836OM08, LM87
| `--*A. iris Newman 1836 [incl. A. newmanni Castelnau 1867, A. splendens Castelnau 1867]LM87
`--+--+--Thalpius LeConte 1851OM08, LM87 [incl. Enaphorus LeConte 1851BB01, Zuphiosoma Laporte de Castelnau 1867BB01]
| | |--*T. pygmaeus (Dejean 1826) [=Helluo pygmaeus, Pseudaptinus (*Thalpius) pygmaeus]LM87
| | |--T. australis (Blackburn 1889) [=Acrogenys australis, Pseudaptinus (Thalpius) australis]LM87
| | `--T. fulvus (Castelnau 1867) [=*Zuphiosoma fulva, Pseudaptinus (Thalpius) fulvus]LM87
| `--+--PlanetinaOM08
| `--+--HelluomorphoidesOM08
| `--+--‘Thalpius’ rufulusOM08
| `--Pseudaptinus Castelnau 1834OM08, Ba14b (see below for synonymy)
| |--*P. albicornis Castelnau 1834LM87
| |--P. brittoni Baehr 1985B86
| |--P. hirsutulus Baehr 1985B86
| |--P. iridescens Baehr 1985B86
| |--P. lecontei (Dejean 1831)BB01
| |--P. monteithi Baehr 1985B86
| `--P. tenuicollis (LeConte 1851)BB01
`--+--+--DryptiniOM08
| `--Pogonoglossus Chaudoir 1862OM08, LM87 (see below for synonymy)
| |--*P. validicornis Chaudoir 1862Ba14a
| |--P. danielsi Baehr 2008Ba14a
| |--P. frater Baehr 2014Ba14a
| |--P. inarmatus Baehr 1988Ba14a
| |--P. inflaticeps (Sloane 1904)Ba14a [=Carpaulum inflaticepsLM87]
| |--P. parvus Darlington 1968Ba14a
| |--P. porosus (Sloane 1904)Ba14a [=Carpaulum porosumLM87]
| |--P. rufopiceus Baehr 1993Ba14a
| `--P. soror Baehr 2014Ba14a
`--+--CatapiesiniBS02
| |--Catapiesis brasiliensisOM08
| `--HomalomorphaBS02
`--+--Mischocephalus [Mischocephalina]OM08
`--Zuphium Latreille 1806OM08, Ba14b [=Zophium Gistel 1839BB01, Zoyphium Agassiz 1847BM76]
|--*Z. olens (Rossi 1790)LM87 [=Carabus olensLM87, Galerita (Zuphium) olensG20]
|--Z. australe Chaudoir 1862LM87
|--Z. castelnaui Gestro 1875LM87
|--Z. fitzroyense Macleay 1888LM87
|--Z. flavumGWW10
|--Z. numidicum Lucas 1846E12
|--Z. pindan Macleay 1888LM87
|--Z. rockhamptonense Castelnau 18967 [=Z. rockhamptonensis]LM87
`--Z. thouzeti Castelnau 1867B86
Zuphiitae incertae sedis:
Helluarchus Sloane 1914M80, LM87
|--*H. robustus Sloane 1914LM87
`--H. whitei Lea 1914LM87
Speozuphium Moore 1995Ba14b
Speothalpius Moore 1995Ba14b
`--S. grayiBa14b
Typhlozuphium Baehr 2014Ba14b
|--*T. humicolum Baehr 2014Ba14b
`--T. longipenne Baehr 2014Ba14b
Parazuphium Jeannel 1942B86, Ba14b
|--*P. (Parazuphium) chevrolati (Castelnau 1833) [=Zuphium chevrolati]Ba14b
`--P. (Austrozuphium Baehr 1985)Ba14b
|--P. (*A.) mastersii (Castelnau 1867) [=Zuphium mastersii]Ba14b
|--P. (A.) flavescens Baehr 1985Ba14b
`--P. (A.) pilbarae Baehr 2014Ba14b
Agastus lineatusB89
*Dicranoglossus resplendens (Castelnau 1867) [=Helluosoma resplendens; incl. Aenigma parvulum Macleay 1873]LM87
Pogonoglossus Chaudoir 1862OM08, LM87 [incl. Carpaulum Sloane 1904LM87; Physocrotaphinae, PhysocrotaphiniBa14a]
Pseudaptinus Castelnau 1834OM08, Ba14b [incl. Diaphorus Dejean 1831 non Meigen 1824BB01, Tiphys Gistel 1848 (preoc.)BB01]
Zuphiini [Galeritini, Galeritinini, Galeritulini, Helluomorphina, Helluonides, Helluoninae, Helluonini, Omphrina, Patriziina, Zuphiina, Zuphiinae]OM08
*Type species of generic name indicated
References
[B86] Baehr, M. 1986. Revision of the Australian Zuphiinae. 6. The genus Planetes Macleay. Supplement to the other genera (Insecta, Coleoptera, Carabidae). Spixiana 9 (2): 151–168.
[Ba14a] Baehr, M. 2014a. Two new species of the beetle genus Pogonoglossus from the Pilbara region of north-western Australia (Coleoptera: Carabidae: Physocrotaphini). Records of the Western Australian Museum 29 (2): 89–94.
[Ba14b] Baehr, M. 2014b. Three new species and a new genus of subterranean Zuphiini from the Pilbara region of north-western Australia (Coleoptera: Carabidae: Harpalinae). Records of the Western Australian Museum 29 (2): 95–104.
Ball, G. E. 1985. Reconstructed phylogeny and geographical history of genera of the tribe Galeritini (Coleoptera: Carabidae). In: Ball, G. E. (ed.) Taxonomy, Phylogeny and Zoogeography of Beetles and Ants: A volume dedicated to the memory of Philip Jackson Darlington, Jr. (1904–1983) pp. 276–321. D. W. Junk Publishers.
[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.
[BS02] Ball, G. E., & D. Shpeley. 2002. Ginemini, Ginema thomasi, new tribe, new genus and new species, from Amazonian Bolivia (Coleoptera: Carabidae: Harpalinae). Transactions of the American Entomological Society 128 (1): 75–98.
[B55] Basilewsky, P. 1955. Carabidae (Coleoptera) de l’Angola (première partie). Diamang Publicações Culturais 27: 91–137.
[B89] Bates, H.-W. 1889. Contributions a la faune Indo-Chinoise. 3e mémoire. Carabidae. Annales de la Société Entomologique de France, 6e série 9: 261–286.
[BM76] Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.
[Bo14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature’s gems. Ivy Press: Lewes (United Kingdom).
[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.
[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.
[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.
[GWW10] Guthrie, N. A., T. Weir & K. Will. 2010. Localised and regional patterns in ground-dwelling beetle assemblages in a semi-tropical arid zone environment. Records of the Western Australian Museum Supplement 78 (1): 169–184.
[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.
[M80] Matthews, E. G. 1980. A Guide to the Genera of Beetles of South Australia. Part 1. Archostemata and Adephaga. South Australian Museum: Australia.
[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.