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Carabidae Latreille, 1802

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Fig. 7.7.4. Larval head structures. A, nasale and adnasalia, Systolosoma lateritium, S. breve, Trachypachus IlOlmbergi; B,antennae, S. lateritium, T.holmbergi;C- E, S. lateritium.C, mandible; D, maxilla, E, labium; F- H, tergite IX. F. S. lateri-tium; G, T. holmbergi; H, S. breve. From Beutel & Arndt (1995), redrawn.

morphies (Arndt & Beutel 1995): sensorial ap-pendage on lateral side of antennomere III ab-sent, replaced by ventral sensorial field, apicalpart of maxillary palpomere 3 with additional se-tae, number of nasal teeth increased (6-8), uro-gomphi fixed, horn-shaped (groundplan), eightlong setae on tergite IX (including those on uro-gomphi). The specific shape of the parameres(Lindroth 1961-69; Beutel 1994) is an autapo-morphy of adults. The absence of the katas-tigma, the specific sculpture of the elytra, thekidney-shaped sensorial field of the larval anten-nomere 3 and the large, ventral sensorial fieldof antennomere 4 are larval autapomorphies ofSystolosoma. The dilated larval distal maxillarypalpomere with two separated sensorial fields isan autapomorphy of Trachypachus.

Acknowledgements

We thank Dr. Vasily V. Grebennikov (Ottawa)for thoroughly reviewing this chapter.

Literature

Arndt, E. & Beutel, R. G. (1995): Larval morphologyof Systolosoma Solier and Trachypachus Motschul-sky (Coleoptera: Trachypachidae) with phyloge-netic consideration. - Entomologica Scandinavica26: 439-446.

Beutel, R. G. (1993): Phylogenetic analysis of Ade-phaga (Coleoptera) based on characters of the lar-val head. - Systematic Entomology 18: 127-147.(1994): Study on the systematic position of Systolo-soma breve Solier (Adephaga: Trachypachidae)based on characters of the thorax. - Studies onNeotropical Fauna and Environment 29 (3): 161-167.

Bils, W. (1976): Das Abdomenende weiblicher, ter-restrisch lebender Adephaga (Coleoptera) und seineBedeutung fur die Phylogenie. - Zoomorphologie84: 113-193.

Crowson, R. A. (1955): The Natural Classification ofthe Families of Coleoptera. 187 pp. Nathaniel Lloydand Co., LTD., London.

Hlavac, T. F. (1975): The prothorax of Coleoptera (ex-cept Bostrichiformia- Cucujiformia). - Bulletinof the Museum of Comparative Zoology 147 (4):137-183.

Lindroth, C. H. (1960): The larva of TrachypachusMtsch., Gehringia Darl., and Opisthius Kirby (Col.Carabidae). - OpusculaEntomologica25: 30-42.(1961- 69): The ground beetles (Carabidae, excl.Cicindelinae) of Canada and Alaska. Parts 1-6. -Opuscula Entomologica XlVIII + 1192 pp. 1961,Part 2, Suppl. 20: 1- 20; 1963, Part 3, Suppl. 24:201-408; 1966, Part 4, Suppl. 29: 409-648; 1968,Part 5, Suppl. 33: 649-944; 1969 Part 6, Suppl. 34:945-1192; 1969 Part I, Suppl. 35 I + XlVIII.

Ponomarenko, A. G. (1977): Suborder Adephaga, etc.Pp. 3-104 In Arnoldy, L. v., Zherikhin, V. V.,Nikritin, L. M. & Ponomarenko, A. G. (eds.) Meso-zoic Coleoptera [in Russian]. - Trudy Paleontolo-gicheskogo Instituta Akademiya Nauk SSSR 161:1-183.

7.8. Carabidae Latreille, 1802

Erik Arndt, Rolf G. Beutel & Kipling Will

In most recent classifications Rhysodidae (-ni),Cicindelinae and Paussinae are included. How-ever, Rhysodidae were excluded by Kryzhanov-skij et al. (1995) and Lawrence & Newton (1995).They are tentatively treated as a separate familyhere (s. 1-7.9).

Distribution. World-wide (except Antarctica),more than 40000 spp. and 1500 genera. Distribu-tion of subgroups see below.

Biology and Ecology. Carabidae live in all ter-restrial habitat types from the subarctic to thewet tropical regions. The majority of species ex-cept those of tropical rain forests and subtropicalmontane forests (e. g., Mexico, Hawaii) is ground

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dwelling. Adults and larvae live in soil, in leaflitter or are active on the ground surface, some-times climbing bushes or plants. Many species,mainly in Trechini, Anillini and Platynini, in-habit caves or deep soil microcaves (mss). Eventhough many species occur in moist habitats, anamphibious way of live is reported in only veryfew cases (e. g., Carabu variolosus at smallstreams in Fagus forests, C. clathratus and someother Carabus species in swamps; Sturani 1962;Cicindelinae in inundation forests; Adis & Mess-ner 1997; Raw/insius in shallow regions of rapidmountain streams; Davidson & Ball 1998). Alarge percentage of species in tropical rain forestsand subtropical montane forests, and few speciesin temperate regions are arboreal and resting un-der leaves or bark. The majority of carabid spe-cies is able to fly and has a great dispersal power.Some species are dimorphic regarding their wingand muscle development; usually the minority ofindividuals is wingless (e. g., most Carabus, An-thiini). Even species exclusively moving on theground have an enormous dispersal power;migrations of 77 m per night and running speedsof 0.16 mls were estimated in Carabus species(Thiele 1977). Carabid beetles show a circadianrhythm and outside the tropical regions also anannual rhythm. The majority of species is noc-turnal; some species change their activity de-pendent from climate or season but some taxaare active during day light (e. g., Cicindelini).The annual rhythm controls the reproductiveperiod; different mechanisms of diapause are de-scribed for species of the temperate and sub-tropical regions (Paarmann 1979). During in-active periods in winter quarters or hiding placesduring day time, aggregations of several hundredspecimens are possible.

Carabidae are bisexual. Females lay eggs (asfar as known) separately or in small groups insmall hollows in substrate or under bark or in

cases made of mud (King 1919; Thiele 1977) oralgae and bark (Will 1998). The number of eggsper female varies between four and several hun-dreds depending on the species (the highest num-bers recorded are 653 for Calosoma sycophanta;Thiele 1977 and 660 for Colpodes buchanani;Paarmann & Bolte 1990). The egg developmentlasts few days to several weeks. Most of the spe-cies have three larval instars. Exceptions are twolarval instars in some representatives of Zabrini,Harpalini, Lebiini and Anthiini (Bily 1975;Capogreco 1989; Arndt & Paarmann 1999),which is regarded as an adaptation to an aridenvironment, and four or five larval instarsrespectively in members of Lebiini and Brachi-nini (Erwin 1967, 1975), which is regarded as anadaptation to an ectoparasitic way of live. Pseu-domorphini are ovoviviparous (Liebherr & Ka-


vanaugh 1985; Baehr 1997). Brood cart:}occursin some species of Pterostichini, were the femalesguard their eggs (Kavanaugh 1998) and in somespecies of Harpalini, where the female lay eggsin individual soil cells prepared with food stores.Pupation usually takes place in moist soil. Thefull-grown last instar larvae dig a pupal chamberusing their head and legs. In one species(Thermophilus sexmaculatum) the first instarlarva digs the pupal chamber, and the final sec-ond instar remains inactive in this chamber(Arndt & Paarmann 1999). The life span in thefield is up to four years; at least carabids of thetemperate regions live usually longer than oneyear.

Adults and larvae have a partial extraoral di-gestion. During the manipulation of the prey,mandibles and maxillae rotate the food itemwhile digestive fluids are expelled onto it. Adultsof most species are omniv-orous (Larochelle1990;Thiele 1977), even though carnivorous nut-rition seems to prevail. A few groups are speciali-sed herbivores, e. g., Zabrus and some represen-tatives of Harpalini; even arboreal species ofAgra were found to feed on flowers and nectar(Arndt et al. 2001). Much less is known on thefeeding preferences of larvae, but carnivoroushabits of the majority of species is likely. Larvaeof several taxa are specialised on specific preysuch as snails (Cychrini, Licinini), springtails(Nebriini, Notiophilini, Loricerini), ants, or antbrood (Anthiini, Graphipterini, Metriini, Ozae-nini). Larvae of Paussini and Pseudomorphiniare probably fed by ants and those of Lebiiniand Brachinini are ectoparasitic on insect eggsand pupae as far as known. Larvae of Cicindef-inae and Ozaenini live in burrows and lie in am-brush for prey at the entrances. Larvae of severalHarpalini are specialised on seeds (Brandmayr1975; Zetto Brandmayr 1990; Arndt et at. 1996).Only very few ground beetles are known as pestinsects. The grain ground beetle (Zabrus tenebri-oides) is the only species of a certain economicimportance, but it became known as serious pestonly in a restricted period and restricted area ineastern Germany, Moravia and the Ukraine.

The ecological importance of the extremely di-verse family Carabidae is not well understood.The ground beetles represent a major part of theinvertebrate predator guild of the soil fauna,e. g., in temperate forests and agrocoenoses. Dueto their abundance and ubiquitous occurrence,the importance of the role of carabids in theseecosystems can be safely assumed.

Insectivores, bats, rodents, birds, amphibians,reptiles, ants, Asilidae, and Aranaea are knownas predators of ground beetles. Sporozoa (Gre-garinida), Nematoda (Mermis, Hexamermis),Nematomorpha (Gordius) are endoparasites, andseveral taxa of mites (Trombidiiformes, Sarcopti-formes) are known as ectoparasites on carabids.


B

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Fig.7.8.1. Carabidae, habitus of adults of different subfamilies. A, Lophyra sumlini (Cicindelinae) (Cassola 1976); B,Omophronaequale (Omophroninae); C, Trechuskurentzowi (Trechinae); D, Pheropsophusjavanus (Brachininae) (B- DLafer, 1989).

'--Several other representatives of Acari (e. g., Par-asitus) occur frequently on Carabidae but theyare phoretic. Parasitoids of carabid larvae in-clude the hymenopteran taxa Proctotrupes spp(Proctotrupidae; on Harpalinae and Carabinae),Microtonus spp (Braconidae; on Harpalinae),Methocha spp (Tiphiidae; on Cicindelinae) andrepresentatives of the dipteran families Larvae-voridae (on several subfamilies) and Bombyli-idae (Anthrax spp; on Cicindelinae). Besidesnon-specific insect parasitizing fungi (Ento-mophtorales, Hypomycetes) 16 genera with sev-eral hundreds of species of Laboulbeniales (As-comycota) infest carabids.

The following references summarise informa-tion on the biology of "these beetles: Den Boer(1977), Thiele (1977), Larochelle (1990), Lin-droth (1992), Turin (2000).

Morphology, Adults (Figs. 7.8.1-7.8.3). 1-85 mm long. Usually flattened and elongate,with distinct pronoto-elytral angle, rarely withdistinctly convex dorsal side and laterally evenlyrounded (e. g., Omophron). Cuticle usuallysmooth and shiny. Colour black or dark in mostspecies, sometimes metallic. Depigmentation ofparts of the integument can result in conspicuouscolour patterns (e. g., Cicindela, Eurynebria,Omophroninae, Lebia). Microsculpture usuallypresent as a fine polygonal meshwork. Stronglyimpressed microsculpture decreases brilliance ofcuticular surface. Long articulated setae (='macrochetes' sensu Jeannel 1941-42; fixed se-tae) distributed with regular patterns (chaeto-taxy). Small articulated hairs with irregular dis-tribution present or absent, sometimes formingpubescent surfaces.

Head (Figs. 7.8.2A, 7.8.3.A) prognathous, re-latively elongate, moderately retracted. Withoutdistinct neck region. Frontal furrows present orabsent. Compound eyes usually well developed

and protruding, but different degrees of reduc-tion and total loss occur. Clypeus trapezoid, withtwo pairs of fixed setae. Supraocular area withone or two pairs of fixed setae and suboculargenal area with one pair of setae. Gula fairlyelongate, moderately broad, distinctly delimitedby gular sutures, always glabrous, without fixedsetae. Median gular apodeme present or absent.Tentorium with all parts well developed, usuallywith median laminatentorium. Labrum variablein shape, transverse, medially emarginate, bi-lobed or rarely trilobed. Anterior margin with sixor eight fixed setae. Anterolateral margin usuallywith shorter curved setae. Ventral side largelyunsclerotized and covered with sensorial struc-tures, with shallow excavations adapted to theconvex dorsal surface of the mandibles. Anten-nae almost always filiform and II-segmented,usually inserted below lateral frontal projection,rarely inserted on dorsal side of head capsule(Cicindelinae). Pubescence present on antenno-meres 4-11 or 3-11, sometimes also on basalsegments but then less dense than on distal an-tennomeres. Mandibles with basic form of a tri-angular pyramid, with dorsal, ventral and exter-nal surface, moderately to strongly (e. g., Leca-nomerus; Acorn & Ball 1991: Fig. 16) elongate.Primarily with distinct apical tooth, terebraltooth, anterior retinacular tooth, posterior reti-nacular tooth, premolar or molar tooth, incisorridge, terebral ridge, retinacular ridge, ventralgroove with row of microtrichiae and basalbrush (Acorn & Ball 1991). Maxillae composedof transverse, short cardo, which articulates withwell developed fossa maxillaris, triangular ba-sistipes, mediostipes, elongate triangular palpiferwith several fixed setae, lacinia, galea and 4-seg-mented palp. Lacinia basally fused with medio-stipes, hook-shaped, with dense row of mesallydirected strong setae and thin hairs. Galea 2-seg-mented, palp-like. Palpomere 1 very short.Palpomeres 3 and 4 vary strongly in size, usually

122 Erik Arndt, Rolf G. Beutel & Kipling Will

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Fig. 7.8.2A- D, Gehringiaolympiaca, adults. A, head, ventral view; B, protibia with antenna cleaning organ; C, meso-ventrite; D, metacoxae; E- F, Therates waagenorumHorn, 1900 (Cicindelinae), adult. E, head, ventral view; F, meso-and metaventrite.

in correlation with labial palpomeres 2 and 3.Submental part of labium T-shaped, fused withgula posteriorly. Transverse anterior part mesallydelimits fossa maxillaris, with two or more pairsof fixed setae. Mentum with distinct lateral lobesand often with a more or less projecting medianbifid or simple process, with one pair of fixedsetae. Prementum variable in shape but usuallyroughly quadrangular, with sclerotized medianpart and more or less distinct, less strongly scle-rotized lateral lobes (paraglossae). Anterior mar-gin with one pair of long, fixed setae close tomedian line and sometimes additional shorter

lateral setae. Parallel-sided palpigers inserted be-tween emargination of mentum and prementum.Palp, 3-segmented, with very short palpomere 1.Preoral cavity usually with well developed filterapparatus composed of rows of hairs on mandi-bles, maxillae, and hypopharynx.

Prothorax (Fig. 7.8.3 C) rounded laterally andmore or less strongly narrowed posteriorly, dis-tinctly narrower than elytra at posterior margin(with few exceptions, e. g., Omophron, Pseudo-morphinae). Pronotum medially divided by lon-gitudinalline, usually with raised lateral margin(indistinct or absent in Dyschirius and Apotomus)

Carabidae Latreille, 1802 123

Fig. 7.8.3A- D, Elaphrussp., adults. A, Head, ventral view;B, protibia with antenna cleaning organ; C, Pro- and meso-ventrite; D, Metacoxae.

and distinct basal impression. Lateral marginwith two or more pairs of fixed setae, one ofthem located close to posterior margin. Proster-nal process either narrow and short (Gehringia),strongly developed, projecting beyond the hindmargin of the procoxae and tapering posteriorly(Metrius, Carabinae, Hiletinae), or shortenedand posteriorly truncate (Paussinae excl. Met-rius, Cicindelinae, Omophroninae, Elaphrinae,Loricerinae etc.). Procoxal cavities open, withoutinternal postcoxal bridge (Gehringia, Carabinae,Hiletinae) or externally closed. Protibiae usuallywith two apical spurs, both inserted apically(Paussinae [= "Isochaeta"], Cicindelinae, Opis-thiini, Carabini) or one spur shifted proximally("Anisochaeta"). Antennal cleaning (Figs.7.8.2 B, 7.8.3 B) organ restricted to apical part oftibia (Cicindelinae, Opisthiini, Carabini) or ex-tended towards base (Pausiinae, Gehringia,Omophroninae, Hiletinae etc.). Scutellar shieldenclosed by elytral bases or shifted anteriorly(e. g., Broscus). Mesoventrite of Gehrinigiini(Fig. 7.8.2 C), Carabinae and Hiletinae short,with hexagonal groove and anterolateral groovesfor reception of pro coxae (= carabine type).Mesoventrite of other groups (Figs. 7.8.2 F,7.8.3C) moderately elongated, without hexago-nal groove and anterolateral grooves, rounded

in cross section, with smooth collar region andarticulating with prothorax in a ball-and-socketmanner (Figs. 7.8.2 F, 7.8.3 C; Paussinae [excl.Metrius], Cicindelinae, Loricerinae, Elaphrinae,Migadopinae, Scaritinae, Trechinae, Harpalinae,Brachininae, Pseudomorphinae) (= harpalinetype). Mesocoxal cavities laterally open, i. e. bor-dered by mesepimeron ('disjunct type') or closed('conjunct type') (Trechinae, Harpalinae, Bra-chininae, Pseudomorphinae). Elytra usually cov-ering abdominal tergites completely, apicallytruncate in some groups (Paussinae excl. Met-rius, Gehringiinae, Lebiini, Odacanthini, Galeri-tini, Brachininae). Basal margin present or ab-sent. Disc primarily with eight striae and nineinterspaces. Setae generally present in inter-spaces 3, 5 (discal setae), and 9, but sometimesalso in 1 and 7. Sutural stria usually not recur-rent at apex. Epipleura broad at humeral region,gradually narrowing posteriorly, usually endingat apical external angle of elytra, not reachingsutural angle. Always without setae but some-times with pubescence. Metanotum usually ofgeneralised adephagan type but shortened andstrongly simplified in forms with completely re-duced flight organs (e. g., Omus). Anepisternumdoes not reach mesocoxal cavity. Epimeron ex-posed, narrow and parallel-sided (Gehringiinae,

124

A B


c D EFig. 7.8.4. Habitus of larvae of different subfamilies. A, Carabusexaratus (Carabinae) (redrawn from Arndt & Makarov2003); B, Platychile pallida (Cicindelinae) (Arndt 1998b); C, Lindrothius sp. (Harpalinae, Platynini) (Arndt & Hurka1992);D, Lionychus quadrillum(Harpalinae, Lebiini) (Arndt 1989);E, Trichognathusmarginipennis(Harpalinae, Galeri-tini) (Arndt & Drechsel 1998).

Omophroninae, Elaphrinae, Loricerinae, Miga-dopinae, Scaritini), concealed (Opisthiini, Carab-inae), or lobate (Hiletinae, Trechinae, Harpal-inae). Metaventrite (7.8.20, F, 7.8.3 D) withdiscrimen and complete transverse suture sepa-rating preepisternum from katepisternum. Meta-coxae (7.8.20, F, 7.8.3 D) not extended crani-ally, medially not fused and not fused to kate-pisternum, mobility partly retained (c. 5°). Later-ally scarcely broader or as broad as posteriormargin of ventrite, not reaching elytral epipleuralaterally. Metafurca well developed. Mm. furcacoxalis anterior and posterior present. Alaesometimes reduced, if well developed with ob-longum but without katastigma (subcubital setalbinding patch).

Abdomen usually with six visible sternites.Sternite II only visible laterally in most groups.Large median piece of sternite II usually presentin species with distinctly separate metacoxae(Metrius, Ozaenini part.). Sternites III + IVfused. Terminal sternite VII posteriorly acumi-nate, rounded, or truncate with rounded lateraledges. Tergites I-VIII with spiracles. Posteriorsegments invaginated and strongly modified.Aedeagus usually with asymmetric parameres.Coxosternum VIII only exposed in Brachininae.Gonocoxae relatively short, probably primarily

unsegmented, but divided into proximal and dis-tal part in most representatives.

Morphology, Larvae (Figs. 7.8.4-7.8.10). cam-podeiform, moderately flattened, subparallel.Sclerotized parts brownish to black. Head prog-nathous, rounded laterally (Paussinae, Cicindel-inae) or roughly quadrangular (Fig. 7.8.5 A); cer-vical groove present laterally in most taxa butabsent in basal groups (e. g., Paussinae, Carab-inae, Cicindelinae, Omophroninae). Head later-ally with six stemmata arranged in two rows. Oc-ular groove present posterior to the stemmata inmany taxa but absent in basal groups (e. g.,Paussinae, Carabinae). Frontal suture almost al-ways sinuate; posterior part of frontale (pars ab-oralis frontalis) usually with pairwise egg burst-ers; coronal suture usually present, oftenstrongly shortened or absent. Anterior clypeola-bral margin primarily with four nasal teeth(Fig. 7.8.5 A), each with one small ventrally di-rected microseta; ventral side of clypeolabral re-gion with row of teeth. Antennae 4-segmented,anteriorly directed. Antennomeres I and 2 sub-cylindrical, antennomere 3 with bulb-like senso-rial appendage laterally, antennomere 4 smaller,rounded apically. Mandibles with retinaculum;terebrum usually with two cutting edges; penicil-lus present or absent. Maxillae articulated with


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Fig. 7.S.SA, Lindrothius horsti, head capsule with mandible and antenna, first instar, dorsal aspect (Arndt & Hurka1992);S, C, Agonum miilleri. S, maxillary palpus, dorsal aspect (Arndt 1993);C, labium and labial palpi, dorsal aspect(Arndt 1993);D, abdominal tergite III, lateral aspect, Carabus nemoralis (left) and Cicindelacampestris (right) (Arndt1997);E, Agonum miilleri,pygopod, lateral aspect (Arndt 1993).

anteroventral margin of head capsule, moveablein all directions; maxillary groove completely re-duced; cardo small, with or without separate me-sal sclerite; stipes usually elongate; mobility be-tween both parts strongly restricted; lacinia usu-ally short (elongate and hook-shaped in Metrius,Ozaenini and Omophron), absent in some groups;galea 2-segmented; palpi 4-segmented (or 3-seg-mented and inserted on palpifer, Fig. 7.8.5 B).Maxillae moved by four longitudinally arrangedextrinsic muscles; craniostipital muscle (levatorof maxilla) probably homologous with M.craniolacinialis. Submentum completely fusedwith remaining parts of head capsule; mentumshort and membranous; prementum usually withligula and 2-segmented palpi (Fig. 7.8.5 C). Pre-mentum retracted by two pairs of muscles withtentorial origin. M. submentopraementalis ab-

sent. Hypopharynx with dense field of micro-trichiae, separated from dorsal surface and bulg-ing or completely flattened. M. tentoriohypo-pharyngalis primarily present, but absent in lar-vae with flattened hypopharynx (e. g., Ptero-stichus, Brachinus). Gula usually elongate, repre-sented by a narrow area enclosed by paired gularsutures, rarely broad (e. g., Loricerinae; Licinus).Functional mouth open but narrow, in close con-tact with anterior hypopharyngeal margin. Preo-ral filter apparatus formed by hypopharyngealtrichiae and trichiae on the mandibular (penicil-lus) and maxillary bases. Prepharyngeal tube al-ways present. M. clypeopalatalis always repre-sented by several bundles. Pharynx fairly narrow,primarily with well developed ventral dilator (M.tentoriopharyngalis) and postcerebral dorsal di-lator (M. verticopharyngalis), but both muscles

126

D E F G


Fig. 7.8.6A-C, Brachinuscrepitans,first instar, dorsal aspect; A, antenna; B, maxilla; C, tergite IX with urogomphi; D,Elaphrus sp., urogomphi, third instar; E, Blethisa multipunctata, urogomphi, third instar; F-G, Omophron limbatum,doral aspect, F, mandible; G, labium; H, Clivinafossor, third instar, urogomphi; I, Dyschiriussp., third instar, urogomphi;J, Pterostichussp., maxilla, dorsal aspect; arrow: lateral membranous notch; (A-G, Arndt 1991;H-J, Arndt 1993).

reduced or absent in many groups (e. g., Harpal-inae, Brachininae).

Thorax with tergites medially divided by nar-row median ecdysial suture; mesonotum andmetanotum with anterior keel; pronotum largerthan following segments. Large spiracle presentbetween pro- and mesothorax. Legs 5-segmented(coxa, trochanter, femur, tibia, tarsus), usuallywith two claws. Four small sclerites present later-ally and dorsally of coxal base (episternum, epi-meron, trochantin, pleurite, the latter two arelacking on the prothorax). Abdominal tergitesI - VIII similar in structure, with anterior keeland median ecdysial suture; ventral and lateralsides with sternites, epipleuron, hypopleuron anda lateral spiracle; sternites consisting of one large(medio-) sternite and smaller paired anterior,inner and outer (Iatero-) sternites (Fig. 7.8.5 D).Abdominal segment IX smaller, with fused ster-nites; tergite IX usually with a pair of urogomphi(Fig. 7.8.6C-E, H, I); urogomphi long and slen-der, moveably attached to tergite and unseg-mented, or segmented and/or fused with tergite,or reduced. Tergite X (pygopod) cylindrical, di-rected downwards (Fig. 7.8.5 E).

First instar larvae with characteristic chaeto-taxy (Bousquet & Goulet 1984; Arndt 1993).Epicranial sclerites with ten dorsolateral andseven ventral setae; frontale with eleven setae,two on anterolateral margin and one very dis-tinctive long seta in anterolateral angle. Anten-nomere III with three long setae, antennomereIV with four long apical setae. Mandible with

one strong seta on outer margin. Cardo with onedistinctive seta; stipes with two large setae onouter side and a field of setae on inner side; laci-nia with one seta subapically (even distinct onstrongly reduced lacinia); palpifer with a distinc-tive seta on the ventral side; setae on galea andpalpi very small. Prementum including ligula pri-marily with seven pairs of setae, but number al-most always reduced or increased; ventral sidewith a pair of very small basal setae, a largecentral seta, and a pit on the apical part of theligula (distinctive in most species). Urogomphiwith five long setae in first instar and nine longsetae in the following instars.

Larvae of several groups of Carabidae arestrongly modified morphologically (e. g., legs re-duced, hyperprognathous, physogastric; seebelow).

The following references summa rise informa-tion on the larval morphology of Carabidae: vanEmden (1942), Bousquet & Goulet (1984), Arndt(1991 a, 1993), Beutel (1993), Luff (1993).

Paussinae (= Metriinae + Ozaeninae + Pro-topaussinae + Paussinae)

Distribution. All zoogeographic regions, butmost diverse in the tropics. Only few species oc-cur in southern parts of the Northern Hemi-sphere. The Metriini are restricted with one ge-nus and two species to western North America.Ozaenini occur with about 14 genera pantropicaland in southern North America, the Protopaus-


sini are restricted with one genus and seven spe-cies to the eastern Oriental region. The Paussiniare the most diverse and widespread tribe.

Bio]ogyand Eco]ogy. Higher paussines (= Proto-paussinae + Paussinae) are characterized by theiradvanced myrmecophi]ous habits, at least in thelarval stage. This unusual life style may haveoriginated from specialised predatory habits oflarvae, which are a ground plan feature of thesubfamily (Nagel 1979). Larvae of Metriini (Met-rius) and Ozaenini produce gland secretions attheir highly modified terminal abdominal seg-ments, which attract other insects. These secre-tions are apparently also attractive for ants, andmay have enabled paussine larvae to enter antsnests as parasites. Adults of Paussini are adaptedto these hostile environments in different ways(defiant type, symphilous type). Myrmecophiloushabits have independently evolved in Physea(Ozaenini).

Morphology, Adults (Darlington 1950; Nagel]997). Colour black (e. g., Metrius) or brown(most Ozaenini, Protopaussini and Paussini).Body laterally rounded (Metrius, Mystropomus)or parallel-sided. Fixed setae (i. e., long mecha-noreceptive setae) present or absent in Ozaenini,always absent in Protopaussini and Paussini. An-tennae filiform or moniliform in Metrius, Ozae-nini, Protopaussus and Megalopaussus. Stronglymodified in other subgroups of Paussini, withpedicellus very small, deeply sunken into apex ofscapus, connected with antennomere 3 by rigidwedging, and flagellomeres broadened and oftentransformed into a compact club. Procoxal pro-cess elongate and tapering posteriorly in Metrius,more or less truncate and posterolaterally con-nected with pleural process or strongly reducedin other paussines. Procoxal cavities alwaysclosed. Both pro tibial spurs in apical position ifpresent, usually obsolete or absent in Paussini.Antenna cleaning organ extended towards baseof tibia or reduced (Paussini). Hind walls of pro-coxae contact anterior excavations of meso-ventrite in Metrius. Connection between pro-and mesoventrite of harpaline type in all otherpaussines, with smooth anterior collar of meso-ventrite and without hexagonal groove. Meso-coxae separate or contiguous. Metacoxae sepa-rate in Metrius and Ozaenini (almost contiguousin Physea), contiguous in Protopaussini andPaussini. Termina] abdominal segment coveredby elytra (Metrius, most Ozaenini, Protopaussus),exposed in Dhanya and most Paussini. Speciali-sed abdominal defense secretion delivery systempresent. Subapical elytral flange present inPaussinae excl. Metrius.

Higher paussines with labrum often extendedlaterally or longitudinally. Mandibles compara-tive]y small and simple. Maxillary and labial

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pa]pi very variable. Mentum usually reduced insize and retracted or obliterated medially. Pro-thorax usually strongly modified. Prothoracicglands almost always present, usually associatedwith concavities or clefts and trichome systems.The trichome system is specialised in Protopaus-sus (Nage] 1997) and absent in Arthropterus andsome other genera (e. g., Homopterus, Carabido-memnus). Prosternal process fairly broad andposteriorly truncate in Protopaussus, distinctly orcompletely reduced in Paussini. Procoxae usuallyvery prominent and contiguous or almost contig-uous (Paussitae sensu Nage] 1997). Femora andtibiae of fore-, midd]e- and hind legs usuallybroadened and flattened (not in Pleuropterus).Protibial antenna cleaning organ vestigial or ab-sent. Protarsomeres usually cylindrical, in somecases partly or completely retractile (Ho-mopterus). Elytra without lateral edge. Meta-ventrite usually without discrimen and transversesuture. Median cell of alae usually triangular(not in Protopaussus). Abdominal sternites 11-IV fused, sutures between sternites faintly indi-cated or absent. Stridulatory file on the ventralabdominal base present or absent. Terminal ab-dominal segments partly exposed and sclero-tised. Parameres slender and asymmetric in ad-vanced paussines.

Morpho]ogy, Larvae (Fig. 7.8.7 A -G). Larvaelive in burrows in soil or in ants nests and arestrongly modified morphologically. Head andposterior part of body bent upwards (dorsad)(Fig. 7.8.7. G). Head capsule s]ightly or distinctlyrounded ]aterally; stemmata reduced to oneocellus or lacking; coronal suture absent (Fig.7.8.7 A); posterior tentorial grooves representedby a single narrow cleft immediately adjacent toa deep posteromedian emargination of theventral wall of the head capsule in Paussini(Arndt & Beutel 1994); nasale present in Metriusbut absent in other groups; fronta]e with mem-branous anterior margin in Paussini (Arndt &Beutel ]994); mandible (Fig. 7.8.7 B) broad atbase, with s]ightly sclerotized appendage (Iaciniamobilis) proximal to retinaculum in Paussini;maxillae with 4-segmented palpi in Metriini but3-segmented palpi in Ozaenini and Paussini;palpomere I more or less comp]etely fused withstipes in the latter group; galea I-segmented(Arndt & Beute] 1994), blade-like in Metriini(Beutel 1992); lacinia absent in Paussini but longand blade-like in Metriini. Prementum elongate,with tubercles. Paussini with posterior tentorialarms originating from narrow, common medianstalk, immediately adjacent with posteromedianemargination of head capsule, with dorsa] armsstrongly flattened and exceptionally short(Arndt & Beutel 1994). Regular fringe of longerhairs absent from cranial part of hypopharynx in

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A...

B

Fig. 7.8.7A, Paussus sp., head capsule, dorsal aspect(Arndt & Beutel 1995); B, Paussus sp., mandible, dorsalaspect (Arndt & Beutel 1995); C- F, Transformation seriesof posterior part of abdomen in representatives of Pauss-inae. C, Metrius contractus (redrawn from Bousquet1986); D, Pachyteles sp. (redrawn from Costa et al. 1988);E, Physea setosa (redrawn from Bousquet 1986); F, Paus-sus sp. (Arndt 1998 a); G, Metrius contractus, habitus, firstinstar (Bousquet 1986).

Paussini (specialised feeding habits); present butsparse in Metriini.

Legs primarily 6-segmented with two claws(anterior claw longer in Metriini and Ozaenini);tibia and tarsus fused to one strongly sclerotizedsegment with numerous strong setae on theventral side and a single small hook-like clawin Paussini. Posterior abdominal segments andurogomphi strongly modified. Epipleurites IX


greatly enlarged and forming a vertically ori-ented anal plate together with tergum VIII. Uro-gomphi branched (Metriini, Ozaenini) or flat-tened, triangular and integrated into anal plate(Paussini) (Fig.7.8.7C-F) (Bousquet 1986;Arndt & Beutel 1994; Vigna-Taglianti et al.1998).

Gehringiinae

Distribution. The only known species, Gehringiaolympiaca, was described by Darlington (1933).It occurs in north-western North America.

Biology and Ecology. G. olympiaca occurs ongravel banks of small to moderate mountainbrooks with cold water. It lives in the moistgravel and coarse sand and avoids coming to thesurface. Oviposition takes place in early summer.

Morphology, Adults (Fig.7.8.2A-D; Lindroth1969; Beutel 1992). 1.6-1.7 mm long, black, up-per surface covered with scattered setigerouspunctures. Compound eyes with large, convexfacets. Antennae short, moniliform, pubescentexcept on scapus. Distal antennomeres trans-verse. Ultimate maxillary palpomere small, subu-late. Pronotum distinctly narrowed posteriorly.Prosternal process short and narrow, not extend-ing beyond hind margin of procoxae posteriorly.Procoxal cavities open. Protibial cleaning organpresent, elongated towards proximal part oftibia. Spurs both in terminal position. Protarsinot dilated in males, proximal three tarsomereswith sparse adhesive setae. Mesoventrite of cara-bine type. Mesocoxal cavity disjunct. Elytra withunderlapping edges (Lindroth 1961- 69), withdeep sutural (and often faintly suggested 2nd and3rd) stria, otherwise with sparse punctures arran-ged in irregular rows. Apex truncate, exposinglast tergite. Metaventrite broad, with shortdiscrimen and laterally obsolete transverse su-ture. Metepimeron parallel-sided and exposed,not lobate. Metacoxae distinctly separate. Alaewell developed, with marginal fringe of hairs.Abdomen without separate sclerite betweenmetacoxae. Terminal sternite VII with almosttruncate, very slightly convex hind margin. Para-meres slightly asymmetric.

Morphology, Larvae. Length of specimen de-scribed by Lindroth (1960) 1.2 mm. Habitus oflarvae and characters generally very similar tothose of Trechinae. Body slightly sclerotized, ab-dominal tergites and sternites not apparent.Head almost square, parallel-sided; sutures in-distinct. Frontale distinctly extended posteriorly;coronal suture short; postocular groove absent;cervical groove absent. Egg-bursters consist oftwo rows of 5-7 minute tubercles on posteriorfrontale. Stemmata absent. Nasale with slightly


bisinuate anterior margin, slightly produced inadvance of inconspicuous adnasale, with foursmall teeth alternating with minute setae. An-tenna with three stout basal segments and slen-der antennomere IV; antennomere III with mesalspine and stalked, globulous sensorial append-age. Mandible sickle-shaped, with one lateralseta and well developed pointed retinaculum;penicillum absent; mesal edge of stipes with onlyone seta; 2-segmented galea with very slenderdistal segment; lacinia absent; basal palpomerewith twisted seta. Prementum subquadrate; palpislender, widely separated; ligula absent. Hypo-pharynx completely flattened. Prothorax dis-tinctly larger than meso- and meta thorax. Legsshort, with equal claws. Urogomphi fused withtergite IX, unsegmented, number of setae re-duced (as in Trechinae). Pygopodium longerthan urogomphi.

Omophroninae

Distribution. Represented by one genus with 60species in the Holarctic, Oriental and Afrotropi-cal regions, as well as Central America.

Biology and Ecology. All species are stronglyhygrophilous and restricted to the immediatevicinity of water. The adults are nocturnal andrun very rapidly when hunting at night. Like thelarvae, they spend the day in burrows in sandor clay.

Morphology, Adults (Fig. 7.8.1 B; Lindroth 1969;Beutel 1992). Characterised by an almost circu-lar outline, long and slender antennae and legs,and an unusual colour pattern with a paleground colour and darker, often metallic mark-ings. Head short and transverse. Prosternal pro-cess broad and apically truncate, covering themesoventrite entirely, thus joining the prothoraximmovably to the pterothorax. Procoxal cavitiesclosed, but pro coxae contact the anterolateralconcavities of the mesoventrite. Protibia of ani-sochaetous type, antennal cleaner distinctly elon-gated towards proximal part of tibia. Two proxi-mal protarsomeres of males strongly dilated.Proximal mesotarsomere moderately dilated.Scutellar shield concealed. Elytra with 14-15striae, but without abbreviated sutural stria.Metepimeron indistinct. Alae well developed.Median lobe sclerotized only in ventral half.Parameres moderately asymmetric.

Morphology, Larvae (Fig. 7.8.6 F -G). With fos-sorial adaptations. Body tapering posteriorly.Head capsule comparatively large and pro-nouncedly wedge-shaped, with strongly protrud-ing, triangular nasale. Stemmata large. Postocu-lar and cervical grooves lacking. Antennae con-spicuously held upward, inclined towards me-

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dian line, forming part of a prey-grasping basket.Mandible with single cutting edge and bidentateretinaculum; penicillus absent (Fig. 7.8.6 F). Pre-epipharynx with V-shaped, sclerotized bar; hypo-pharyngeal bulge reduced. M. tentoriohypopha-ryngalis medialis absent. Maxilla with very longlacinia and almost equally long 2-segmented ga-lea. Prementum small; ligula long, moderatelywide at base and extended as digitiform pro-jection apically (Fig. 7.8.6 G). Ligula and labialpalps subequal in length. Posterior tentorialgrooves V-shaped, shifted to posterior margin ofhead capsule. Tentorial bridge absent. Pro thoraxbroad basally, narrowed toward head, widerthan other segments. Legs rather long, withshortened tibia; trochanter, femur and tibia withapical whirls of stout setae; claws subequal, eachwith a long seta. Abdominal segments with nu-merous long setae. Hypopleurites not apparentin first instar. Urogomphi fused to tergite IX, notsegmented, about 3 times longer than tergite IX.Apex of urogomphi formed by a hyaline append-age in first instar (Landry & Bousquet 1984).

Cicindelinae

Distribution. World-wide, with five tribes, ap-proximately 130 genera and 2000 species. TheCtenostomatini comprise two genera, Pogonos-loma in Madagascar, and Clenosloma in tropicalSouth America. The Collyrini are restricted withfew genera to India and South East Asia, theManticorini are restricted with two genera toSouthern Africa. Megacephalini and Cicindeliniare the most diverse and widespread tribes.

Biology and Ecology. Adults of several genera arecharacterised by diurnal habits, very rapid ter-restriallocomotion, and excellent flying abilities.The larvae life in vertical or horizontal burrowsin soil, sandy substrates or rotten wood and arepredators with a highly specialised ambush strat-egy and unusual morphological adaptations.

Morphology, Adults (Fig. 7.8.1 A, 7.8.2E-F).10-70mm. Body black in basal groups (e. g., Omus,Amblycheila), with comparatively stout legs.Other genera with conspicuous colour pattern,metallic areas and long and very slender legs(Cicindela, Megacephala) or a very slender, ant-like body (arboricolous genera). Compound eyeslarge and protruding in diurnal cicindelines (e. g.,Cicindela auct.). Labrum large and broadened.Antennae inserted on dorsal side of head. Man-dibles with several pointed apices, long, inter-crossing in resting position. Prothorax elon-gated, prosternal process and postcoxal bridgeunusually broad. Antennal cleaner and protibialspurs terminal. Elytra without stria. Alae com-pletely reduced (Omus) or well developed. Ob-longum absent. Thoracic segments otherwise

130 Erik Arndt, Rolf G. Beutel & Kipling Will

Fig. 7.8.8A-C, Platychilepallida, first instar, dorsal aspect (Arndt 1998b). A, head capsule; B, nasaIe;C, maxilla, sb -sclerotized bar; D- E, Eucalliaboussingaulltii(redrawn from Arndt et al. 1996). D, labium; E, abdominal tergite V, ih -inner hook, mh - median hook, oh - outer hook.

similar to Elaphrinae and Loricerinae. Para-meres symmetrical, connected by jugal sc1erite.

Morphology, Larvae (Arndt 1998b; Arndt &Putchkov 1997; Breyer 1989; Hamilton 1925;Knisley & Pearson 1984; Putchkov & Arndt1994) (Fig. 7.8.8). Strongly modified morpho-logically in correlation with their life habits.Head and pronotum strongly enlarged andstrongly sc1erotized in contrast to long and slen-der rest of body. Dorsal side of hyperprogna-thous head and protergum form a functionalunit. Both parts together form a lid of the bur-

rows prior to the capture of prey. Head stronglyrounded laterally; ventral side strongly convex.Six stemmata of different size present; two pairson dorsal side of head strongly enlarged (Fig.7.8.8 A). Frons distinctly extended posteriorly,coronal suture very short or absent. Posterodor-sal margin of head capsule emarginate, thusnearly reaching or reaching posterior margin offrontale. Ridge on caudal part of parietale con-nected with ridge on caudal part of frontale inManticorini and Megacephalini, but separate inother taxa. Nasale (Fig. 7.8.8 B) strongly pro-truding, shovel-shaped, with anterior margin


smooth and subtruncate or sinuate. Antennaewith elongate, nearly equally sized antennomeres1- 3. Antennomere 4 slightly shorter. Sensorialappendage of antennomere 3 replaced by smallfield of pores. Mandible slender; apical partlonger than basal part including triangular reti-naculum; penicillus absent. M. craniomandibu-laris internus unusually large and complex. Addi-tional protergal muscle inserts on adductor ten-don (Cicindela; Breyer 1989). Cardo triangular;stipes slender, curved, strongly sclerotized, withone or more spines on mesobasal margin and amembranous field on dorsal side; lacinia smallor absent; strongly sclerotized bar with threebristles present between stipes and palpomere 1;palpomere 1 large, fused with or at least attachedto galeomere; galea as large as maxillary palp;both galeomeres with thick bristles (Fig. 7.8.8 C).Prementum dorsally covered by multisetosehypopharynx; ligula prominent, with two pairsof setae; bipartite ventral sclerite between pre-mentum and labial palpomere 1 (Fig. 7.8.8 D)present or absent (Megacephalini and Manticor-ini). Premental retractors very strong, arrangedin an unusual manner (Cicindela; Breyer 1989).Dense preoral filter present, apparently arisingfrom upper part of ligula (Cicindela; Breyer1989: Fig. 5). Hypopharynx separated form dor-sal premental surface by a distinct fold but al-most completely flattened. M. tentoriohypopha-ryngalis absent. Gular suture anteriorly limitedby an Y-shaped posterior tentorial groove in Ci-cindelini; posterior tentorial groove T-shaped inall other taxa. Most parts of tentorium stronglyflattened (dorsal arms) or thin; posterior armsbasally fused; thin U-shaped tentorial bridgewith attached to posterodorsal margin of headcapsule by thin processes; anterior arms stronglydeveloped, broadly connected with head capsule(Cicindela; Breyer 1989). Pronotum rhomboid(Fig. 7.8.4 B), distinctly different from smaller,laterally rounded meso- and metanota. Legsstout, with very short tarsus; anterior claw largerthan posterior claw. Abdominal segments I-IVand VI - IX subequal, but segment V distinctlymodified, dorsally enlarged as an "abdominalhump", with separate anterior-, lateral- and cau-dal sclerites and 2- 3 hooks inserted betweenthem (Fig. 7.8.8 E). Urogomphi absent; pygopodshort, conical and multisetose. Chaetotaxystrongly modified. Unusual, flattened, split setaeon head and pronotum present in many taxa.

The following characters distinguish Manti-corini from other larvae of Cicindelinae: anten-nal base inserted anteroventrad of stemmata andseparate from base of mandible by wide sclero-tized area; antennomere I very thin; tibia dis-tinctly curved with a field of numerous short se-tae on posterolateral side; number of stemmatareduced, two enlarged stemmat a situated onprominent region of head capsule (Oberprieler &

131

Arndt 2000). Larvae of Collyrini and Ctenosto-matini live in rotten wood contrary to all othergroups and are characterised by several apo-morphies: width of nasale reduced, body flat-tened, and (except in Pogonostoma) claws fusedwith tarsus (Arndt & Putchkov 1997).

Carabinae

The subfamily comprises 9 tribes, four of which(Carabini, Ceroglossini, Pamborini, and Cych-rini) contain large and colourful species. Theadults, especially of the largest genus Carabus,are often characterised by metallic cuticle withconspicuous elytral patterns. The group is para-or polyphyletic, comprising with Cicindini, oneof the most peculiar and enigmatic tribes.

Distribution. World-wide, but the great majorityof species live in the Northern Hemisphere. Themonotypic Notiokasiini are restricted to thewarm-temperate Neotropical region, Ceroglos-sini to the Neotropical Andes, Pamborini to Aus-tralia and New Zealand. The Cicindini containtwo monotypic genera, Cicindis from Argentinaand Archeocindis from Persian Gulf.

Biology and Ecology. Most representatives in-cluding Carabus and Ceroglossus live on theground as more or less generalized predators,and nearly all of them are flightless. Adults andlarvae of Cychrini are specialized predators ofsnails. Adults and larvae of Calosoma are veryactive predators of caterpillars and adults andlarvae of some species climb trees. Mass flightsto caterpillar outbreaks are recorded e. g., forCalosoma jrigidum. Larvae of Notiophilus arehighly specialised predators of springtails.

Morphology, Adults. Size ranging from small(e. g., Notiophilus) to very large (e. g., Carabussubgenus Procerus). Protibial spurs and antennacleaning organ terminal (Opisthiini, Carabini,Cychrini) or one spur subterminal and antennacleaning organ slightly prolonged proximally.Prosternal process distinctly extending beyondhind margin of procoxae and apically tapering.Procoxal cavities open. Mesoventrite of carabinetype. Metepimeron concealed.

Morphology, Larvae (Fig. 7.8.4 A, 7.8.5 D). Lar-vae of Carabinae are remarkably diverse mor-phologically. None of the features which charac-terize the subfamily as a whole are autapo-morphic: cervical and ocular grooves absent; hy-podon (central tooth) usually present (absent inOpisthius); antennal muscles not intercrossing;mandible usually with penicillum; hypopharynxdistinctly bulging, with preoral filter; M. tentori-ohypopharyngalis medialis usually present (ten-

132

torial bridge interrupted and M. 42m absent inNebria); Mm. vertiCopharyngalis and tentorio-pharyngalis well developed (Beutel 1992).

Ceroglossini, Carabini, Pamborini and Cych-rini are characterised by strong sclerotization, re-duced number of setae on tergites and sternites,a remarkably increased number of pores, and areduced sensorial appendage of antennomere 3(Arndt 1998a; Priiser & Arndt 1995). The uro-gomphi are markedly sclerotized, shortened,fused with tergite IX and increasingly reduced inthe latter three taxa. The tergites are often ex-tended laterally (e. g., Cychrini). Nasal teeth andsetae FRIO,II are absent in Cychrini and manyspecies of Carabus (Arndt & Makarov 2003).Larvae of Nebriini, Opisthiini, and Notiophiliniare characterized by reduced setae TE6, PRs andMEz, and the presence of a roughly pointedmicrosculpture on all tergites. The urogomphiare thin, straight and moveably attached to ter-gite IX (Arndt 1993). Antennomere III is en-larged and multisetose in Opisthiini (Bousquet &Smetana 1991). A strongly protruding nasal re-gion with long, sharp and prominent teeth ischaracteristiC for Notiophilini and some taxa ofNebriini. A sclerotized bar is present between thestipes and maxillary palpomere I in Notiophilini(as in Cicindelinae; Arndt 1993).

Hiletinae

Distribution. Hiletinae (Hiletini) are a tropicalgroup considered as rare and enigmatic (Erwin &Stork 1985). Approximately 20 known speciesare arranged in 2 genera, Hiletus and Eucamara-gnathus. Hiletus species occur in tropical Africaand species of Eucamaragnathus in Africa,Madagascar, south-eastern Asia, and in SouthAmerica east of the Andes from Ecuador to low-land eastern Brazil (Erwin & Stork 1985).

Biology and Ecology. All species frequent lato-solic soils in broadleaf evergreen and deciduousforests or in grassland savannahs with scatteredtrees.

Morphology, Adults (Erwin & Stork 1985; Beutel1992). Head large and robust in proportion topronotum, with transverse sulcus whiChconnectsthe sulcate frontal furrows. Antennae geniculate,with scapus as long as antennomeres 2-4. Sca-pus fits in groove below compound eyes. Mandi-bles large and markedly down turned distally,creating a hollow concavity beneath. Each man-dible with eight or nine triangular teeth, whichincrease in size distally. Lacinia enlarged later-ally, resembling an asymmetric club. Dorsal edgewith numerous spatulate spiculae. Galea longand finger-like. Mentum with deep median con-cavity, anteromesally produced into simple (Eu-


camaragnathus) or bifurcate (Hiletus) projec-tions. Profemora of males almost always withventral tooth (not in Southeast Asian Eucamara-gnathus). Protibia of anisochaetous type. Anten-nal cleaner poorly developed. Tarsi of all legsslender and distally tapering. Adhesive setaepresent on protarsomeres 1-3 and mesotarso-meres 1- 2. Prosternal process long and narrow-ing posteriorly. Procoxal cavity open. Meso-ventrite with hexagonal and anterolateralgrooves (carabine type). Mesocoxal cavities ofdisjunct type. Elytral pattern diverse. Metepim-eron lobate. Alae well developed, with anteriorsector cell much larger than 3rd radial cell. Me-dian lobe unspecialised. Parameres asymmetriC,usually brushy or multisetiferous (not in E. bra-siliensis).

The thoracic structures display a unique com-bination of "primitive" (prosternal process, pro-coxal cavity, mesoventrite) and derived features(lobate metepimeron).

Larvae. unknown

Loricerinae

Distribution. One tribe with two genera, the Hol-arctic Loricera (9 spp.) and the monotypic Ellip-tosoma (Madeira).

Biology and Ecology. Species of Loricera aremore or less hygrophilous and characterised byspecialised prey-catching techniques of larvaeand adults, which involve the modified antennaeor maxillae, respectively. They feed on spring-tails.

Morphology, Adults (Jeannel 1941-42; Lindroth1961-69). In general outline resembling a mid-dle-sized Agonum (Loricera). Frons with 2 largefoveae and a posterior median sulcus, emanatingfrom a deep transverse constriction behind theeyes. Antennae with long scapus and very long,erect setae on pedicellus and antennomeres 3-6.Protibia of anisochaetous type. Procoxal cavityposteriorly closed by narrow bridge. Protho-racic-mesothoracic connection of harpaline type.Mesocoxal cavity of disjunct type. Elytra with 12regular stria (without abbreviated sutural stria).Metepimeron parallel-sided and narrow. Copula-tory organ short and oval, median lobe withwide open basal orifice and elongated, asymmet-riC apical part. Parameres apically rounded,without fringes of hairs. Left paramere longer.

Morphology, Larvae (Arndt 1993, Luff 1993)(Fig. 7.8.9 A-B). Head capsule rounded later-ally. Cervical and ocular grooves absent. Nasalregion with two acute teeth and a row of smallsharp teeth beneath them. Antenna 2 X longerthan mandibles. Mandible siCkle-shaped; retina-


o

A B

133

oFig. 7.8.9A- B, Loricera pilicornis, third instar (redrawn from Arndt 1991). A, mandible; B, maxilla; C- D, Siagona sp.(Grebennikov 1999 a); A, habitus. B, antenna.

culum large, serrate along inner edge; penicillumpresent (Fig. 7.8.9 A). Stipes very large, almostentirely sclerotized; galea mesally directed atright angle to stipes, longer than maxillary palp;apical galeomere with divided into a swollen,finely granular basal portion and a long, whip-like apical portion; distal part covered by hyalinesecretion (Fig. 7.8.9 B); field of setae on maxil-lary palp reduced; stipital field of setae reduced,its vestiges shifted to the middle of the innermargin. Labial palp slender, with secondarilysubdivided apical palpomere (Loricera sp.;Thompson 1979); ligula unusually large, broadlyrounded anteriorly, multisetose. Chaetotaxy offrontale and tergites modified; flattened, split se-tae on pronotum, mesonotum and metanotumpresent (see Cicindelinae). Legs long; tarsi elon-gated, with 2 unequal claws; anterior claw larger,with basal seta; seta absent from posterior claw.Urogomphi slender and nearly as long as headand thorax combined, fused to medially dividedtergite IX, in later instars with numerous setifer-ous nodes. Pygopod short and conical.

Elaphrinae

Distribution. One tribe comprising 3 genera, Di-acheila, Blethisa and Elaphrus, restricted to theHolarctic region.

Biology and Ecology. Elaphrines are usuallystrongly hygrophilous, and often riparian andvery active during the daylight.

Morphology, Adults (Fig. 7.8.3; Jeannel 1941-42; Lindroth 1961-69; Beutel 1992). Middle-sized, always with metallic lustre. Eyes large andprominent. Prothorax with coarse punctuation.

Protarsomeres I - IV or I- III dilated in males.

Elytral striae feeble, irregular, usually disturbedby fovea or tubercles. Thoracic structures other-wise similar to Loricerinae. Copulatory organwith a long sclerotized stylet protruding throughthe basal orifice of the median lobe when in re-pose. Parameres with fringes of hairs at the apexand ventral edge, less dense on the left paramerein Elaphrus.

Morphology, Larvae (Goulet 1983, Luff 1993)(Fig. 7.8.6 D- E). Body slender and subcylindri-cal (Luff 1993: Fig. 21). Head without ocular andcervical grooves (Luff 1993: Fig. 22). Coronal su-ture distinct, moderately long or short. Nasalecentrally produced, triangular, with serrate(Elaphrus) or smooth (Blethisa) lateral edges.Mandible with large retinaculum and penicillus.Cutting edge of mandible finely serrate in manyspecies. Lacinia reduced, very short or ring-shaped. Pro thorax largest segment of body; pro-notum quadrangular. Other segments laterallyrounded. Equal claws elongate, more than 0.5 aslong as tarsus. Urogomphi fused with tergite IX,in second and third instars multisetose with setaeinserting on small tubercles or with 10 long setaeon very large nodes (Fig. 7.8.6 D- E).

Migadopinae

Distribution. A relatively small group of twotribes with 15 genera mainly distributed in thesubantarctic region (Andes, southern part ofSouth America, Falkland Islands, southern Aus-tralia and Tasmania, Southern New Zealand,Auckland Islands) (Jeannel 1938; Moret 1990;Baehr 1999).

134

Biology and Ecology. Most species of this tribelive in cool and south temparate rain forests andmoorland. They live on the ground, and areflightless with the exception of one species.

Morphology, Adults (Jeannel 1938). Mediumsized beetles ranging from 6-20 mm, with gla-brous surface. Almost always with one supraor-bital seta (absent in Aquilex diabolicola; Moret1990). Pronotum and elytral disc without setae.Elytra with one additional stria between Isl and2nd primary striae. Wings always absent. Tho-racic structures otherwise similar to Loricerinae.Parameres well developed, at least right para-mere fringed with hairs.

Morphology, Larvae (Johns 1974). Three dif-ferent larvae of Loxomerus and the larva of anundetermined genus were described so far (Johns1974). Internal features are unknown. Head withmore or less constricted neck region. Ocular andcervical groove absent. Coronal suture very long.Nasale with three or five teeth including a hypo-don; mandible straight and slender, with largeretinaculum; retinaculum with additional sub-basal tooth; penicillus present; maxilla with sti-pes, palpi and galea long and slender; lacinia ab-sent. Prementum wide; ligula small. Legs withtwo claws, the anterior longer than the posteriorone. Urogomphi short, articulating on membra-nous region of tergite IX. Hypopleurite with cap-like gland on dorsal margin in second and thirdinstar larvae (absent in first instar).

Siagoninae

A small group comprising two tribes, Enceladinirepresented by the monotypic genus Enceladus(northern South America) and Siagonini re-presented by the genera Siagona (southern Palae-arctic including Malay Archipelago and Philip-pines, Africa; 80 species), Cymbionotum (south-ern Palaearctic excluding Malay Archipelago,Africa) and Luperca (Africa, India; two species)(Darlington 1967; Erwin 1978).

Biology and Ecology. Species of Cymbionotumand Siagona live on the ground in wet areas withdecaying vegetation or like Enceladus underbark. Some occur on mud and in soil clefts ofdesiccating waters. Luperca were found in termi-taria.

Morphology, Adults (Jeannel 1941-42). Moder-ately depressed and glabrous (Enceladus, Luper-ca) or strongly depressed and pubescent (Sia-gona). Body always strongly constricted betweenpro- and mesothorax. Mandibles short, withoutseta. Palps short and narrow. Prothorax stronglynarrowed posteriorly. Pro thoracic-meso thoracicconnection of harpaline type. Mesocoxal cavities


disjunct. Elytra distinctly flattened, withoutbasal margin, with 8 stria (indistinct in Siagona).Metepimeron parallel-sided, not lobate. Aedea-gus large, laterally compressed. Parameres de-void of hairs in Luperca and Enceladus.

Morphology, Larvae (Fig. 7.8.9 C- D) (Greben-nikov 1999a). The larvae of Siagoninae are char-acterised by many apomorphic features. Headwith more or less constricted neck; cervicalgroove absent (Siagona, Fig. 7.8.9 C) or com-pleted to a dorsal parietal keel (Enceladus); ocu-lar groove lacking; coronal suture long; 6 stem-mata present or number of stemmata reduced totwo or one. Nasal region protruding, serrate,shovel-shaped; adnasale with a dense group ofsetae. Antennomere III without sensorial ap-pendage; antennomeres III and IV multisetosewith membranous area on ventral side or ex-tremely elongate and whip-like (Fig. 7.8.9 D).Mandible with large retinaculum; subapical setaMN2 longer than retinaculum; penicillus andseta MN I on outer mandibular margin absent.Maxillary and labial palpi strongly elongate (Sia-gona), or of normal size but with terminal seg-ments enlarged and with extended sensorialfields (Enceladus). Lacinia present. Ligulabroadly rounded. Legs slender, with two claws;setae of claws shifted to pretarsal sclerite. Ab-dominal tergite IX distinctly reduced, dividedmedially, fused with multisetose and extremelyelongate urogomphi (Fig. 7.8.9 C). Urogomphiabout 0.5-1.0 as long as remaining body.

Scaritinae

Scaritinae, which are probably not monophyletic(s. below), comprise small to very large species.They are are always strongly constricted betweenthe pro- and mesothorax.

Biology and Ecology. Scaritinae are mainly char-acterised by their habits of burrowing in soil orsandy substrates.

Distribution. Scaritini (sensu lato) occur worldwide, but most of taxa are concentrated in theSouthern Hemisphere. Promecognathini withfive genera are restricted to southernmost Africaand northwestern North America. Salcediini oc-cur with three genera in the pantropical region.

Morphology, Adults (Jeannel 1941-42). Dis-tinctly depressed (e. g., Scarites) or almost cylin-drical (e. g., Dyschirius). Pro thorax strongly nar-rowed posteriorly, thus body distinctly con-stricted between pro- and pterothorax. Protho-racic-mesothoracic connection of harpaline type.Mesocoxal cavities disjunct. Metepimeron paral-lel-sided, not lobate.


Morphology, Larvae (Fig. 7.8.6 H, I). One of themost heterogenous groups concerning larvalmorphology (Thompson & Allen 1974; Peyrieras1976; Thompson 1979; Nichols 1986; Bous-quet & Smetana 1986; Arndt 1991a; LufT 1993;Moore & Lawrence 1994).

Promecognathini: Only the first instar larva ofone species of Promecognathus is known and dif-fers strongly from all other taxa in this subfamily(Bousquet & Smetana 1986). Head quadrangu-lar, parallel-sided; stemmata reduced; cervicalgroove absent; coronal suture absent; posteriorangles of frontale bulging, extended to posteriormargin of head. Nasale straight, smooth, notprotruding; antenna slender; antennomere IIIwithout sensorial appendage but with sensorialfield apically on ventral side; mandible slender,with small retinaculum; penicillus lacking in con-trast to other known scaritine larvae; lacinia andligula lacking; palpi short and stout, urogomphiclub-like, attached to tergite IX by a membra-nous area; body and urogomphi multisetose evenin first instar.

Scaritine subtribe Carenina: Body stronglysclerotized; lateral margins of head subparallel;six stemmata present; coronal suture long; cervi-cal groove present; anterior margin of frontalesmooth, straight, nasale not protruding; anten-nae extremely short; mandible stout, withoutretinaculum; lacinia elongate, as long as galea;maxillary palpi modified, only penultimatearticle of normal shape; ligula present; labialpalpi short and wide, with very large terminalsensorial field; epipleurites and hypopleuritesfused on all abdominal segments; urogomphi ab-sent (Moore & Lawrence 1994).

Subtribe Pasimachina: Body slender, subparal-lel, strongly sclerotized and dark; head widerthan long; six stemmata present; cervical groovelacking; coronal suture short; nasale protruding,shovel-shaped, with smooth anterior margin; an-tenna long and slender, with sensorial appendageabsent from article III; mandible slender withsmall retinaculum; lacinia and ligula absent; bothpalpi of normal structure; urogomphi fused totergite IX, turned inside, longer than pygopod(Thompson & Allen 1974).

Subtribe Scaritina: Larvae strongly sclero-tized, slender, elongate, subparallel; head quad-rangular; coronal suture very long; cervicalgroove shallow; nasale trapezoid; antenna longand slender with antennomere 3 flattened latero-apically, sensorial field present but appendageabsent on antennomere III; mandible long andslender; lacinia absent; galea and palpi slender,of normal structure; ligula present; urogomphislender, fused with tergite IX (Peyrieras 1976;Thompson 1979; Nichols 1986; Arndt 1991a).

Clivina and related groups: Body yellowish,not strongly sclerotized; stemmata absent; coro-nal suture and cervical groove present; nasale

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wide, with several teeth, slightly protruding; an-tenna short and stout; antennomere III with sen-sorial appendage; mandible slender, with smallretinaculum; finely serrate on inner margin; laci-nia and ligula present; legs as in Dyschirius andin contrast to all other known Scaritinae withsingle claw; urogomphi fused with tergite IX, aslong as pygidium, but flattened in dorso-ventraldirection (Fig. 7.8.6 H); muItisetose in secondand third instars or urogomphi and pygopodwith several large horns (Thompson 1979).

Dyschirius and related groups: Larvae stronglysclerotized, brown to black coloured, only headsometimes red. Head wider than long with sixstemmata, nasale protruding, crown-like, poste-rior angles of frontale wide or slender and long;antenna stout but sensorial appendage on anten-nomere III present; mandible slender with retina-culum present; lacinia absent; legs with a singleclaw; urogomphi short and stout, much shorterthan pygopod or tergite IX (Fig. 7.8.6 I; Arndt1991a; LufT 1993).

The larvae of several subtribes and of the tribeSalcediini are still unknown.

Trechinae

Trechinae are a much disputed problematicgroup and probably a paraphyletic assemblage(see below). It is unclear whether Patrobini, Apo-tomus, Broscini and Psydrini should be included.The inclusion of Melaenini and Cymbionotini(Lawrence & Newton 1995) is also problematic.

Distribution. World wide, a very diverse group.

Biology and Ecology. Highly variable, s. above.

Morphology, Adults. (Fig. 7.8.1 C; Jeannel 1941-42). Male protarsi often with unilaterally dilatedproximal tarsomeres. Mesepimeron broad ornarrow. Otherwise harpaline type character com-bination of thoracic sclerites, i. e. closed procoxalcavities, apically truncate prosternal process,harpaline type mesoventrite, conjunct mesocoxalcavities, lobate metepimeron.

Morphology, Larvae. The larvae of Trechinae areclose to the ground plan of "higher Carabidae"(Arndt 1993). Body moderately flattened, paral-lel-sided. Head roughly quadrangular; cervicalgroove present; head laterally with 6 stemmataarranged in two rows, number reduced in cavedwelling taxa. Ocular groove present; frontal su-ture sinuate; egg bursters consisting of a keel,isolated teeth or microspines. Nasale serrate,protruding. Antennomeres I and II subcylindri-cal; antennomere III with bulb-like sensorialappendage inserted laterally; antennomere IVsmaller, rounded apically. Mandibles with retina-culum; terebrum with single cutting edge,

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smooth, or serrate in some taxa of Anillini andBembidiini; penicillus present or absent. Stipesslender; lacinia absent. Prementum with ligula.Hypopharynx separated from dorsal surface,flattened, with dense field of microtrichiae. An-tennal muscles arranged crosswise (Beutel 1993).

Thorax with large pronotum and smaller, sub-equal meso- and metanotum, the latter two withanterior keel. Legs slender, with single claw in alltaxa except Perileptus, Amblystogenium, Thalas-sophilus, and Patrobini (s. Psydrini, Gehringiini).Lateral and dorsolateral area above coxal basewith four small sclerites (episternum, epimeron,trochantin, pleurite, the latter two are lacking onprothorax). Abdominal tergites I- VIII similar instructure; tergites with anterior keel and medianecdysial sutures; sternites consisting of one largemedian plate and smaller paired anterior, innerand outer (latero- )sternites. Abdominal segmentIX smaller, with fused sternal plates; tergite IXfused with urogomphi. Abdominal tergite Xcylindrical, directed downwards (pygopod).Number of setae on urogomphi in second andthird instars reduced, seta URj3 always lacking.Setae TA3-6 lacking in Trechini, Zolini, Bembi-diini (incl. Anillina), and Pogonini (Arndt et al.1999, Grebennikov 1999b; Arndt 2000; Greben-nikov & Maddison 2000). Larvae of Apotomini,Melaenini and Cymbiotonini are unknown.

Harpalinae

Distribution. World wide, the most diverse groupwith more than 30 tribes and 20000 species.

Biology and Ecology. Highly variable, s. above.

Morphology, Adults (Jeannel 1941-42). Mandi-bles without scrobal seta. Male pro tarsi neverwith unilaterally dilated proximal tarsomeres.Procoxal cavities uni- or biperforated. Elytracomplete or apically truncate (e. g., Odacan-thini, Lebiini).

Morphology, Larvae (Figs.7.8.4C-E, 7.8.6J).The tremendous number of species and supra-specific taxa included in this subfamily is re-flected by an extreme diversity of larval charac-ters. Taxa which are presumably close to theground plan of the subfamily, such as somePterostichini and Platynini, are similar to Trech-inae in most features described above, but havea short lacinia and double claws. All larvae ofthis subfamily are characterised by the reductionof the second ligular seta to a pore (Arndt 1993).The antennal muscles are arranged crosswise asin larvae of Trechinae and Brachininae (Beutel1993). A membranous band is present ventrolat-erally on the stipes in many representatives (Mo-rionini, Pterostichini, Zabrini, Panagaeini, Pe-leciini, Callistini, Oodini, Licinini, Harpalini;


Arndt 1993; Fig. 7.8.6J). The membranous notchis regarded as rudiment of a complete transversedivision of stipes as occuring in Cnemalobus(Roig-Jufient 1993).

The subgroups with most derived larval char-acters are the Licinini, Panagaeini and relatedgroups (van Emden 1942; Liebherr & Ball 1990;Arndt 1991b), and the so-called "Truncatipen-nia", e. g., Anthiini (Arndt & Paarmann 1999),Helluonini, Orthogoniini (Makarov 1998), Gra-phipterini (Zetto Brandmayr et al. 1993), Dryp-tini (Habu & Sadanaga 1965),Galeritini (Arndt &Drechsel 1998), and Lebiini (part.) (Lieftinck &Wiebes 1968;Arndt 1989;Capogreco 1989;Arndtet al. 2001).

Pseudomorphinae

Distribution. An unusual group of carabids dis-tributed in the Oriental region, Australia (Adelo-topus, Cryptocephalomorpha, Cainogenion, Paus-sotropus, Sphallomorpha) and the New World(Pseudomorpha).

Biology and Ecology. Pseudomorphine adults aremainly found under bark (Baehr 1994). Theknown larvae (except Sphallomorpha) are physo-gastric due to their myrmecophilous habits. Theknown larvae of Sphallomorpha dig holes in theground around ant nests and prey on ants in a sim-ilar manner like cicindeline larvae. Pseudo-morphines are the only known carabids withovoviviparity (Liebherr & Kavanaugh 1985).

Morphology, Adults (Notman 1925; Baehr 1992).Body form elongate and more or less parallel andmoderately convex (Pseudomorpha), very elon-gate and cylindrical, resembling Scolytidae (Ade-lotopus part.), or broadly oval and depressed(Sphallomorpha). Colour dark piceous to nearlyblack (Sphallomorpha part.) or variegated withmaculae, vittae or pale margins. Head deflexed(Cryptocephalomorpha) or horizontal, with deepantennal grooves. Compound eyes on dorsal sideof head (Adeloptopus, Cainogenion) or lateral,continuous border beneath eye present in Adeloto-pus. Scapus partly visible from above. Mandibleswithout visible scrobes, very small in Cryptoceph-alomorpha. Maxilla with (Cainogenion, Paussotro-pus) or without a large lateral lobe in most genera.Submento-mental suture absent. Prosternal pro-cess present or absent (Paussotropus). Legs short,with strongly developed femora. Configuration ofthoracic sclerites ofharpaline type (see above). Sixabdominal sternites visible. Parameres setose orglabrous, fairly symmetrical or highly asymmetri-cal but not 'balteate' (Baehr 1992).

Morphology, Larvae (Fig. 7.8.10) (Erwin 1981,Baehr 1997). Larvae with abdomen curved in lat-eral view and rows of spines on tergites V-VII


B

Fig. 7.S.IOA, Adelotopus dytiscoides, habitus, first instar;B, Adelotopusrubiginosus,head, first instar, dorsal aspect;C, ventral aspect (A-C, Baehr 1997).

(Sphallomorpha, Moore 1974) or physogastricalwith membranous, poorly sclerotized body(Fig. 7.8.IOA). Head subquadrate (Fig. 7.8.10B-C) or elongate and slightly narrowing anteriorly,without stemm ata and egg bursters. Frontalevery wide, frontal suture not sinuate, broadlyreaching posterior margin of head; coronal su-ture, ocular and cervical groove absent. Nasaleflat, convex or triangular, smooth, without teeth.Mandible without retinaculum or penicillus.Lacinia absent; galea 1- or 2-segmented, more orless reduced; maxillary palp 4-segmented, largein relation to other maxillary parts, longer thanstipes. Ligula broadly rounded or absent. Pro no-turn sclerotized; other tergites largely or entirelymembranous. Legs with double claws of dif-ferent length. Urogomphi absent. Chaetotaxystrongly modified, larvae often multisetose orwith large club-like, flattened or split setae onhead (Fig. 7.8.10 B-C), dorsal head appendages,thoracic and abdominal tergites (Baehr 1997; Er-win 1981).

Phylogeny and Taxonomy. A branching pattern(Sphallomorpha + (Pseudomorpha + (Crypto-cephalomorpha + (Adelotopus + (Cainogenion +Paussotropus»») was proposed by Baehr (1994).

Brachininae

Mainly characterised by a complex abdominalexplosive mechanism. A comparable apparatusis only present in Paussinae. However, distinctstructural differences in both groups were

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pointed out by Forsyth (1972) and others (seebelow).

Distribution. Crepidogastrini with 7 genera(more than 100 species) are distributed in tropi-cal and southern Africa, two species of Tyroniaoccur in India and Sri Lanka. Brachinini with 8genera (and more than 500 species) occur worldwide, but most of the species are distributed intropical and subtropical regions.

Biology and Ecology. The known larvae of Bra-chininae are parasitic.

Morphology, Adults (Fig. 7.8.1 D; Erwin 1970,Jeannel 1941-42; Lindroth 1961-69; Lieb-herr & Will 1998). Mandibles with a long seta atthe anterior part of the external scrobe. Protho-rax narrow, not or only slightly wider than head.Elytra very broad, abruptly truncate at apex,with (Brachinus) or without a pale, narrow mem-brane. Other thoracic features of harpaline type,with narrow mesepimeron. Abdomen with 7 vis-ible sternites in females and 8 visible sternites inmales. Copulatory apparatus with the medianlobe often deformed, the right paramere stronglyreduced, and the left paramere strongly sclero-tized. Female genital tract with appended sper-mathecal gland, segmented gonocoxae, gono-coxal rami absent, and in some taxa, a digitiformdiverticulum of the spermathecal duct present(Liebherr & Will 1998).

Morphology, Larvae (Fig.7.8.6A-C). There islittle known about larvae of Brachininae. OnlyBrachinini are known in the larval stage. Fewfirst instar larvae are described (see Arndt 1993for a review), and a single species is known in allinstars (Erwin 1967).

Following characters distinguish Brachininaefrom the ground plan of Carabidae and are re-garded as au tap omorphic character states: ante-rior margin of frontale membranous, teeth andsetae lacking. Posterior angle of frontale veryslender, V-shaped, egg bursters lacking in first in-star. Setal group gMX lacking, Lacinia absent(7.8.11 B). Ligula and setae LA3.4.6lacking. Uro-gomphi reduced or absent (7.8.11 C). Later instarlarvae physogastrical, legs and head appendages:t reduced (Arndt 1993). The antennal musclesare arranged crosswise, a feature shared by lar-vae of Brachininae, Harpalinae and Trech-inae(BeuteI1993). Other internal head structuresare similar to those of harpaline larvae exam-ined. Hypopharynx completely flattened. Func-tional mouth narrow. Mm. tentoriohypopharyn-galis, verticopharyngalis and tentoriopharyngalisabsent. The shift of the brain to the prothorax(Beutel 1993: Fig. 27) is a result of miniaturisa-tion.

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Phylogeny and Taxonomy. Phylogenetic hypothe-ses for Carabidae, originally simply the classifi-cation of presumed natural groups, have contin-ued to develop and be refined as advances intechnology have permitted researchers to usemore and finer details of behaviors, morphology,chemistry and gene sequence data. The extensiveknowledge of carabid beetles makes them wellsuited to cladistic methodology, which is morefrequently being used to rigorously test hypo-theses of relationships previously posited. Ball(1979) and Ball et al. (1998) provide an excellentdetailed review of the history of classificationand phylogenetics for the family. Brief and usefulsynopses with notes on relationships and distri-bution of each tribe were published by Bous-quet & Larochelle (1993).

Recent phylogenetic analyses that bear on re-lationships within Adephaga, affecting our un-derstanding of the root for the family, or for ex-emplars from across many tribal level taxa incarabids, include a variety of character systems.Notably, 18S rDNA sequence data (Maddisonet al. 1998, 1999; Shull et at. 2001), female repro-ductive tract (Liebherr & Will 1998), larval mor-phology (Arndt 1993, 1998), cuticular and mus-cular morphology (Beutel 1992, 1993; Beutel &Haas 1996) have all been analysed. Each of thesestudies, with their strengths and limitations, hasprovided some insight into the evolutionary his-tory of the family.

The placement of Trachypachidae relative toCarabidae and Hydradephaga has varied overtime and among authors. Some analyses have fa-voured a sister-group relationship to Dytiscoidea(e. g., Arndt 1993, 1998; Beutel 1993; Arndt &Beutel 1995; Beutel & Haas 2000) while othersplace them as sister to Carabidae, including rhy-sodines (e. g., Kavanaugh 1986; Beutel 1998).Given the conflicting results from analyses basedon both adult and larval characters, Shull et al.(2001) used 18S rDNA sequence data to attemptto resolve the relationships among the families ofAdephaga. In all but one alignment and analysisstrategy used by Shull et al. (2001) Trachypach-idae was associated with the remaining Geade-phaga and not Dytiscoidea. Ultimately, 18SrDNA sequence data, additional sequence datafrom other regions and characters from mor-phology should be combined in a single analysisto see if sufficient overall agreement can befound among character sets. Although analysesthus far do not unequivocally establish the rela-tionships among adephagan families, all rein-force the distinctness of the trachypachid lineage,its great age and relictual nature.

Rhysodidae is treated in this volume as a sepa-rate family from Carabidae, however, many au-thors have treated them as a subfamilial memberof Carabidae, potentially related to some groupof Clivinini (Bell 1998) (s. 1-7.9). Many authors


have noted that their extraordinary life historyinvolving wedging through wood and feeding onslime-molds has likely resulted in members ofthis group being highly apomorphic both asadults and larvae. The apparent deviation fromthe primitive form makes it difficult or impos-sible to clearly assess the homology of manyadult and especially larval structures. Rhyso-dines are generally separate from Carabidae be-cause of their distinctly divergent form, however,little evidence exist that Carabidae excludingRhysodidae is a monophyletic group. The ar-rangement of prehypopharyngeal setae in thelarva (Beutel 1993) and development of pubes-cence of the antennomeres in adults (Beutel1995) have been suggested as possible synapo-morphies for Carabidae not including rhyso-dines. These characters were presented to sup-port a working hypothesis that excludes rhyso-dines from carabids. Unfortunately, neither canbe considered very convincing given the varia-tion involved in these structures.

Various external structures found in adult rhy-sodines such as the form of the medial septumthat separates the procoxal cavities, the closureof the procoxal cavities and so-called ani-sochaete form of antennal cleaner, seem to indi-cate a placement within Carabidae (Bell 1967;Beutel & Haas 1996). This is consistent withcharacters of the female reproductive tract (Lieb-herr & Will 1998), though this character systemdid not provide any decisive synapomorphies fora specific sister-group relationship, the prefer-red phylogenetic hypothesis placed rhysodineswithin Carabidae in a grade with Clivinini. Theshared condition of a divided gonocoxite IX isthe only evidence that supports the inclusion ofrhysodines within anisochaetous carabid taxa.

Bell (1998) concluded that rhysodines, andwhat he considered a subtribe of scaritines, thesalcediines (Clivinini), were sister taxa. However,Scaritini is most likely not monophyletic, clivi-nines may only be distantly related to Scaritinisensu stricto, with some evidence supporting acloser relationship of rhysodines to Scaritini thanto Clivinini (see below). Molecular sequence evi-dence from the 18s gene (Maddison et al. 1998,1999; Shull et al. 2001) are problematic due tolikely convergence of several potentially aberrantsequences including rhysodines, scaritines, cicin-delines and paussines. Although the authors viewthese associations as potentially spurious, addi-tional taxon sampling and various methods overthe course of the several papers has not alteredthis result. None of these analyses have associ-ated rhysodines and clivinines nor has anyplaced rhysodines outside of Carabidae. Asnoted (Maddison et al. 1999) it is likely that thisparticular sequence cannot properly place basalcarabid taxa. It is also possible that some ofthese taxa are in fact related. A relationship be-


tween Rhysodini and Scaritini, exclusive of Clivi-nini, should not be discounted as these taxa alsoshare a peculiar structure, the vesicula seminalis(referred to as "mesidemia" by SmrZ (1981,1985)), not found in any other carabid tribe(SmrZ 1981, 1985; Will unpubl.).

The larval form is strikingly different in rhy-sodines, unlike any carabid taxon and salcediineshave apparently typical carabid larvae (Arndt,pers. obs.). Bell's (1998) proposed association ofrhysodines and salcediines (Clivinini), or any as-sociation with a particular tribe within Carab-idae, is not supported by larval characters (s.1-7.9).

Subdivisions within Carabidae have beenplaced at a variety of levels over the years (Ball1979; Ball et at. 1998). The subfamilies used byLawrence & Newton (1995), which have beenadopted for use in this volume, vary greatly inthe type and relative support for their mono-phyly. Several of the subfamilies contain rela-tively few species and only one or a smallnumber of genera, e. g., Gehringiinae, Omophron-inae, Hiletinae, Loricerinae, Elaphrinae and Sia-goninae. Each of these is well supported by apo-morphic characters and are distinct in form, buttheir relationships to other subfamilies are notwell understood. Other subfamilies have manymore taxa and vary considerably in regard to thesupport for their monophyly and evidence forsister-group relationships.

Although the monophyly of Cicindelinae iswell supported, the position of tiger beetles inCarabidae is still a question as contentious as therelationships for the families Trachypachidaeand Rhysodidae discussed above. Although nocharacter analysis has placed them outside ofCarabidae, they are still frequently classified thatway, with some authors maintaining them as aseparate family. However, this issue in particular,and the phylogenetic structure of Carabidae ingeneral, is dependent on the choice of the rootingpoint for the family. Recently it was proposed(Pearson & Vogler 2001) that Cicindelinae bemaintained as a family separate from Carabidaegiven results from studies by Bils (1976) andNichols (1985). However, neither Bils nor Nich-ols tested the monophyly of Carabidae excludingCicindelidae. Rather they chose Cicindelinae(-idae) as the outgroup and so fixed the rootingpoint between tiger beetles and other carabids.Therefore these analyses provide no evidence forthe exclusion of Cicindelinae. In fact, re-rootingthe phylogenies presented by Bils and Nicholsusing Trachypachidae, Gyrinidae or Haliplidae,as suggested by results of recent studies on ade-phagan families (Beutel 1998; Beutel & Haas2000; Shull et al. 2001), places Cicindelinae wellwithin the family Carabidae. Adult morphologi-cal structures consistently place tiger beetleswithin Carabidae and often associate them with

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some Carabinae (Carabini, Cychrini, Pamborini)(Beutel 1998; Liebherr & Will 1998) a relation-ship that suggests the unique form of the tigerbeetle female abdomen is derived from the car-abine type (Liebherr and Will 1998). Addition-ally, both Cicindelinae and some carabines(Calosoma) produce aromatic aldehydes as a ma-jor constituent of their defensive secretions(Moore & Wallbank 1968; Schildknecht et al.1968). However, pygidial gland secretory cell ag-gregation type and structure are more similar tothe typical hydradephagan form (Forsyth 1970).Larval characters suggest a close relationship ofCicindelinae and Loricerinae (Arndt 1993).Combined analysis of adult and larval charactersplaces them as a member of a more derivedgrade than carabines together with loricerines,elaphrines and scaritines (Beutel & Haas 1996).

Published analyses of 18S ribosomal DNA(Maddison et al. 1999; Shull et al. 200 I) areproblematic with regard to the placement ofcicindelines within carabids, and their study sug-gests that the 18S rDNA sequence is probablynot sufficient for resolving relationships of thesubfamily. Results of these analyses of sequencedata are consistent with a placement of cicinde-lines within Carabidae, but are not consideredconclusive as the sequence divergence of Cicinde-lini is likely to strongly effect results due to con-vergence.

Paussinae is a very well supported monophy-letic group based on morphological characteris-tics including aspects of the pygidial glands andsubstantial larval characters (Beutel 1995). Theyshare with Brachininae similar chemical com-pounds and delivery of these defensive secretionsby crepitating. However, the form of the glands,details of the secretory cells and mode of deliverydiffers dramatically between these groups (Eisneret al. 2000, 2001). Paussinae is generally consid-ered adelphotaxon to the remaining carabid sub-families or one of the basal-most lineages, how-ever, evidence for this is limited. This basal posi-tion is supported by female reproductive trackand larval characters (Arndt 1993; Liebherr &Will 1998). However, DNA sequence data (Shullet at. 2001) consistently places Paussinae as amuch more derived group and never as sister tothe remaining Carabidae. Shull et al. (2001) sug-gest the intriguing, and only slightly less parsi-monious, possibility that a clade of Paussinae +Brachininae lies at or near the base of Harpal-inae. This would, as pointed out by Shull et al.,require the reversal of a large number morpho-logical characters thought to define the generalpattern of relationships in the family.

Gehringiinae, like nebriines and carabines,have what is thought to be the plesiomorphicthoracic configuration for Carabidae: confluent,internally unbridged and open procoxae, meso-ventrite with anterolateral grooves for reception

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of the procoxae and with a median hexagonalgroove). As this is the plesiomorphic arrange-ment these structures do not provide groupinginformation and emphasis for classification hasbeen placed on whether the arrangement of thepro tibial spurs and antennal cleaning organ rep-resents an isochaete or anisochaete condition(Bell 1967; Beutel 1992; Beutel & Haas 1996).The condition of the gehringiine tibia is notclearly assigned. If considered as isochaetousGehringiinae is placed as sister to Paussinae(Beutel & Haas 1996), if anisochaetous, theycould be sister to Nebriitae. Placement withinanisochaete taxa and near nebriines is consistentwith the dimerous condition of the gonocoxiteand dual spermatheca in the female tract (Lieb-herr & Will 1998). Larval characters have notproven decisive for grouping Gehringiinae andno results have been published based on DNAsequence data.

The rounded body shape of species in Omo-phroninae immediately sets them apart from allother adephagans. Details of their adult mor-phology are equally unique and this has resultedin a wide range of proposed relationships for thesubfamily. Some authors have even placed themoutside Carabidae with Hydradephaga (Bils1976; Nichols 1985), but like Cicindelinae dis-cussed above, re-rooting these phylogenetichypotheses maintains Omophroninae as a mem-ber of Carabidae. Larval characters clearly sup-port their inclusion within carabids (Beutel 1991;Arndt 1993) and suggest a position near the baseof Anisochaeta or with Nebriitae. A close rela-tionship between Nebriitae, Omophroninae andGehringiinae was suggested by Liebherr and Will(1998) based on a hypothesised substitution ofspermathecal function in these taxa. Analyses of18S rDNA sequence data clearly place Omo-phroninae in Carabidae and in the basal gradeof non-Harpalinae taxa (Maddison et al. 1998,1999; Shull et al. 2001). However, no consistentsister-group relationships were found in the vari-ous analyses.

Carabinae as delimited by Lawrence & New-ton (1995) is almost certainly not monophyletic.As discussed above Cicindelinae

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~/.A' - o,,~kiplingw/FULLPAPERS/24.pdf · 2008. 9. 5. · We thank Dr. Vasily V. Grebennikov...