Carabidae Latreille, 1802 119

~

~/.A' .~..A ---:: o,,~ ~~~ ~ ~~- ~~ I ~ A B

Fig. 7.7.4. Larval head structures. A, nasale and adnasalia, lateritium, S. breve, 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- Hlavac, T. F. (1975): The prothorax of Coleoptera (ex- pendage on lateral side of antennomere III ab- cept - ). - Bulletin sent, replaced by ventral sensorial field, apical of the Museum of Comparative Zoology 147 (4): 137-183. part of maxillary palpomere 3 with additional se- tae, number of nasal teeth increased (6-8), uro- Lindroth, C. H. (1960): The of Trachypachus Mtsch., Gehringia Darl., and Opisthius Kirby (Col. gomphi fixed, horn-shaped (groundplan), eight Carabidae). - OpusculaEntomologica25: 30-42. long setae on tergite IX (including those on uro- (1961- 69): The ground (Carabidae, excl. gomphi). The specific shape of the parameres Cicindelinae) of Canada and Alaska. Parts 1-6. - (Lindroth 1961-69; Beutel 1994) is an autapo- Opuscula Entomologica XlVIII + 1192 pp. 1961, morphy of adults. The absence of the katas- Part 2, Suppl. 20: 1- 20; 1963, Part 3, Suppl. 24: tigma, the specific sculpture of the elytra, the 201-408; 1966, Part 4, Suppl. 29: 409-648; 1968, kidney-shaped sensorial field of the larval anten- Part 5, Suppl. 33: 649-944; 1969 Part 6, Suppl. 34: nomere 3 and the large, ventral sensorial field 945-1192; 1969 Part I, Suppl. 35 I + XlVIII. of antennomere 4 are larval autapomorphies of Ponomarenko, A. G. (1977): Suborder , etc. Systolosoma. The dilated larval distal maxillary Pp. 3-104 In Arnoldy, L. v., Zherikhin, V. V., Nikritin, L. M. & Ponomarenko, A. G. (eds.) Meso- palpomere with two separated sensorial fields is zoic Coleoptera [in Russian]. - Trudy Paleontolo- an autapomorphy of Trachypachus. gicheskogo Instituta Akademiya Nauk SSSR 161: 1-183. Acknowledgements We thank Dr. Vasily V. Grebennikov (Ottawa) for thoroughly reviewing this chapter. 7.8. Carabidae Latreille, 1802 Literature Erik Arndt, Rolf G. Beutel & Kipling Will Arndt, E. & Beutel, R. G. (1995): Larval morphology of Systolosoma Solier and Trachypachus Motschul- sky (Coleoptera: ) with phyloge- In most recent classifications (-ni), netic consideration. - Entomologica Scandinavica Cicindelinae and Paussinae are included. How- 26: 439-446. Beutel, R. G. (1993): Phylogenetic analysis of Ade- ever, Rhysodidae were excluded by Kryzhanov- phaga (Coleoptera) based on characters of the lar- skij et al. (1995) and Lawrence & Newton (1995). val head. - Systematic Entomology 18: 127-147. They are tentatively treated as a separate family (1994): Study on the systematic position of Systolo- here (s. 1-7.9). soma breve Solier (Adephaga: Trachypachidae) based on characters of the thorax. - Studies on Distribution. World-wide (except Antarctica), Neotropical Fauna and Environment 29 (3): 161- more than 40000 spp. and 1500 genera. Distribu- 167. tion of subgroups see below. Bils, W. (1976): Das Abdomenende weiblicher, ter- restrisch lebender Adephaga (Coleoptera) und seine Bedeutung fur die Phylogenie. - Zoomorphologie Biology and Ecology. Carabidae live in all ter- 84: 113-193. restrial habitat types from the subarctic to the Crowson, R. A. (1955): The Natural Classification of wet tropical regions. The majority of ex- the Families of Coleoptera. 187 pp. Nathaniel Lloyd cept those of tropical rain forests and subtropical and Co., LTD., London. montane forests (e. g., Mexico, Hawaii) is ground 120 Erik Arndt, Rolf G. Beutel & Kipling Will dwelling. Adults and larvae live in soil, in leaf vanaugh 1985; Baehr 1997). Brood cart:}occurs litter or are active on the ground surface, some- in some species of Pterostichini, were the females times climbing bushes or plants. Many species, guard their eggs (Kavanaugh 1998) and in some mainly in Trechini, Anillini and , in- species of Harpalini, where the female lay eggs habit caves or deep soil microcaves (mss). Even in individual soil cells prepared with food stores. though many species occur in moist habitats, an Pupation usually takes place in moist soil. The amphibious way of live is reported in only very full-grown last instar larvae dig a pupal chamber few cases (e. g., Carabu variolosus at small using their head and legs. In one species streams in Fagus forests, C. clathratus and some (Thermophilus sexmaculatum) the first instar other species in swamps; Sturani 1962; larva digs the pupal chamber, and the final sec- Cicindelinae in inundation forests; Adis & Mess- ond instar remains inactive in this chamber ner 1997; Raw/insius in shallow regions of rapid (Arndt & Paarmann 1999). The life span in the mountain streams; Davidson & Ball 1998). A field is up to four years; at least carabids of the large percentage of species in tropical rain forests temperate regions live usually longer than one and subtropical montane forests, and few species year. in temperate regions are arboreal and resting un- Adults and larvae have a partial extraoral di- gestion. During the manipulation of the prey, der leaves or bark. The majority of carabid spe- mandibles and maxillae rotate the food item cies is able to fly and has a great dispersal power. while digestive fluids are expelled onto it. Adults Some species are dimorphic regarding their wing of most species are omniv-orous (Larochelle and muscle development; usually the minority of 1990;Thiele 1977), even though carnivorous nut- individuals is wingless (e. g., most Carabus, An- rition seems to prevail. A few groups are speciali- thiini). Even species exclusively moving on the sed herbivores, e. g., Zabrus and some represen- ground have an enormous dispersal power; tatives of Harpalini; even arboreal species of migrations of 77 m per night and running speeds were found to feed on flowers and nectar of 0.16 mls were estimated in Carabus species (Arndt et al. 2001). Much less is known on the (Thiele 1977). Carabid beetles show a circadian feeding preferences of larvae, but carnivorous rhythm and outside the tropical regions also an habits of the majority of species is likely. Larvae annual rhythm. The majority of species is noc- of several taxa are specialised on specific prey turnal; some species change their activity de- such as snails (, ), springtails pendent from climate or season but some taxa (Nebriini, Notiophilini, Loricerini), ants, or ant are active during day light (e. g., ). brood (Anthiini, Graphipterini, Metriini, Ozae- The annual rhythm controls the reproductive nini). Larvae of Paussini and Pseudomorphini period; different mechanisms of diapause are de- are probably fed by ants and those of scribed for species of the temperate and sub- and Brachinini are ectoparasitic on eggs tropical regions (Paarmann 1979). During in- and pupae as far as known. Larvae of Cicindef- active periods in winter quarters or hiding places inae and Ozaenini live in burrows and lie in am- during day time, aggregations of several hundred brush for prey at the entrances. Larvae of several specimens are possible. Harpalini are specialised on seeds (Brandmayr Carabidae are bisexual. Females lay eggs (as 1975; Zetto Brandmayr 1990; Arndt et at. 1996). far as known) separately or in small groups in Only very few ground beetles are known as pest small hollows in substrate or under bark or in . The grain ground (Zabrus tenebri- cases made of mud (King 1919; Thiele 1977) or oides) is the only species of a certain economic algae and bark (Will 1998). The number of eggs importance, but it became known as serious pest per female varies between four and several hun- only in a restricted period and restricted area in eastern Germany, Moravia and the Ukraine. dreds depending on the species (the highest num- The ecological importance of the extremely di- bers recorded are 653 for sycophanta; verse family Carabidae is not well understood. Thiele 1977 and 660 for Colpodes buchanani; The ground beetles represent a major part of the Paarmann & Bolte 1990). The egg development invertebrate predator guild of the soil fauna, lasts few days to several weeks. Most of the spe- e. g., in temperate forests and agrocoenoses. Due cies have three larval instars. Exceptions are two to their abundance and ubiquitous occurrence, larval instars in some representatives of , the importance of the role of carabids in these Harpalini, Lebiini and Anthiini (Bily 1975; ecosystems can be safely assumed. Capogreco 1989; Arndt & Paarmann 1999), Insectivores, , rodents, birds, amphibians, which is regarded as an adaptation to an arid reptiles, ants, Asilidae, and Aranaea are known environment, and four or five larval instars as predators of ground beetles. Sporozoa (Gre- respectively in members of Lebiini and Brachi- garinida), Nematoda (Mermis, Hexamermis), nini (Erwin 1967, 1975), which is regarded as an Nematomorpha (Gordius) are endoparasites, and adaptation to an ectoparasitic way of live. Pseu- several taxa of mites (Trombidiiformes, Sarcopti- domorphini are ovoviviparous (Liebherr & Ka- formes) are known as ectoparasites on carabids. Carabidae Latreille, 1802 121

B

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

'-- Several other representatives of Acari (e. g., Par- and protruding, but different degrees of reduc- asitus) occur frequently on Carabidae but they tion and total loss occur. Clypeus trapezoid, with are phoretic. Parasitoids of carabid larvae in- two pairs of fixed setae. Supraocular area with clude the hymenopteran taxa Proctotrupes spp one or two pairs of fixed setae and subocular (Proctotrupidae; on and ), genal area with one pair of setae. Gula fairly Microtonus spp (Braconidae; on Harpalinae), elongate, moderately broad, distinctly delimited Methocha spp (Tiphiidae; on Cicindelinae) and by gular sutures, always glabrous, without fixed representatives of the dipteran families Larvae- setae. Median gular apodeme present or absent. voridae (on several subfamilies) and Bombyli- Tentorium with all parts well developed, usually idae (Anthrax spp; on Cicindelinae). Besides with median laminatentorium. Labrum variable non-specific insect parasitizing fungi (Ento- in shape, transverse, medially emarginate, bi- mophtorales, Hypomycetes) 16 genera with sev- lobed or rarely trilobed. Anterior margin with six eral hundreds of species of Laboulbeniales (As- or eight fixed setae. Anterolateral margin usually comycota) infest carabids. with shorter curved setae. Ventral side largely The following references summarise informa- unsclerotized and covered with sensorial struc- tion on the biology of "these beetles: Den Boer tures, with shallow excavations adapted to the (1977), Thiele (1977), Larochelle (1990), Lin- convex dorsal surface of the mandibles. Anten- droth (1992), Turin (2000). nae almost always filiform and II-segmented, usually inserted below lateral frontal projection, Morphology, Adults (Figs. 7.8.1-7.8.3). 1- rarely inserted on dorsal side of head capsule 85 mm long. Usually flattened and elongate, (Cicindelinae). Pubescence present on antenno- meres 4-11 or 3-11, sometimes also on basal with distinct pronoto-elytral angle, rarely with segments but then less dense than on distal an- distinctly convex dorsal side and laterally evenly tennomeres. Mandibles with basic form of a tri- rounded (e. g., Omophron). Cuticle usually angular pyramid, with dorsal, ventral and exter- smooth and shiny. Colour black or dark in most nal surface, moderately to strongly (e. g., Leca- species, sometimes metallic. Depigmentation of nomerus; Acorn & Ball 1991: Fig. 16) elongate. parts of the integument can result in conspicuous Primarily with distinct apical tooth, terebral colour patterns (e. g., Cicindela, Eurynebria, tooth, anterior retinacular tooth, posterior reti- Omophroninae, ). Microsculpture usually nacular tooth, premolar or molar tooth, incisor present as a fine polygonal meshwork. Strongly ridge, terebral ridge, retinacular ridge, ventral impressed microsculpture decreases brilliance of groove with row of microtrichiae and basal cuticular surface. Long articulated setae (= brush (Acorn & Ball 1991). Maxillae composed 'macrochetes' sensu Jeannel 1941-42; fixed se- of transverse, short cardo, which articulates with tae) distributed with regular patterns (chaeto- well developed fossa maxillaris, triangular ba- taxy). Small articulated hairs with irregular dis- sistipes, mediostipes, elongate triangular palpifer tribution present or absent, sometimes forming with several fixed setae, lacinia, galea and 4-seg- pubescent surfaces. mented palp. Lacinia basally fused with medio- Head (Figs. 7.8.2A, 7.8.3.A) prognathous, re- stipes, hook-shaped, with dense row of mesally latively elongate, moderately retracted. Without directed strong setae and thin hairs. Galea 2-seg- distinct neck region. Frontal furrows present or mented, palp-like. Palpomere 1 very short. absent. Compound eyes usually well developed Palpomeres 3 and 4 vary strongly in size, usually 122 Erik Arndt, Rolf G. Beutel & Kipling Will

7/

" f ~ , ,~ ..'..\

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. lateral setae. Parallel-sided palpigers inserted be- Submental part of labium T-shaped, fused with tween emargination of mentum and prementum. gula posteriorly. Transverse anterior part mesally Palp, 3-segmented, with very short palpomere 1. delimits fossa maxillaris, with two or more pairs Preoral cavity usually with well developed filter of fixed setae. Mentum with distinct lateral lobes apparatus composed of rows of hairs on mandi- and often with a more or less projecting median bles, maxillae, and hypopharynx. bifid or simple process, with one pair of fixed Prothorax (Fig. 7.8.3 C) rounded laterally and setae. Prementum variable in shape but usually more or less strongly narrowed posteriorly, dis- roughly quadrangular, with sclerotized median tinctly narrower than elytra at posterior margin part and more or less distinct, less strongly scle- (with few exceptions, e. g., Omophron, Pseudo- rotized lateral lobes (paraglossae). Anterior mar- morphinae). Pronotum medially divided by lon- gin with one pair of long, fixed setae close to gitudinalline, usually with raised lateral margin median line and sometimes additional shorter (indistinct or absent in 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 margin in cross section, with smooth collar region and with two or more pairs of fixed setae, one of articulating with prothorax in a ball-and-socket them located close to posterior margin. Proster- manner (Figs. 7.8.2 F, 7.8.3 C; Paussinae [excl. nal process either narrow and short (Gehringia), ], Cicindelinae, , , strongly developed, projecting beyond the hind Migadopinae, , Trechinae, Harpalinae, margin of the procoxae and tapering posteriorly Brachininae, ) (= harpaline (Metrius, Carabinae, Hiletinae), or shortened type). Mesocoxal cavities laterally open, i. e. bor- and posteriorly truncate (Paussinae excl. Met- dered by mesepimeron ('disjunct type') or closed rius, Cicindelinae, Omophroninae, Elaphrinae, ('conjunct type') (Trechinae, Harpalinae, Bra- Loricerinae etc.). Procoxal cavities open, without chininae, Pseudomorphinae). Elytra usually cov- internal postcoxal bridge (Gehringia, Carabinae, ering abdominal tergites completely, apically Hiletinae) or externally closed. Protibiae usually truncate in some groups (Paussinae excl. Met- with two apical spurs, both inserted apically rius, Gehringiinae, Lebiini, Odacanthini, Galeri- (Paussinae [= "Isochaeta"], Cicindelinae, Opis- tini, Brachininae). Basal margin present or ab- thiini, ) or one spur shifted proximally sent. Disc primarily with eight striae and nine ("Anisochaeta"). Antennal cleaning (Figs. interspaces. Setae generally present in inter- 7.8.2 B, 7.8.3 B) organ restricted to apical part of spaces 3, 5 (discal setae), and 9, but sometimes tibia (Cicindelinae, Opisthiini, Carabini) or ex- also in 1 and 7. Sutural stria usually not recur- tended towards base (Pausiinae, Gehringia, rent at apex. Epipleura broad at humeral region, Omophroninae, Hiletinae etc.). Scutellar shield gradually narrowing posteriorly, usually ending enclosed by elytral bases or shifted anteriorly at apical external angle of elytra, not reaching (e.g., Broscus). Mesoventrite of Gehrinigiini sutural angle. Always without setae but some- (Fig. 7.8.2 C), Carabinae and Hiletinae short, times with pubescence. Metanotum usually of with hexagonal groove and anterolateral grooves generalised adephagan type but shortened and for reception of pro coxae (= carabine type). strongly simplified in forms with completely re- Mesoventrite of other groups (Figs. 7.8.2 F, duced flight organs (e. g., Omus). Anepisternum 7.8.3C) moderately elongated, without hexago- does not reach mesocoxal cavity. Epimeron ex- nal groove and anterolateral grooves, rounded posed, narrow and parallel-sided (Gehringiinae, 124 Erik Arndt, Rolf G. Beutel & Kipling Will

A B c D E Fig. 7.8.4. Habitus of larvae of different subfamilies. A, Carabusexaratus (Carabinae) (redrawn from Arndt & Makarov 2003); B, Platychile pallida (Cicindelinae) (Arndt 1998b); C, Lindrothius sp. (Harpalinae, Platynini) (Arndt & Hurka 1992);D, quadrillum(Harpalinae, Lebiini) (Arndt 1989);E, Trichognathusmarginipennis(Harpalinae, Galeri- tini) (Arndt & Drechsel 1998).

Omophroninae, Elaphrinae, Loricerinae, Miga- unsegmented, but divided into proximal and dis- dopinae, Scaritini), concealed (Opisthiini, Carab- tal part in most representatives. inae), or lobate (Hiletinae, Trechinae, Harpal- inae). Metaventrite (7.8.20, F, 7.8.3 D) with Morphology, Larvae (Figs. 7.8.4-7.8.10). cam- discrimen and complete transverse suture sepa- podeiform, moderately flattened, subparallel. rating preepisternum from katepisternum. Meta- Sclerotized parts brownish to black. Head prog- coxae (7.8.20, F, 7.8.3 D) not extended crani- nathous, rounded laterally (Paussinae, Cicindel- ally, medially not fused and not fused to kate- inae) or roughly quadrangular (Fig. 7.8.5 A); cer- pisternum, mobility partly retained (c. 5°). Later- vical groove present laterally in most taxa but ally scarcely broader or as broad as posterior absent in basal groups (e. g., Paussinae, Carab- margin of ventrite, not reaching elytral epipleura inae, Cicindelinae, Omophroninae). Head later- laterally. Metafurca well developed. Mm. furca ally with six stemmata arranged in two rows. Oc- coxalis anterior and posterior present. Alae ular groove present posterior to the stemmata in sometimes reduced, if well developed with ob- many taxa but absent in basal groups (e. g., longum but without katastigma (subcubital setal Paussinae, Carabinae). Frontal suture almost al- binding patch). ways sinuate; posterior part of frontale (pars ab- oralis frontalis) usually with pairwise egg burst- Abdomen usually with six visible sternites. ers; coronal suture usually present, often Sternite II only visible laterally in most groups. strongly shortened or absent. Anterior clypeola- Large median piece of sternite II usually present bral margin primarily with four nasal teeth in species with distinctly separate metacoxae (Fig. 7.8.5 A), each with one small ventrally di- (Metrius, Ozaenini part.). Sternites III + IV rected microseta; ventral side of clypeolabral re- fused. Terminal sternite VII posteriorly acumi- gion with row of teeth. Antennae 4-segmented, nate, rounded, or truncate with rounded lateral anteriorly directed. Antennomeres I and 2 sub- edges. Tergites I-VIII with spiracles. Posterior cylindrical, antennomere 3 with bulb-like senso- segments invaginated and strongly modified. rial appendage laterally, antennomere 4 smaller, Aedeagus usually with asymmetric parameres. rounded apically. Mandibles with retinaculum; Coxosternum VIII only exposed in Brachininae. terebrum usually with two cutting edges; penicil- Gonocoxae relatively short, probably primarily lus present or absent. Maxillae articulated with Carabidae Latreille, 1802 125

B E A tergite ~

:-~ %~le' ::cIJ, spiracle I J _ C~--< iD /

:I ~ -:-, eplpleumu - ,I ht ~G->fs V I

: \.L' :) ;- hypopleuron ~ \ \" /~, I ~.~ I --:-- laterosternite ~ ~ ~ ~, , -' , (=coxallobe) I ___ ~ I , l;lJ~/ ~ ~ _ I : I r' ! \ '- ( . , I D ~ /' \ ~ ~1\ inner sternite mediosternite inner sternite mediosternite Carabus sp. Cicindela sp. Fig. 7.S.SA, Lindrothius horsti, head capsule with mandible and antenna, first instar, dorsal aspect (Arndt & Hurka 1992);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, Carabusnemoralis (left) and Cicindelacampestris (right) (Arndt 1997);E, Agonum miilleri,pygopod, lateral aspect (Arndt 1993). anteroventral margin of head capsule, moveable sent. Hypopharynx with dense field of micro- in all directions; maxillary groove completely re- trichiae, separated from dorsal surface and bulg- duced; cardo small, with or without separate me- ing or completely flattened. M. tentoriohypo- sal sclerite; stipes usually elongate; mobility be- pharyngalis primarily present, but absent in lar- tween both parts strongly restricted; lacinia usu- vae with flattened hypopharynx (e. g., Ptero- ally short (elongate and hook-shaped in Metrius, stichus, ). Gula usually elongate, repre- Ozaenini and Omophron), absent in some groups; sented by a narrow area enclosed by paired gular galea 2-segmented; palpi 4-segmented (or 3-seg- sutures, rarely broad (e. g., Loricerinae; Licinus). mented and inserted on palpifer, Fig. 7.8.5 B). Functional mouth open but narrow, in close con- Maxillae moved by four longitudinally arranged tact with anterior hypopharyngeal margin. Preo- extrinsic muscles; craniostipital muscle (levator ral filter apparatus formed by hypopharyngeal of maxilla) probably homologous with M. trichiae and trichiae on the mandibular (penicil- craniolacinialis. Submentum completely fused lus) and maxillary bases. Prepharyngeal tube al- with remaining parts of head capsule; mentum ways present. M. clypeopalatalis always repre- short and membranous; prementum usually with sented by several bundles. Pharynx fairly narrow, ligula and 2-segmented palpi (Fig. 7.8.5 C). Pre- primarily with well developed ventral dilator (M. mentum retracted by two pairs of muscles with tentoriopharyngalis) and postcerebral dorsal di- tentorial origin. M. submentopraementalis ab- lator (M. verticopharyngalis), but both muscles 126 Erik Arndt, Rolf G. Beutel & Kipling Will

DEF G

Fig. 7.8.6A-C, Brachinuscrepitans,first instar, dorsal aspect; A, antenna; B, maxilla; C, tergite IX with urogomphi; D, sp., urogomphi, third instar; E, 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- one strong seta on outer margin. Cardo with one inae, Brachininae). distinctive seta; stipes with two large setae on Thorax with tergites medially divided by nar- outer side and a field of setae on inner side; laci- row median ecdysial suture; mesonotum and nia with one seta subapically (even distinct on metanotum with anterior keel; pronotum larger strongly reduced lacinia); palpifer with a distinc- than following segments. Large spiracle present tive seta on the ventral side; setae on galea and between pro- and mesothorax. Legs 5-segmented palpi very small. Prementum including ligula pri- (coxa, trochanter, femur, tibia, tarsus), usually marily with seven pairs of setae, but number al- with two claws. Four small sclerites present later- most always reduced or increased; ventral side ally and dorsally of coxal base (episternum, epi- with a pair of very small basal setae, a large meron, trochantin, pleurite, the latter two are central seta, and a pit on the apical part of the lacking on the prothorax). Abdominal tergites ligula (distinctive in most species). Urogomphi I - VIII similar in structure, with anterior keel with five long setae in first instar and nine long and median ecdysial suture; ventral and lateral setae in the following instars. sides with sternites, epipleuron, hypopleuron and Larvae of several groups of Carabidae are a lateral spiracle; sternites consisting of one large strongly modified morphologically (e. g., legs re- (medio-) sternite and smaller paired anterior, duced, hyperprognathous, physogastric; see inner and outer (Iatero-) sternites (Fig. 7.8.5 D). below). Abdominal segment IX smaller, with fused ster- The following references summa rise informa- nites; tergite IX usually with a pair of urogomphi tion on the larval morphology of Carabidae: van (Fig. 7.8.6C-E, H, I); urogomphi long and slen- Emden (1942), Bousquet & Goulet (1984), Arndt der, moveably attached to tergite and unseg- (1991 a, 1993), Beutel (1993), Luff (1993). mented, or segmented and/or fused with tergite, or reduced. Tergite X (pygopod) cylindrical, di- Paussinae (= Metriinae + Ozaeninae + Pro- rected downwards (Fig. 7.8.5 E). topaussinae + Paussinae) First instar larvae with characteristic chaeto- taxy (Bousquet & Goulet 1984; Arndt 1993). Distribution. All zoogeographic regions, but Epicranial sclerites with ten dorsolateral and most diverse in the tropics. Only few species oc- seven ventral setae; frontale with eleven setae, cur in southern parts of the Northern Hemi- two on anterolateral margin and one very dis- sphere. The Metriini are restricted with one ge- tinctive long seta in anterolateral angle. Anten- nus and two species to western North America. nomere III with three long setae, antennomere Ozaenini occur with about 14 genera pantropical IV with four long apical setae. Mandible with and in southern North America, the Protopaus- Carabidae Latreille, 1802 127 sini are restricted with one and seven spe- pa]pi very variable. Mentum usually reduced in cies to the eastern Oriental region. The Paussini size and retracted or obliterated medially. Pro- are the most diverse and widespread tribe. thorax usually strongly modified. Prothoracic glands almost always present, usually associated Bio]ogyand Eco]ogy. Higher paussines (= Proto- with concavities or clefts and trichome systems. paussinae + Paussinae) are characterized by their The trichome system is specialised in Protopaus- advanced myrmecophi]ous habits, at least in the sus (Nage] 1997) and absent in Arthropterus and larval stage. This unusual life style may have some other genera (e. g., Homopterus, Carabido- originated from specialised predatory habits of memnus). Prosternal process fairly broad and larvae, which are a ground plan feature of the posteriorly truncate in Protopaussus, distinctly or subfamily (Nagel 1979). Larvae of Metriini (Met- completely reduced in Paussini. Procoxae usually rius) and Ozaenini produce gland secretions at very prominent and contiguous or almost contig- their highly modified terminal abdominal seg- uous (Paussitae sensu Nage] 1997). Femora and ments, which attract other insects. These secre- tibiae of fore-, midd]e- and hind legs usually tions are apparently also attractive for ants, and broadened and flattened (not in Pleuropterus). may have enabled paussine larvae to enter ants Protibial antenna cleaning organ vestigial or ab- nests as parasites. Adults of Paussini are adapted sent. Protarsomeres usually cylindrical, in some to these hostile environments in different ways cases partly or completely retractile (Ho- (defiant type, symphilous type). Myrmecophilous mopterus). Elytra without lateral edge. Meta- habits have independently evolved in Physea ventrite usually without discrimen and transverse (Ozaenini). suture. Median cell of alae usually triangular (not in Protopaussus). Abdominal sternites 11- Morphology, Adults (Darlington 1950; Nagel IV fused, sutures between sternites faintly indi- ]997). Colour black (e. g., Metrius) or brown cated or absent. Stridulatory file on the ventral (most Ozaenini, Protopaussini and Paussini). abdominal base present or absent. Terminal ab- Body laterally rounded (Metrius, Mystropomus) dominal segments partly exposed and sclero- or parallel-sided. Fixed setae (i. e., long mecha- tised. Parameres slender and asymmetric in ad- noreceptive setae) present or absent in Ozaenini, vanced paussines. always absent in Protopaussini and Paussini. An- tennae filiform or moniliform in Metrius, Ozae- Morpho]ogy, Larvae (Fig. 7.8.7 A -G). Larvae nini, Protopaussus and Megalopaussus. Strongly live in burrows in soil or in ants nests and are modified in other subgroups of Paussini, with strongly modified morphologically. Head and pedicellus very small, deeply sunken into apex of posterior part of body bent upwards (dorsad) scapus, connected with antennomere 3 by rigid wedging, and flagellomeres broadened and often (Fig. 7.8.7. G). Head capsule s]ightly or distinctly rounded ]aterally; stemmata reduced to one transformed into a compact club. Procoxal pro- cess elongate and tapering posteriorly in Metrius, ocellus or lacking; coronal suture absent (Fig. more or less truncate and posterolaterally con- 7.8.7 A); posterior tentorial grooves represented nected with pleural process or strongly reduced by a single narrow cleft immediately adjacent to in other paussines. Procoxal cavities always a deep posteromedian emargination of the closed. Both pro tibial spurs in apical position if ventral wall of the head capsule in Paussini present, usually obsolete or absent in Paussini. (Arndt & Beutel 1994); nasale present in Metrius Antenna cleaning organ extended towards base but absent in other groups; fronta]e with mem- of tibia or reduced (Paussini). Hind walls of pro- branous anterior margin in Paussini (Arndt & coxae contact anterior excavations of meso- Beutel ]994); mandible (Fig. 7.8.7 B) broad at ventrite in Metrius. Connection between pro- base, with s]ightly sclerotized appendage (Iacinia and mesoventrite of harpaline type in all other mobilis) proximal to retinaculum in Paussini; paussines, with smooth anterior collar of meso- maxillae with 4-segmented palpi in Metriini but ventrite and without hexagonal groove. Meso- 3-segmented palpi in Ozaenini and Paussini; coxae separate or contiguous. Metacoxae sepa- palpomere I more or less comp]etely fused with rate in Metrius and Ozaenini (almost contiguous stipes in the latter group; galea I-segmented in Physea), contiguous in Protopaussini and (Arndt & Beute] 1994), blade-like in Metriini Paussini. Termina] abdominal segment covered (Beutel 1992); lacinia absent in Paussini but long by elytra (Metrius, most Ozaenini, Protopaussus), and blade-like in Metriini. Prementum elongate, exposed in Dhanya and most Paussini. Speciali- with tubercles. Paussini with posterior tentorial sed abdominal defense secretion delivery system arms originating from narrow, common median present. Subapical elytral flange present in stalk, immediately adjacent with posteromedian Paussinae excl. Metrius. emargination of head capsule, with dorsa] arms Higher paussines with labrum often extended strongly flattened and exceptionally short laterally or longitudinally. Mandibles compara- (Arndt & Beutel 1994). Regular fringe of longer tive]y small and simple. Maxillary and labial hairs absent from cranial part of hypopharynx in 128 Erik Arndt, Rolf G. Beutel & Kipling Will

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

Biology and Ecology. G. olympiaca occurs on gravel banks of small to moderate mountain brooks with cold water. It lives in the moist gravel and coarse sand and avoids coming to the surface. Oviposition takes place in early summer.

Morphology, Adults (Fig.7.8.2A-D; Lindroth 1969; Beutel 1992). 1.6-1.7 mm long, black, up- per surface covered with scattered setigerous punctures. Compound eyes with large, convex facets. Antennae short, moniliform, pubescent except 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 organ present, elongated towards proximal part of tibia. Spurs both in terminal position. Protarsi not dilated in males, proximal three tarsomeres with sparse adhesive setae. Mesoventrite of cara- bine type. Mesocoxal cavity disjunct. Elytra with underlapping edges (Lindroth 1961- 69), with deep sutural (and often faintly suggested 2nd and 3rd) stria, otherwise with sparse punctures arran- ged in irregular rows. Apex truncate, exposing last tergite. Metaventrite broad, with short discrimen and laterally obsolete transverse su- Fig. 7.8.7A, Paussus sp., head capsule, dorsal aspect ture. Metepimeron parallel-sided and exposed, (Arndt & Beutel 1995); B, Paussus sp., mandible, dorsal not lobate. Metacoxae distinctly separate. Alae aspect (Arndt & Beutel 1995); C- F, Transformation series well developed, with marginal fringe of hairs. of posterior part of abdomen in representatives of Pauss- inae. C, (redrawn from Bousquet Abdomen without separate sclerite between 1986); D, Pachyteles sp. (redrawn from Costa et al. 1988); metacoxae. Terminal sternite VII with almost E, Physea setosa (redrawn from Bousquet 1986); F, Paus- truncate, very slightly convex hind margin. Para- sus sp. (Arndt 1998 a); G, Metrius contractus, habitus, first meres slightly asymmetric. instar (Bousquet 1986). Morphology, Larvae. Length of specimen de- scribed by Lindroth (1960) 1.2 mm. Habitus of Paussini (specialised feeding habits); present but larvae and characters generally very similar to sparse in Metriini. those of Trechinae. Body slightly sclerotized, ab- Legs primarily 6-segmented with two claws dominal tergites and sternites not apparent. (anterior claw longer in Metriini and Ozaenini); Head almost square, parallel-sided; sutures in- tibia and tarsus fused to one strongly sclerotized distinct. Frontale distinctly extended posteriorly; segment with numerous strong setae on the coronal suture short; postocular groove absent; ventral side and a single small hook-like claw cervical groove absent. Egg-bursters consist of in Paussini. Posterior abdominal segments and two rows of 5-7 minute tubercles on posterior urogomphi strongly modified. Epipleurites IX frontale. Stemmata absent. Nasale with slightly Carabidae Latreille, 1802 129 bisinuate anterior margin, slightly produced in dian line, forming part of a prey-grasping basket. advance of inconspicuous adnasale, with four Mandible with single cutting edge and bidentate small teeth alternating with minute setae. An- retinaculum; penicillus absent (Fig. 7.8.6 F). Pre- tenna with three stout basal segments and slen- epipharynx with V-shaped, sclerotized bar; hypo- der antennomere IV; antennomere III with mesal pharyngeal bulge reduced. M. tentoriohypopha- spine and stalked, globulous sensorial append- ryngalis medialis absent. Maxilla with very long age. Mandible sickle-shaped, with one lateral lacinia and almost equally long 2-segmented ga- seta and well developed pointed retinaculum; lea. Prementum small; ligula long, moderately penicillum absent; mesal edge of stipes with only wide at base and extended as digitiform pro- one seta; 2-segmented galea with very slender jection apically (Fig. 7.8.6 G). Ligula and labial distal segment; lacinia absent; basal palpomere palps subequal in length. Posterior tentorial with twisted seta. Prementum subquadrate; palpi grooves V-shaped, shifted to posterior margin of slender, widely separated; ligula absent. Hypo- head capsule. Tentorial bridge absent. Pro thorax pharynx completely flattened. Prothorax dis- broad basally, narrowed toward head, wider tinctly larger than meso- and meta thorax. Legs than other segments. Legs rather long, with short, with equal claws. Urogomphi fused with shortened tibia; trochanter, femur and tibia with tergite IX, unsegmented, number of setae re- apical whirls of stout setae; claws subequal, each duced (as in Trechinae). Pygopodium longer with a long seta. Abdominal segments with nu- than urogomphi. merous long setae. Hypopleurites not apparent in first instar. Urogomphi fused to tergite IX, not Omophroninae segmented, about 3 times longer than tergite IX. Apex of urogomphi formed by a hyaline append- Distribution. Represented by one genus with 60 age in first instar (Landry & Bousquet 1984). species in the Holarctic, Oriental and Afrotropi- cal regions, as well as Central America. Cicindelinae

Biology and Ecology. All species are strongly Distribution. World-wide, with five tribes, ap- hygrophilous and restricted to the immediate proximately 130 genera and 2000 species. The vicinity of water. The adults are nocturnal and Ctenostomatini comprise two genera, Pogonos- run very rapidly when hunting at night. Like the loma in Madagascar, and Clenosloma in tropical larvae, they spend the day in burrows in sand South America. The Collyrini are restricted with or clay. few genera to India and South East Asia, the Manticorini are restricted with two genera to Morphology, Adults (Fig. 7.8.1 B; Lindroth 1969; Southern Africa. Megacephalini and Cicindelini Beutel 1992). Characterised by an almost circu- are the most diverse and widespread tribes. lar outline, long and slender antennae and legs, and an unusual colour pattern with a pale Biology and Ecology. Adults of several genera are ground colour and darker, often metallic mark- characterised by diurnal habits, very rapid ter- ings. Head short and transverse. Prosternal pro- restriallocomotion, and excellent flying abilities. cess broad and apically truncate, covering the The larvae life in vertical or horizontal burrows mesoventrite entirely, thus joining the prothorax in soil, sandy substrates or rotten wood and are immovably to the pterothorax. Procoxal cavities predators with a highly specialised ambush strat- closed, but pro coxae contact the anterolateral egy and unusual morphological adaptations. concavities of the mesoventrite. Protibia of ani- sochaetous type, antennal cleaner distinctly elon- Morphology, Adults (Fig. 7.8.1 A, 7.8.2E-F).10- gated towards proximal part of tibia. Two proxi- 70mm. Body black in basal groups (e. g., Omus, mal protarsomeres of males strongly dilated. Amblycheila), with comparatively stout legs. Proximal mesotarsomere moderately dilated. Other genera with conspicuous colour pattern, Scutellar shield concealed. Elytra with 14-15 metallic areas and long and very slender legs striae, but without abbreviated sutural stria. (Cicindela, ) or a very slender, ant- Metepimeron indistinct. Alae well developed. like body (arboricolous genera). Compound eyes Median lobe sclerotized only in ventral half. large and protruding in diurnal cicindelines (e. g., Parameres moderately asymmetric. Cicindela auct.). Labrum large and broadened. Antennae inserted on dorsal side of head. Man- Morphology, Larvae (Fig. 7.8.6 F -G). With fos- dibles with several pointed apices, long, inter- sorial adaptations. Body tapering posteriorly. crossing in resting position. Prothorax elon- Head capsule comparatively large and pro- gated, prosternal process and postcoxal bridge nouncedly wedge-shaped, with strongly protrud- unusually broad. Antennal cleaner and protibial ing, triangular nasale. Stemmata large. Postocu- spurs terminal. Elytra without stria. Alae com- lar and cervical grooves lacking. Antennae con- pletely reduced (Omus) or well developed. Ob- spicuously held upward, inclined towards me- 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- rows prior to the capture of prey. Head strongly meres symmetrical, connected by jugal sc1erite. rounded laterally; ventral side strongly convex. Six stemmata of different size present; two pairs Morphology, Larvae (Arndt 1998b; Arndt & on dorsal side of head strongly enlarged (Fig. Putchkov 1997; Breyer 1989; Hamilton 1925; 7.8.8 A). Frons distinctly extended posteriorly, Knisley & Pearson 1984; Putchkov & Arndt coronal suture very short or absent. Posterodor- 1994) (Fig. 7.8.8). Strongly modified morpho- sal margin of head capsule emarginate, thus logically in correlation with their life habits. nearly reaching or reaching posterior margin of Head and pronotum strongly enlarged and frontale. Ridge on caudal part of parietale con- strongly sc1erotized in contrast to long and slen- nected with ridge on caudal part of frontale in der rest of body. Dorsal side of hyperprogna- Manticorini and Megacephalini, but separate in thous head and protergum form a functional other taxa. Nasale (Fig. 7.8.8 B) strongly pro- unit. Both parts together form a lid of the bur- truding, shovel-shaped, with anterior margin Carabidae Latreille, 1802 131

smooth and subtruncate or sinuate. Antennae Arndt 2000). Larvae of Collyrini and Ctenosto- with elongate, nearly equally sized antennomeres matini live in rotten wood contrary to all other 1- 3. Antennomere 4 slightly shorter. Sensorial groups and are characterised by several apo- appendage of antennomere 3 replaced by small morphies: width of nasale reduced, body flat- field of pores. Mandible slender; apical part tened, and (except in Pogonostoma) claws fused longer than basal part including triangular reti- with tarsus (Arndt & Putchkov 1997). naculum; penicillus absent. M. craniomandibu- laris internus unusually large and complex. Addi- Carabinae tional protergal muscle inserts on adductor ten- don (Cicindela; Breyer 1989). Cardo triangular; The subfamily comprises 9 tribes, four of which stipes slender, curved, strongly sclerotized, with (Carabini, Ceroglossini, Pamborini, and Cych- one or more spines on mesobasal margin and a rini) contain large and colourful species. The membranous field on dorsal side; lacinia small adults, especially of the largest genus Carabus, or absent; strongly sclerotized bar with three are often characterised by metallic cuticle with bristles present between stipes and palpomere 1; conspicuous elytral patterns. The group is para- palpomere 1 large, fused with or at least attached or polyphyletic, comprising with Cicindini, one to galeomere; galea as large as maxillary palp; of the most peculiar and enigmatic tribes. both galeomeres with thick bristles (Fig. 7.8.8 C). Prementum dorsally covered by multisetose Distribution. World-wide, but the great majority hypopharynx; ligula prominent, with two pairs of species live in the Northern Hemisphere. The of setae; bipartite ventral sclerite between pre- monotypic Notiokasiini are restricted to the mentum and labial palpomere 1 (Fig. 7.8.8 D) warm-temperate Neotropical region, Ceroglos- present or absent (Megacephalini and Manticor- sini to the Neotropical Andes, Pamborini to Aus- ini). Premental retractors very strong, arranged tralia and New Zealand. The Cicindini contain in an unusual manner (Cicindela; Breyer 1989). two monotypic genera, Cicindis from Argentina Dense preoral filter present, apparently arising and Archeocindis from Persian Gulf. from upper part of ligula (Cicindela; Breyer 1989: Fig. 5). Hypopharynx separated form dor- Biology and Ecology. Most representatives in- sal premental surface by a distinct fold but al- cluding Carabus and live on the most completely flattened. M. tentoriohypopha- ground as more or less generalized predators, ryngalis absent. Gular suture anteriorly limited and nearly all of them are flightless. Adults and by an Y-shaped posterior tentorial groove in Ci- larvae of Cychrini are specialized predators of cindelini; posterior tentorial groove T-shaped in snails. Adults and larvae of Calosoma are very all other taxa. Most parts of tentorium strongly active predators of caterpillars and adults and flattened (dorsal arms) or thin; posterior arms larvae of some species climb trees. Mass flights basally fused; thin U-shaped tentorial bridge to caterpillar outbreaks are recorded e. g., for with attached to posterodorsal margin of head Calosoma jrigidum. Larvae of Notiophilus are capsule by thin processes; anterior arms strongly highly specialised predators of springtails. developed, broadly connected with head capsule (Cicindela; Breyer 1989). Pronotum rhomboid Morphology, Adults. Size ranging from small (Fig. 7.8.4 B), distinctly different from smaller, (e. g., Notiophilus) to very large (e. g., Carabus laterally rounded meso- and metanota. Legs stout, with very short tarsus; anterior claw larger subgenus Procerus). Protibial spurs and antenna than posterior claw. Abdominal segments I-IV cleaning organ terminal (Opisthiini, Carabini, Cychrini) or one spur subterminal and antenna and VI - IX subequal, but segment V distinctly modified, dorsally enlarged as an "abdominal cleaning organ slightly prolonged proximally. hump", with separate anterior-, lateral- and cau- Prosternal process distinctly extending beyond dal sclerites and 2- 3 hooks inserted between hind margin of procoxae and apically tapering. them (Fig. 7.8.8 E). Urogomphi absent; pygopod Procoxal cavities open. Mesoventrite of carabine short, conical and multisetose. Chaetotaxy type. Metepimeron concealed. strongly modified. Unusual, flattened, split setae on head and pronotum present in many taxa. Morphology, Larvae (Fig. 7.8.4 A, 7.8.5 D). Lar- The following characters distinguish Manti- vae of Carabinae are remarkably diverse mor- corini from other larvae of Cicindelinae: anten- phologically. None of the features which charac- nal base inserted anteroventrad of stemmata and terize the subfamily as a whole are autapo- separate from base of mandible by wide sclero- morphic: cervical and ocular grooves absent; hy- tized area; antennomere I very thin; tibia dis- podon (central tooth) usually present (absent in tinctly curved with a field of numerous short se- Opisthius); antennal muscles not intercrossing; tae on posterolateral side; number of stemmata mandible usually with penicillum; hypopharynx reduced, two enlarged stemmat a situated on distinctly bulging, with preoral filter; M. tentori- prominent region of head capsule (Oberprieler & ohypopharyngalis medialis usually present (ten- 132 Erik Arndt, Rolf G. Beutel & Kipling Will torial bridge interrupted and M. 42m absent in camaragnathus) or bifurcate (Hiletus) projec- Nebria); Mm. vertiCopharyngalis and tentorio- tions. Profemora of males almost always with pharyngalis well developed (Beutel 1992). ventral tooth (not in Southeast Asian Eucamara- Ceroglossini, Carabini, Pamborini and Cych- gnathus). Protibia of anisochaetous type. Anten- rini are characterised by strong sclerotization, re- nal cleaner poorly developed. Tarsi of all legs duced number of setae on tergites and sternites, slender and distally tapering. Adhesive setae a remarkably increased number of pores, and a present on protarsomeres 1-3 and mesotarso- reduced sensorial appendage of antennomere 3 meres 1- 2. Prosternal process long and narrow- (Arndt 1998a; Priiser & Arndt 1995). The uro- ing posteriorly. Procoxal cavity open. Meso- gomphi are markedly sclerotized, shortened, ventrite with hexagonal and anterolateral fused with tergite IX and increasingly reduced in grooves (carabine type). Mesocoxal cavities of the latter three taxa. The tergites are often ex- disjunct type. Elytral pattern diverse. Metepim- tended laterally (e. g., Cychrini). Nasal teeth and eron lobate. Alae well developed, with anterior setae FRIO,II are absent in Cychrini and many sector cell much larger than 3rd radial cell. Me- species of Carabus (Arndt & Makarov 2003). dian lobe unspecialised. Parameres asymmetriC, Larvae of Nebriini, Opisthiini, and Notiophilini usually brushy or multisetiferous (not in E. bra- are characterized by reduced setae TE6, PRs and siliensis). MEz, and the presence of a roughly pointed The thoracic structures display a unique com- microsculpture on all tergites. The urogomphi bination of "primitive" (prosternal process, pro- are thin, straight and moveably attached to ter- coxal cavity, mesoventrite) and derived features gite IX (Arndt 1993). Antennomere III is en- (lobate metepimeron). larged and multisetose in Opisthiini (Bousquet & Larvae. unknown Smetana 1991). A strongly protruding nasal re- gion with long, sharp and prominent teeth is Loricerinae characteristiC for Notiophilini and some taxa of Nebriini. A sclerotized bar is present between the Distribution. One tribe with two genera, the Hol- stipes and maxillary palpomere I in Notiophilini arctic (9 spp.) and the monotypic Ellip- (as in Cicindelinae; Arndt 1993). tosoma (Madeira).

Hiletinae Biology and Ecology. Species of Loricera are Distribution. Hiletinae (Hiletini) are a tropical more or less hygrophilous and characterised by group considered as rare and enigmatic (Erwin & specialised prey-catching techniques of larvae and adults, which involve the modified antennae Stork 1985). Approximately 20 known species or maxillae, respectively. They feed on spring- are arranged in 2 genera, Hiletus and Eucamara- tails. gnathus. Hiletus species occur in tropical Africa and species of Eucamaragnathus in Africa, Madagascar, south-eastern Asia, and in South Morphology, Adults (Jeannel 1941-42; Lindroth America east of the Andes from Ecuador to low- 1961-69). In general outline resembling a mid- land eastern Brazil (Erwin & Stork 1985). dle-sized Agonum (Loricera). Frons with 2 large foveae and a posterior median sulcus, emanating from a deep transverse constriction behind the Biology and Ecology. All species frequent lato- eyes. Antennae with long scapus and very long, solic soils in broadleaf evergreen and deciduous erect setae on pedicellus and antennomeres 3-6. forests or in grassland savannahs with scattered trees. Protibia of anisochaetous type. Procoxal cavity posteriorly closed by narrow bridge. Protho- racic-mesothoracic connection of harpaline type. Morphology, Adults (Erwin & Stork 1985; Beutel Mesocoxal cavity of disjunct type. Elytra with 12 1992). Head large and robust in proportion to regular stria (without abbreviated sutural stria). pronotum, with transverse sulcus whiChconnects Metepimeron parallel-sided and narrow. Copula- the sulcate frontal furrows. Antennae geniculate, tory organ short and oval, median lobe with with scapus as long as antennomeres 2-4. Sca- wide open basal orifice and elongated, asymmet- pus fits in groove below compound eyes. Mandi- riC apical part. Parameres apically rounded, bles large and markedly down turned distally, without fringes of hairs. Left paramere longer. creating a hollow concavity beneath. Each man- dible with eight or nine triangular teeth, which Morphology, Larvae (Arndt 1993, Luff 1993) increase in size distally. Lacinia enlarged later- (Fig. 7.8.9 A-B). Head capsule rounded later- ally, resembling an asymmetric club. Dorsal edge ally. Cervical and ocular grooves absent. Nasal with numerous spatulate spiculae. Galea long region with two acute teeth and a row of small and finger-like. Mentum with deep median con- sharp teeth beneath them. Antenna 2 X longer cavity, anteromesally produced into simple (Eu- than mandibles. Mandible siCkle-shaped; retina- Carabidae Latreille, 1802 133

o

AB o Fig. 7.8.9A- B, , 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; penicillum Protarsomeres I - IV or I- III dilated in males. present (Fig. 7.8.9 A). Stipes very large, almost Elytral striae feeble, irregular, usually disturbed entirely sclerotized; galea mesally directed at by fovea or tubercles. Thoracic structures other- right angle to stipes, longer than maxillary palp; wise similar to Loricerinae. Copulatory organ apical galeomere with divided into a swollen, with a long sclerotized stylet protruding through finely granular basal portion and a long, whip- the basal orifice of the median lobe when in re- like apical portion; distal part covered by hyaline pose. Parameres with fringes of hairs at the apex secretion (Fig. 7.8.9 B); field of setae on maxil- and ventral edge, less dense on the left paramere lary palp reduced; stipital field of setae reduced, in Elaphrus. its vestiges shifted to the middle of the inner margin. Labial palp slender, with secondarily Morphology, Larvae (Goulet 1983, Luff 1993) subdivided apical palpomere (Loricera sp.; (Fig. 7.8.6 D- E). Body slender and subcylindri- Thompson 1979); ligula unusually large, broadly cal (Luff 1993: Fig. 21). Head without ocular and rounded anteriorly, multisetose. Chaetotaxy of cervical grooves (Luff 1993: Fig. 22). Coronal su- frontale and tergites modified; flattened, split se- ture distinct, moderately long or short. Nasale tae on pronotum, mesonotum and metanotum centrally produced, triangular, with serrate present (see Cicindelinae). Legs long; tarsi elon- (Elaphrus) or smooth (Blethisa) lateral edges. gated, with 2 unequal claws; anterior claw larger, Mandible with large retinaculum and penicillus. with basal seta; seta absent from posterior claw. Cutting edge of mandible finely serrate in many Urogomphi slender and nearly as long as head species. Lacinia reduced, very short or ring- and thorax combined, fused to medially divided shaped. Pro thorax largest segment of body; pro- tergite IX, in later instars with numerous setifer- notum quadrangular. Other segments laterally ous nodes. Pygopod short and conical. rounded. Equal claws elongate, more than 0.5 as long as tarsus. Urogomphi fused with tergite IX, Elaphrinae in second and third instars multisetose with setae Distribution. One tribe comprising 3 genera, Di- inserting on small tubercles or with 10 long setae acheila, Blethisa and Elaphrus, restricted to the on very large nodes (Fig. 7.8.6 D- E). Holarctic region. Migadopinae Biology and Ecology. Elaphrines are usually strongly hygrophilous, and often riparian and Distribution. A relatively small group of two very active during the daylight. tribes with 15 genera mainly distributed in the subantarctic region (Andes, southern part of Morphology, Adults (Fig. 7.8.3; Jeannel 1941- South America, Falkland Islands, southern Aus- 42; Lindroth 1961-69; Beutel 1992). Middle- tralia and Tasmania, Southern New Zealand, sized, always with metallic lustre. Eyes large and Auckland Islands) (Jeannel 1938; Moret 1990; prominent. Prothorax with coarse punctuation. Baehr 1999). 134 Erik Arndt, Rolf G. Beutel & Kipling Will

Biology and Ecology. Most species of this tribe disjunct. Elytra distinctly flattened, without live in cool and south temparate rain forests and basal margin, with 8 stria (indistinct in Siagona). moorland. They live on the ground, and are Metepimeron parallel-sided, not lobate. Aedea- flightless with the exception of one species. gus large, laterally compressed. Parameres de- void of hairs in Luperca and Enceladus. Morphology, Adults (Jeannel 1938). Medium sized beetles ranging from 6-20 mm, with gla- Morphology, Larvae (Fig. 7.8.9 C- D) (Greben- brous surface. Almost always with one supraor- nikov 1999a). The larvae of Siagoninae are char- bital seta (absent in Aquilex diabolicola; Moret acterised by many apomorphic features. Head 1990). Pronotum and elytral disc without setae. with more or less constricted neck; cervical Elytra with one additional stria between Isl and groove absent (Siagona, Fig. 7.8.9 C) or com- 2nd primary striae. Wings always absent. Tho- pleted to a dorsal parietal keel (Enceladus); ocu- racic structures otherwise similar to Loricerinae. lar groove lacking; coronal suture long; 6 stem- Parameres well developed, at least right para- mata present or number of stemmata reduced to mere fringed with hairs. two or one. Nasal region protruding, serrate, shovel-shaped; adnasale with a dense group of Morphology, Larvae (Johns 1974). Three dif- setae. Antennomere III without sensorial ap- ferent larvae of Loxomerus and the larva of an pendage; antennomeres III and IV multisetose undetermined genus were described so far (Johns with membranous area on ventral side or ex- 1974). Internal features are unknown. Head with tremely elongate and whip-like (Fig. 7.8.9 D). more or less constricted neck region. Ocular and Mandible with large retinaculum; subapical seta cervical groove absent. Coronal suture very long. MN2 longer than retinaculum; penicillus and Nasale with three or five teeth including a hypo- seta MN I on outer mandibular margin absent. don; mandible straight and slender, with large Maxillary and labial palpi strongly elongate (Sia- retinaculum; retinaculum with additional sub- gona), or of normal size but with terminal seg- basal tooth; penicillus present; maxilla with sti- ments enlarged and with extended sensorial pes, palpi and galea long and slender; lacinia ab- fields (Enceladus). Lacinia present. Ligula sent. Prementum wide; ligula small. Legs with broadly rounded. Legs slender, with two claws; two claws, the anterior longer than the posterior setae of claws shifted to pretarsal sclerite. Ab- one. Urogomphi short, articulating on membra- dominal tergite IX distinctly reduced, divided nous region of tergite IX. Hypopleurite with cap- medially, fused with multisetose and extremely like gland on dorsal margin in second and third elongate urogomphi (Fig. 7.8.9 C). Urogomphi instar larvae (absent in first instar). about 0.5-1.0 as long as remaining body.

Siagoninae Scaritinae A small group comprising two tribes, Enceladini Scaritinae, which are probably not monophyletic represented by the monotypic genus Enceladus (northern South America) and Siagonini re- (s. below), comprise small to very large species. presented by the genera Siagona (southern Palae- They are are always strongly constricted between arctic including Malay Archipelago and Philip- the pro- and mesothorax. pines, Africa; 80 species), Cymbionotum (south- ern Palaearctic excluding Malay Archipelago, Biology and Ecology. Scaritinae are mainly char- Africa) and Luperca (Africa, India; two species) acterised by their habits of burrowing in soil or (Darlington 1967; Erwin 1978). sandy substrates.

Biology and Ecology. Species of Cymbionotum Distribution. Scaritini (sensu lato) occur world and Siagona live on the ground in wet areas with wide, but most of taxa are concentrated in the decaying vegetation or like Enceladus under Southern Hemisphere. Promecognathini with bark. Some occur on mud and in soil clefts of five genera are restricted to southernmost Africa desiccating waters. Luperca were found in termi- and northwestern North America. Salcediini oc- taria. cur with three genera in the pantropical region.

Morphology, Adults (Jeannel 1941-42). Moder- Morphology, Adults (Jeannel 1941-42). Dis- ately depressed and glabrous (Enceladus, Luper- tinctly depressed (e. g., ) or almost cylin- ca) or strongly depressed and pubescent (Sia- drical (e. g., Dyschirius). Pro thorax strongly nar- gona). Body always strongly constricted between rowed posteriorly, thus body distinctly con- pro- and mesothorax. Mandibles short, without stricted between pro- and pterothorax. Protho- seta. Palps short and narrow. Prothorax strongly racic-mesothoracic connection of harpaline type. narrowed posteriorly. Pro thoracic-meso thoracic Mesocoxal cavities disjunct. Metepimeron paral- connection of harpaline type. Mesocoxal cavities lel-sided, not lobate. Carabidae Latreille, 1802 135

Morphology, Larvae (Fig. 7.8.6 H, I). One of the wide, with several teeth, slightly protruding; an- most heterogenous groups concerning larval tenna short and stout; antennomere III with sen- morphology (Thompson & Allen 1974; Peyrieras sorial appendage; mandible slender, with small 1976; Thompson 1979; Nichols 1986; Bous- retinaculum; finely serrate on inner margin; laci- quet & Smetana 1986; Arndt 1991a; LufT 1993; nia and ligula present; legs as in Dyschirius and Moore & Lawrence 1994). in contrast to all other known Scaritinae with Promecognathini: Only the first instar larva of single claw; urogomphi fused with tergite IX, as one species of Promecognathus is known and dif- long as pygidium, but flattened in dorso-ventral fers strongly from all other taxa in this subfamily direction (Fig. 7.8.6 H); muItisetose in second (Bousquet & Smetana 1986). Head quadrangu- and third instars or urogomphi and pygopod lar, parallel-sided; stemmata reduced; cervical with several large horns (Thompson 1979). groove absent; coronal suture absent; posterior Dyschirius and related groups: Larvae strongly angles of frontale bulging, extended to posterior sclerotized, brown to black coloured, only head margin of head. Nasale straight, smooth, not sometimes red. Head wider than long with six protruding; antenna slender; antennomere III stemmata, nasale protruding, crown-like, poste- without sensorial appendage but with sensorial rior angles of frontale wide or slender and long; field apically on ventral side; mandible slender, antenna stout but sensorial appendage on anten- with small retinaculum; penicillus lacking in con- nomere III present; mandible slender with retina- trast to other known scaritine larvae; lacinia and culum present; lacinia absent; legs with a single ligula lacking; palpi short and stout, urogomphi claw; urogomphi short and stout, much shorter club-like, attached to tergite IX by a membra- than pygopod or tergite IX (Fig. 7.8.6 I; Arndt nous area; body and urogomphi multisetose even 1991a; LufT 1993). in first instar. The larvae of several subtribes and of the tribe Scaritine subtribe Carenina: Body strongly Salcediini are still unknown. sclerotized; lateral margins of head subparallel; six stemmata present; coronal suture long; cervi- Trechinae cal groove present; anterior margin of frontale Trechinae are a much disputed problematic smooth, straight, nasale not protruding; anten- nae extremely short; mandible stout, without group and probably a paraphyletic assemblage (see below). It is unclear whether Patrobini, Apo- retinaculum; lacinia elongate, as long as galea; tomus, and Psydrini should be included. maxillary palpi modified, only penultimate The inclusion of Melaenini and Cymbionotini article of normal shape; ligula present; labial (Lawrence & Newton 1995) is also problematic. palpi short and wide, with very large terminal sensorial field; epipleurites and hypopleurites fused on all abdominal segments; urogomphi ab- Distribution. World wide, a very diverse group. sent (Moore & Lawrence 1994). Subtribe Pasimachina: Body slender, subparal- Biology and Ecology. Highly variable, s. above. lel, strongly sclerotized and dark; head wider than long; six stemmata present; cervical groove Morphology, Adults. (Fig. 7.8.1 C; Jeannel 1941- lacking; coronal suture short; nasale protruding, 42). Male protarsi often with unilaterally dilated shovel-shaped, with smooth anterior margin; an- proximal tarsomeres. Mesepimeron broad or tenna long and slender, with sensorial appendage narrow. Otherwise harpaline type character com- absent from article III; mandible slender with bination of thoracic sclerites, i. e. closed procoxal small retinaculum; lacinia and ligula absent; both cavities, apically truncate prosternal process, palpi of normal structure; urogomphi fused to harpaline type mesoventrite, conjunct mesocoxal tergite IX, turned inside, longer than pygopod cavities, lobate metepimeron. (Thompson & Allen 1974). Subtribe Scaritina: Larvae strongly sclero- Morphology, Larvae. The larvae of Trechinae are tized, slender, elongate, subparallel; head quad- close to the ground plan of "higher Carabidae" rangular; coronal suture very long; cervical (Arndt 1993). Body moderately flattened, paral- groove shallow; nasale trapezoid; antenna long lel-sided. Head roughly quadrangular; cervical and slender with antennomere 3 flattened latero- groove present; head laterally with 6 stemmata apically, sensorial field present but appendage arranged in two rows, number reduced in cave absent on antennomere III; mandible long and dwelling taxa. Ocular groove present; frontal su- slender; lacinia absent; galea and palpi slender, ture sinuate; egg bursters consisting of a keel, of normal structure; ligula present; urogomphi isolated teeth or microspines. Nasale serrate, slender, fused with tergite IX (Peyrieras 1976; protruding. Antennomeres I and II subcylindri- Thompson 1979; Nichols 1986; Arndt 1991a). cal; antennomere III with bulb-like sensorial and related groups: Body yellowish, appendage inserted laterally; antennomere IV not strongly sclerotized; stemmata absent; coro- smaller, rounded apically. Mandibles with retina- nal suture and cervical groove present; nasale culum; terebrum with single cutting edge, 136 Erik Arndt, Rolf G. Beutel & Kipling Will smooth, or serrate in some taxa of Anillini and Arndt 1993; Fig. 7.8.6J). The membranous notch ; penicillus present or absent. Stipes is regarded as rudiment of a complete transverse slender; lacinia absent. Prementum with ligula. division of stipes as occuring in Cnemalobus Hypopharynx separated from dorsal surface, (Roig-Jufient 1993). flattened, with dense field of microtrichiae. An- The subgroups with most derived larval char- tennal muscles arranged crosswise (Beutel 1993). acters are the Licinini, Panagaeini and related Thorax with large pronotum and smaller, sub- groups (van Emden 1942; Liebherr & Ball 1990; equal meso- and metanotum, the latter two with Arndt 1991b), and the so-called "Truncatipen- anterior keel. Legs slender, with single claw in all nia", e. g., Anthiini (Arndt & Paarmann 1999), taxa except Perileptus, Amblystogenium, Thalas- , Orthogoniini (Makarov 1998), Gra- sophilus, and Patrobini (s. Psydrini, Gehringiini). phipterini (Zetto Brandmayr et al. 1993), Dryp- Lateral and dorsolateral area above coxal base tini (Habu & Sadanaga 1965),Galeritini (Arndt & with four small sclerites (episternum, epimeron, Drechsel 1998), and Lebiini (part.) (Lieftinck & trochantin, pleurite, the latter two are lacking on Wiebes 1968;Arndt 1989;Capogreco 1989;Arndt prothorax). Abdominal tergites I- VIII similar in et al. 2001). structure; tergites with anterior keel and median ecdysial sutures; sternites consisting of one large Pseudomorphinae median plate and smaller paired anterior, inner and outer (latero- )sternites. Abdominal segment Distribution. An unusual group of carabids dis- IX smaller, with fused sternal plates; tergite IX tributed in the Oriental region, Australia (Adelo- fused with urogomphi. Abdominal tergite X topus, Cryptocephalomorpha, Cainogenion, Paus- cylindrical, directed downwards (pygopod). sotropus, Sphallomorpha) and the New World Number of setae on urogomphi in second and (). third instars reduced, seta URj3 always lacking. Setae TA3-6 lacking in Trechini, Zolini, Bembi- Biology and Ecology. Pseudomorphine adults are diini (incl. Anillina), and Pogonini (Arndt et al. mainly found under bark (Baehr 1994). The 1999, Grebennikov 1999b; Arndt 2000; Greben- known larvae (except Sphallomorpha) are physo- nikov & Maddison 2000). Larvae of Apotomini, gastric due to their myrmecophilous habits. The Melaenini and Cymbiotonini are unknown. known larvae of Sphallomorpha dig holes in the ground around ant nests and prey on ants in a sim- ilar manner like cicindeline larvae. Pseudo- Harpalinae morphines are the only known carabids with Distribution. World wide, the most diverse group ovoviviparity (Liebherr & Kavanaugh 1985). with more than 30 tribes and 20000 species. Morphology, Adults (Notman 1925; Baehr 1992). Biology and Ecology. Highly variable, s. above. Body form elongate and more or less parallel and moderately convex (Pseudomorpha), very elon- Morphology, Adults (Jeannel 1941-42). Mandi- gate and cylindrical, resembling Scolytidae (Ade- bles without scrobal seta. Male pro tarsi never lotopus part.), or broadly oval and depressed with unilaterally dilated proximal tarsomeres. (Sphallomorpha). Colour dark piceous to nearly Procoxal cavities uni- or biperforated. Elytra black (Sphallomorpha part.) or variegated with complete or apically truncate (e. g., Odacan- maculae, vittae or pale margins. Head deflexed thini, Lebiini). (Cryptocephalomorpha) or horizontal, with deep antennal grooves. Compound eyes on dorsal side Morphology, Larvae (Figs.7.8.4C-E, 7.8.6J). of head (Adeloptopus, Cainogenion) or lateral, The tremendous number of species and supra- continuous border beneath eye present in Adeloto- specific taxa included in this subfamily is re- pus. Scapus partly visible from above. Mandibles flected by an extreme diversity of larval charac- without visible scrobes, very small in Cryptoceph- ters. Taxa which are presumably close to the alomorpha. Maxilla with (Cainogenion, Paussotro- ground plan of the subfamily, such as some pus) or without a large lateral lobe in most genera. Pterostichini and Platynini, are similar to Trech- Submento-mental suture absent. Prosternal pro- inae in most features described above, but have cess present or absent (Paussotropus). Legs short, a short lacinia and double claws. All larvae of with strongly developed femora. Configuration of this subfamily are characterised by the reduction thoracic sclerites ofharpaline type (see above). Six of the second ligular seta to a pore (Arndt 1993). abdominal sternites visible. Parameres setose or The antennal muscles are arranged crosswise as glabrous, fairly symmetrical or highly asymmetri- in larvae of Trechinae and Brachininae (Beutel cal but not 'balteate' (Baehr 1992). 1993). A membranous band is present ventrolat- erally on the stipes in many representatives (Mo- Morphology, Larvae (Fig. 7.8.10) (Erwin 1981, rionini, Pterostichini, Zabrini, Panagaeini, Pe- Baehr 1997). Larvae with abdomen curved in lat- leciini, Callistini, Oodini, Licinini, Harpalini; eral view and rows of spines on tergites V-VII Carabidae Latreille, 1802 137

pointed out by Forsyth (1972) and others (see below).

Distribution. Crepidogastrini with 7 genera (more than 100 species) are distributed in tropi- cal and southern Africa, two species of Tyronia occur in India and Sri Lanka. Brachinini with 8 genera (and more than 500 species) occur world B wide, but most of the species are distributed in tropical 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 at the 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, Fig. 7.S.IOA, dytiscoides, habitus, first instar; with narrow mesepimeron. Abdomen with 7 vis- B, Adelotopusrubiginosus,head, first instar, dorsal aspect; ible sternites in females and 8 visible sternites in C, ventral aspect (A-C, Baehr 1997). males. Copulatory apparatus with the median lobe often deformed, the right paramere strongly reduced, and the left paramere strongly sclero- (Sphallomorpha, Moore 1974) or physogastrical tized. Female genital tract with appended sper- with membranous, poorly sclerotized body mathecal gland, segmented gonocoxae, gono- (Fig. 7.8.IOA). Head subquadrate (Fig. 7.8.10B- coxal rami absent, and in some taxa, a digitiform C) or elongate and slightly narrowing anteriorly, diverticulum of the spermathecal duct present without stemm ata and egg bursters. Frontale (Liebherr & Will 1998). very wide, frontal suture not sinuate, broadly reaching posterior margin of head; coronal su- Morphology, Larvae (Fig.7.8.6A-C). There is ture, ocular and cervical groove absent. Nasale little known about larvae of Brachininae. Only flat, convex or triangular, smooth, without teeth. Brachinini are known in the larval stage. Few Mandible without retinaculum or penicillus. first instar larvae are described (see Arndt 1993 Lacinia absent; galea 1- or 2-segmented, more or for a review), and a single species is known in all less reduced; maxillary palp 4-segmented, large instars (Erwin 1967). in relation to other maxillary parts, longer than Following characters distinguish Brachininae stipes. Ligula broadly rounded or absent. Pro no- from the ground plan of Carabidae and are re- turn sclerotized; other tergites largely or entirely garded as autap omorphic character states: ante- membranous. Legs with double claws of dif- rior margin of frontale membranous, teeth and ferent length. Urogomphi absent. Chaetotaxy setae lacking. Posterior angle of frontale very strongly modified, larvae often multisetose or slender, V-shaped, egg bursters lacking in first in- with large club-like, flattened or split setae on star. Setal group gMX lacking, Lacinia absent head (Fig. 7.8.10 B-C), dorsal head appendages, (7.8.11 B). Ligula and setae LA3.4.6lacking. Uro- thoracic and abdominal tergites (Baehr 1997; Er- gomphi reduced or absent (7.8.11 C). Later instar win 1981). larvae physogastrical, legs and head appendages :t reduced (Arndt 1993). The antennal muscles Phylogeny and . A branching pattern are arranged crosswise, a feature shared by lar- (Sphallomorpha + (Pseudomorpha + (Crypto- vae of Brachininae, Harpalinae and Trech- cephalomorpha + (Adelotopus + (Cainogenion + inae(BeuteI1993). Other internal head structures Paussotropus»») was proposed by Baehr (1994). are similar to those of harpaline larvae exam- Brachininae ined. Hypopharynx completely flattened. Func- tional mouth narrow. Mm. tentoriohypopharyn- Mainly characterised by a complex abdominal galis, verticopharyngalis and tentoriopharyngalis explosive mechanism. A comparable apparatus absent. The shift of the brain to the prothorax is only present in Paussinae. However, distinct (Beutel 1993: Fig. 27) is a result of miniaturisa- structural differences in both groups were tion. 138 Erik Arndt, Rolf G. Beutel & Kipling Will

Phylogeny and Taxonomy. Phylogenetic hypothe- have noted that their extraordinary life history ses for Carabidae, originally simply the classifi- involving wedging through wood and feeding on cation of presumed natural groups, have contin- slime-molds has likely resulted in members of ued to develop and be refined as advances in this group being highly apomorphic both as technology have permitted researchers to use adults and larvae. The apparent deviation from more and finer details of behaviors, morphology, the primitive form makes it difficult or impos- chemistry and gene sequence data. The extensive sible to clearly assess the homology of many knowledge of carabid beetles makes them well adult and especially larval structures. Rhyso- suited to cladistic methodology, which is more dines are generally separate from Carabidae be- frequently being used to rigorously test hypo- cause of their distinctly divergent form, however, theses of relationships previously posited. Ball little evidence exist that Carabidae excluding (1979) and Ball et al. (1998) provide an excellent Rhysodidae is a monophyletic group. The ar- detailed review of the history of classification rangement of prehypopharyngeal setae in the and phylogenetics for the family. Brief and useful larva (Beutel 1993) and development of pubes- synopses with notes on relationships and distri- cence of the antennomeres in adults (Beutel bution of each tribe were published by Bous- 1995) have been suggested as possible synapo- quet & Larochelle (1993). morphies for Carabidae not including rhyso- Recent phylogenetic analyses that bear on re- dines. These characters were presented to sup- lationships within Adephaga, affecting our un- port a working hypothesis that excludes rhyso- derstanding of the root for the family, or for ex- dines from carabids. Unfortunately, neither can emplars from across many tribal level taxa in be considered very convincing given the varia- carabids, include a variety of character systems. tion involved in these structures. Notably, 18S rDNA sequence data (Maddison Various external structures found in adult rhy- et al. 1998, 1999; Shull et at. 2001), female repro- sodines such as the form of the medial septum ductive tract (Liebherr & Will 1998), larval mor- that separates the procoxal cavities, the closure phology (Arndt 1993, 1998), cuticular and mus- of the procoxal cavities and so-called ani- cular morphology (Beutel 1992, 1993; Beutel & sochaete form of antennal cleaner, seem to indi- Haas 1996) have all been analysed. Each of these cate a placement within Carabidae (Bell 1967; studies, with their strengths and limitations, has Beutel & Haas 1996). This is consistent with provided some insight into the evolutionary his- characters of the female reproductive tract (Lieb- tory of the family. herr & Will 1998), though this character system The placement of Trachypachidae relative to did not provide any decisive synapomorphies for Carabidae and Hydradephaga has varied over a specific sister-group relationship, the prefer- time and among authors. Some analyses have fa- red phylogenetic hypothesis placed rhysodines voured a sister-group relationship to Dytiscoidea within Carabidae in a grade with . The (e. g., Arndt 1993, 1998; Beutel 1993; Arndt & shared condition of a divided gonocoxite IX is Beutel 1995; Beutel & Haas 2000) while others the only evidence that supports the inclusion of place them as sister to Carabidae, including rhy- rhysodines within anisochaetous carabid taxa. sodines (e. g., Kavanaugh 1986; Beutel 1998). Bell (1998) concluded that rhysodines, and Given the conflicting results from analyses based what he considered a subtribe of scaritines, the on both adult and larval characters, Shull et al. salcediines (Clivinini), were sister taxa. However, (2001) used 18S rDNA sequence data to attempt Scaritini is most likely not monophyletic, clivi- to resolve the relationships among the families of nines may only be distantly related to Scaritini Adephaga. In all but one alignment and analysis sensu stricto, with some evidence supporting a strategy used by Shull et al. (2001) Trachypach- closer relationship of rhysodines to Scaritini than idae was associated with the remaining Geade- to Clivinini (see below). Molecular sequence evi- phaga and not Dytiscoidea. Ultimately, 18S dence from the 18s gene (Maddison et al. 1998, rDNA sequence data, additional sequence data 1999; Shull et al. 2001) are problematic due to from other regions and characters from mor- likely convergence of several potentially aberrant phology should be combined in a single analysis sequences including rhysodines, scaritines, cicin- to see if sufficient overall agreement can be delines and paussines. Although the authors view found among character sets. Although analyses these associations as potentially spurious, addi- thus far do not unequivocally establish the rela- tional taxon sampling and various methods over tionships among adephagan families, all rein- the course of the several papers has not altered force the distinctness of the trachypachid lineage, this result. None of these analyses have associ- its great age and relictual nature. ated rhysodines and clivinines nor has any Rhysodidae is treated in this volume as a sepa- placed rhysodines outside of Carabidae. As rate family from Carabidae, however, many au- noted (Maddison et al. 1999) it is likely that this thors have treated them as a subfamilial member particular sequence cannot properly place basal of Carabidae, potentially related to some group carabid taxa. It is also possible that some of of Clivinini (Bell 1998) (s. 1-7.9). Many authors these taxa are in fact related. A relationship be- Carabidae Latreille, 1802 139 tween and Scaritini, exclusive of Clivi- some Carabinae (Carabini, Cychrini, Pamborini) nini, should not be discounted as these taxa also (Beutel 1998; Liebherr & Will 1998) a relation- share a peculiar structure, the vesicula seminalis ship that suggests the unique form of the tiger (referred to as "mesidemia" by SmrZ (1981, beetle female abdomen is derived from the car- 1985)), not found in any other carabid tribe abine type (Liebherr and Will 1998). Addition- (SmrZ 1981, 1985; Will unpubl.). ally, both Cicindelinae and some carabines The larval form is strikingly different in rhy- (Calosoma) produce aromatic aldehydes as a ma- sodines, unlike any carabid taxon and salcediines jor constituent of their defensive secretions have apparently typical carabid larvae (Arndt, (Moore & Wallbank 1968; Schildknecht et al. pers. obs.). Bell's (1998) proposed association of 1968). However, pygidial gland secretory cell ag- rhysodines and salcediines (Clivinini), or any as- gregation type and structure are more similar to sociation with a particular tribe within Carab- the typical hydradephagan form (Forsyth 1970). idae, is not supported by larval characters (s. Larval characters suggest a close relationship of 1-7.9). Cicindelinae and Loricerinae (Arndt 1993). Subdivisions within Carabidae have been Combined analysis of adult and larval characters placed at a variety of levels over the years (Ball places them as a member of a more derived 1979; Ball et at. 1998). The subfamilies used by grade than carabines together with loricerines, Lawrence & Newton (1995), which have been elaphrines and scaritines (Beutel & Haas 1996). adopted for use in this volume, vary greatly in Published analyses of 18S ribosomal DNA the type and relative support for their mono- (Maddison et al. 1999; Shull et al. 200 I) are phyly. Several of the subfamilies contain rela- problematic with regard to the placement of tively few species and only one or a small cicindelines within carabids, and their study sug- number of genera, e. g., Gehringiinae, Omophron- gests that the 18S rDNA sequence is probably inae, Hiletinae, Loricerinae, Elaphrinae and Sia- not sufficient for resolving relationships of the goninae. Each of these is well supported by apo- subfamily. Results of these analyses of sequence morphic characters and are distinct in form, but data are consistent with a placement of cicinde- their relationships to other subfamilies are not lines within Carabidae, but are not considered well understood. Other subfamilies have many conclusive as the sequence divergence of Cicinde- more taxa and vary considerably in regard to the lini is likely to strongly effect results due to con- support for their monophyly and evidence for vergence. sister-group relationships. Paussinae is a very well supported monophy- Although the monophyly of Cicindelinae is letic group based on morphological characteris- well supported, the position of tiger beetles in tics including aspects of the pygidial glands and Carabidae is still a question as contentious as the substantial larval characters (Beutel 1995). They relationships for the families Trachypachidae share with Brachininae similar chemical com- and Rhysodidae discussed above. Although no pounds and delivery of these defensive secretions character analysis has placed them outside of by crepitating. However, the form of the glands, Carabidae, they are still frequently classified that details of the secretory cells and mode of delivery way, with some authors maintaining them as a differs dramatically between these groups (Eisner separate family. However, this issue in particular, et al. 2000, 2001). Paussinae is generally consid- and the phylogenetic structure of Carabidae in ered adelphotaxon to the remaining carabid sub- general, is dependent on the choice of the rooting families or one of the basal-most lineages, how- point for the family. Recently it was proposed ever, evidence for this is limited. This basal posi- (Pearson & Vogler 2001) that Cicindelinae be tion is supported by female reproductive track maintained as a family separate from Carabidae and larval characters (Arndt 1993; Liebherr & given results from studies by Bils (1976) and Will 1998). However, DNA sequence data (Shull Nichols (1985). However, neither Bils nor Nich- et at. 2001) consistently places Paussinae as a ols tested the monophyly of Carabidae excluding much more derived group and never as sister to Cicindelidae. Rather they chose Cicindelinae the remaining Carabidae. Shull et al. (2001) sug- (-idae) as the outgroup and so fixed the rooting gest the intriguing, and only slightly less parsi- point between tiger beetles and other carabids. monious, possibility that a clade of Paussinae + Therefore these analyses provide no evidence for Brachininae lies at or near the base of Harpal- the exclusion of Cicindelinae. In fact, re-rooting inae. This would, as pointed out by Shull et al., the phylogenies presented by Bils and Nichols require the reversal of a large number morpho- using Trachypachidae, Gyrinidae or , logical characters thought to define the general as suggested by results of recent studies on ade- pattern of relationships in the family. phagan families (Beutel 1998; Beutel & Haas Gehringiinae, like nebriines and carabines, 2000; Shull et al. 2001), places Cicindelinae well have what is thought to be the plesiomorphic within the family Carabidae. Adult morphologi- thoracic configuration for Carabidae: confluent, cal structures consistently place tiger beetles internally unbridged and open procoxae, meso- within Carabidae and often associate them with ventrite with anterolateral grooves for reception 140 Erik Arndt, Rolf G. Beutel & Kipling Will

of the procoxae and with a median hexagonal Several of the subfamilies (Hiletinae, Ela- groove). As this is the plesiomorphic arrange- phrinae, Migadopinae, Siagoninae and Scarit- ment these structures do not provide grouping inae) are taxa thought to occupy the mid-grade information and emphasis for classification has of the carabid phylogeny. Overall, these taxa been placed on whether the arrangement of the show a trend of extensive reconfiguration of the pro tibial spurs and antennal cleaning organ rep- thorax (Beutel 1992; but not in Hiletinae) and resents an isochaete or anisochaete condition female abdomen (Deuve 1993; Liebherr & Will (Bell 1967; Beutel 1992; Beutel & Haas 1996). 1998) from the organisation found in the basal The condition of the gehringiine tibia is not taxa leading up to the Harpalinae radiation. This clearly assigned. If considered as isochaetous set of taxa (excluding Clivinini from Scaritinae), Gehringiinae is placed as sister to Paussinae Carabinae (excluding some Nebriitae), Cicindel- (Beutel & Haas 1996), if anisochaetous, they inae and some members of Trechinae, have a could be sister to Nebriitae. Placement within sclerotized ramus mesad of the base of the fe- anisochaete taxa and near nebriines is consistent male gonocoxite that is considered a significant with the dimerous condition of the gonocoxite synapomorphy. There is little agreement among and dual spermatheca in the female tract (Lieb- various data set as to the relationships of this herr & Will 1998). Larval characters have not heterogeneous set of taxa. Certainly some of the proven decisive for grouping Gehringiinae and subfamilies that fall along this grade are not no results have been published based on DNA likely to be monophyletic. sequence data. Scaritinae, excluding Promecognathini, is only The rounded body shape of species in Omo- weakly supported as monophyletic in some phroninae immediately sets them apart from all analyses of 18S rDNA sequence data (Shull et al. other adephagans. Details of their adult mor- 2001) and thought to be polyphyletic based on phology are equally unique and this has resulted female reproductive structures. Larval character- in a wide range of proposed relationships for the istics offer no synapomorphies for Clivinini + subfamily. Some authors have even placed them Scaritini (Arndt 1993). No recent analysis con- outside Carabidae with Hydradephaga (Bils firms the placement of Promecognathini with 1976; Nichols 1985), but like Cicindelinae dis- other members of Scaritinae as established by cussed above, re-rooting these phylogenetic Lawrence & Newton (1995). As noted above, the peculiar form of the vesicula seminalis that is hypotheses maintains Omophroninae as a mem- known from Scaritini sensu stricto but not found ber of Carabidae. Larval characters clearly sup- Clivinini is known from Rhysodidae (SmrZ 1981, port their inclusion within carabids (Beutel 1991; Arndt 1993) and suggest a position near the base 1985; Will unpubl.). This is consistent with a re- of Anisochaeta or with Nebriitae. A close rela- lationship of rhysodines and scaritines found with 18S rDNA sequence data (Maddison et al. tionship between Nebriitae, Omophroninae and 1999; Shull et al. 200 I), but interpreted as arte- Gehringiinae was suggested by Liebherr and Will fact by those authors. (1998) based on a hypothesised substitution of As suggested by Erwin (1985) Migadop- spermathecal function in these taxa. Analyses of inae(ini) has not been supported as monophy- 18S rDNA sequence data clearly place Omo- letic in any recent analyses. Liebherr & Will phroninae in Carabidae and in the basal grade (1998) placed Amarotypini as sister to Promec- of non-Harpalinae taxa (Maddison et al. 1998, ognathini while Migadopini was placed in a 1999; Shull et al. 2001). However, no consistent grade sister to Carabidae Limbata of Jeannel sister-group relationships were found in the vari- (1946). Based on 18S sequences data Maddison ous analyses. et al. (1999) found that Amarotypus was best Carabinae as delimited by Lawrence & New- placed near but not as sister to genera of Miga- ton (1995) is almost certainly not monophyletic. dopini. A novel relationship of Loricerini + Mi- As discussed above Cicindelinae may be closely gadopini (excluding Amarotypus) was strongly related to some carabine but there is no sugges- supported in those analyses. tion of a Carabinae + Cicindelinae clade that in- The traditional arrangement of Trechinae (in- cludes nebriite taxa. There is moderate support cluding Apotomini, Broscini, Melaenini, Psyd- for the monophyly of Nebriitae from both DNA rini, Bembidiini, Pogonini, Patrobini) or Sty- sequence data (Maddison et al. 1999) and mor- lifera of Jeannel (1941) is clearly not monophy- phological data (Kavanaugh & Nt~gre 1982; Ka- letic. In addition to the various hypotheses for vanaugh 1996). However, there is no evidence some of these taxa discussed above, Apotomines beyond plesiomorphic similarity that Carabitae were included by Jeannel (1946) based on an in- and Nebriitae form a monophylum. Addition- correct assessment of the arrangement of the ally, female genital tract characters suggest that thoracic sclerites. Both morphological characters Siagoninae may be sister to the carabine + Ci- and 18S sequence data are in agreement that cindelinae clade, though this relationship is not apotomines are not particularly closely related to supported by other character systems. these taxa, however, an alternate placement is Carabidae Latreille, 1802 141 not clear. It has been demonstrated repeatedly published. Some highly distinctive taxa are un- that Psydrini is not monophyletic (Arndt 1993, doubtedly monophyletic but clearly some, per- 1998; Baehr 1998; Liebherr & Will 1998; Maddi- haps most, of the major tribes are not. Arndt son et al. 1999; Shull et al. 2001). Psydrines fall (1998) presents evidence for grouping some taxa out into at least two groups, neither thought based on larval characters. A group including to be closely related to other Trechinae taxa. A Pterostichitae, Panagaeitae, Callistitae, and Har- monophylum of Trechitae + Patrobini is quite palitae is supported by a shared condition of a plausible. Trechitae monophyly is supported by membranous band on the lateroventral surface achiasmatic male meiosis (Serrano 1981) and a of the larval stipes. The villous canal extending sister-group relationship of Patrobini is indicated along the common oviduct in the female repro- by larvae (Arndt 1993), female abdominal struc- ductive tract was suggested as a possible synapo- ture (Deuve 1993) and molecular sequence data morphy for Orthogoniini, Panagaeini, Melanchi- (Maddison et al. 1999). tonini, Graphipterini, Licinini and Loxandrini. Members of Brachininae like Paussinae, use The presence of a basal sclerite of the sperma- an explosive discharge of hot quinones for chem- theca is synapomorphy from the female tract ical defence. Based on this alone it has been sug- grouping Geobaenini, Pseudomorphini, Lachno- gested that these two groups are closely related. phorini and Odacanthini (Liebherr & Will 1998). However, as discussed above, they differ signifi- Defensive chemical data and structures of the cantly in both adult and larval features. Brachin- pygidial gland system may also provide clues to inae larval characters have been interpreted as the affinities of Harpalinae tribes. Orthogonitae primitive (Arndt 1993), but no synapomorphies as circumscribed by Erwin (1985, 1991) is cer- tying them to Paussinae have been identified tainly a polyphyletic group with member taxa (Arndt 1998). Although Arndt (1998) pointed most likely related to a variety of tribes. Abdom- out that there are no external synapomorphies in inal structures in Catapiesis Solier, Brachidius larvae for these two groups, Beutel (1993) iden- Chaudoir Oxyglychus Straneo and Cratocerus tified an antennal musculature arrangement that Dejean are probably synapomorphic for these was synapomorphic for Brachininae and Harpal- taxa, exclusive of Orthogoniini (Will et al. 2000). inae. The female reproductive tract characters Defensive chemical data and molecular sequence and form of the abdomen (Liebherr & Will data recently collected for Catapiesis (Will et al. 1998), thoracic structure and larval characteris- 2000, unpubl.) and Brachidius (Will unpubl.) are tics (Beutel 1992, 1993) all seem to place this consistent with a sister-group relationship of group in or near Harpalinae. Confirmation of these taxa. Secondary chemical compounds sup- hypotheses from these morphological systems is port grouping these with members of Dryptitae found in 18S sequence data (Maddison et al. and Perigonini. 1999), where moving Brachininae from sister of Harpalinae to a basal position adds forty-nine Acknowledgements steps to the parsimony tree. We wish to thank James Liebherr (Cornell Uni- The great subfamily Harpalinae contains the versity) and Dave Kavanaugh (California Acad- bulk of species and greatest range of character emy of Sciences) for helpful comments on earlier variation. Despite considerable heterogeneity of the member taxa in terms of body form and life versions of this chapter. history, this is one of the most clearly monophy- letic subfamilies in Carabidae based on a variety Literature of character systems including the form of the female abdomen and reproductive tract (Deuve 1993; Liebherr & Will 1998), larvae (Arndt Acorn, J., H. & Ball, G. E. (1991): The mandibles of some adult ground beetles: structure, function, and 1993), and molecular sequence data (Maddison the evolution of herbivory (Coleoptera: Carabidae). et al. 1998, 1999;Shull et al. 200 I). The only seri- - Canadian Journal of Zoology 63 (3): 638-650. ous challenge to the monophyly is Brachininae Adis, J. & B. Messner (1997): Adaptations to life under as discussed above. Pseudomorphinae have a water: Tiger Beetles and millipeds. Pp. 319-330 in highly apomorphic form apparently in response Junk, W. J. (ed.) The Central Amazon Floodplain. to the group's evolution into myrmecophily. Its Ecological Studies 126. Springer, Berlin, Heidel- placement within Harpalinae, best treated as a berg. tribe therein, has been confirmed in all recent Arndt, E. (1989): Beschreibung der Larve von Lio- analyses. nychus quadrillum (Duft.) und Bemerkungen zur Relationships of taxa within Harpalinae, the Larvalsystematik der mitteleuropaischen Gruppen des Subtribus Dromiina (Insecta, Coleoptera, presumed radiation of the majority Carabidae: ). - Reichenbachia Museum of extant carabid taxa, remains delphic. No Tierkunde, Dresden 27: 47-56. analysis of any significant number of harpaline (1991 a): Carabidae. Pp. 45-141 in Klausnitzer, B. taxa for a character system that is able to provide (ed.) Die Larven der Kafer Mitteleuropas, Bd. 1. resolution among the included tribes has been Goecke & Evers, Krefeld. 142 Erik Arndt, Rolf G. Beutel & Kipling Will

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sification of Caraboidea (Coleoptera: Adephaga). - Sioanoglymmiina Bol/ettino del Museo Regionale di Scienze Naturali, Sloanoglymmius Bell & Bell, 1991: Australia ATTI, Torino. Clinidiina: Will, K.W. (1998): Oviposition behavior in Grouvellina Bell & Bell, 1978: Madacascar, piceus Dejean (Coleoptera: Carabidae, Lebiini). - Mayotte (Comoro Islands) Fabreries 23 (3-4): 120-123. Will, K. w., Attygalle, A. B. & Herath, K. (2000): New Rhyzodiastes Fairmaire, 1895: Oriental and defensive chemical data for ground beetles (Coleo- Australian regions, eastern South America ptera: Carabidae): Interpretations in a phylogenetic Rhyzotetrops Bell & Bell, 1985: Fiji framework. - Biological Journal of the Linnean So- Rhyzoarca Bell & Bell, 1985: New Caledonia, ciety 71: 459-481. New Zealand, Australia Zetto Brandmayr, T. (1990): Spermatophagous (seed- Temoana Bell & Bell, 1985: Taiwan, Pacific Is- eating) ground beetles: first comparison of the diet lands, Australia westwards to Andamans, and ecology of the harpaline genera Harpalus and eastern India, western China Ophonus (Col., Carabidae). Pp 307-316 in Stork, Rhyzostrix Bell & Bell, 1985: northern South N. (ed.) The Role of Ground Beetles in Ecological America and Environmental Studies. Intercept, Andover, Rhyzodiastes s. str.: eastern Brazil Hampshire. Zetto Brandmayr, T., Brandmayr, P., Marano, I. & Kirby 1895: Holarctic region, Central America, northern South America Paarmann, W. (1993): The larva of serrator (Forskiil 1775) (Coleoptera Carabidae Mexiclinidium Bell & Bell, 1978: highlands of Graphipterini): description and functional mor- Mexico and Guatemala phology. - Tropical Zoology 6: 299- 312. Protainoa Bell & Bell, 1978: western Cuba Tainoa Bell & Bell, 1978: Greater Antilles Arctoclinidium Bell & Bell, ]978: Europe, Iran, Japan, western North America, eastern USA 7.9. Family Rhysodidae Laporte, 1840 Clinidium s. str.: Central America, West Indies, northern and western parts of South (= Rhysodini s. Bell) America Rolf G. Beutel Omoglymmiina Xhosores Bell & Bell, 1978: Natal and extreme eastern part of Cape Province, South Africa Yamatosa Bell & Bell, 1979 substituted for Ya- Distribution. World-wide, with a higher diversity matoa Bell & Bell, 1978 (preoccupied): Ori- in tropical regions. Especially prominent in insu- ental Region, from the Hima]ayas to Japan lar faunas. and Java Leoglymmiina Bell & Bell, 1978:confined to Aus- Shyrodes Grouvelle, ]903: Burma tralia Srimara Bell & Bell, ]978: Vietnam Dhysorina: Plesioglymmius Bell & Bell, 1978: Greater Tangarona Bell, 1982 substituted for Tangaroa Sunda Islands, Malay Peninsula, Mindanao, Bell & Bell, 1978 (preoccupied): North Is- Brazi], Cuba land of New Zealand Ameroglymmius Bell & Bell, ]979: Cuba, Vene- Dhysores Grouvelle, 1903: Africa zue]a, southern Brazi] Neodhysores Bell & Bell, 1978: Brazil Juxtaglymmius Bell & Bell, 1979: Borneo, Ma- Medisorina: lay Peninsula, Java Medisores Bell & Bell, 1987: South Africa Plesioglymmius s. str.: Sumatra, Borneo, Cele- Rhysodina: bes, Mindanao Rhysodes Dalman, 1823: Europe, Asia Arrowina Bell & Bell, ]978: southern India, Sri Kupeus Bell & Bell, 1982 substituted for Ku- Lanka, southern Japan poea Bell & Bell, 1978 (preoccupied): North (sensu lato) Ganglbauer, 1892: Island of New Zealand by far the largest genus of Rhysodidae, al- Bell & Bell, 1978: Pacific islands most cosmopolitan, but absent from Mada- from New Caledonia and New Zealand to gascar and New Zealand Australia and Tasmania, westwards to Min- Hemoglymmius Bell & Bell, 1978: Greater danao and Sulawesi Sunda Is]ands, Africa (I species) Angekiva Bell & Bell, 1979: Australia Boreoglymmius Bell & Bell, 1982: North Amer- Ingevaka Bell & Bell, 1979: North Island of ica, Japan New Zealand Pyxiglymmius Bell & Bell, ]978: Oriental Re- Vakeinga Bell & Bell, 1979: North Island of gion, from Japan to Java, Sumatra, Anda- New Zealand and New Caledonia man Islands and Luzon Kaveinga s. str.: Pacific islands from the Solo- Laminoglymmius Bell & Bell, 1982: Malay mons and Australia (Cape York) to Minda- Peninsula, Sumatra, Borneo, New Guinea nao and Sulawesi Navita Bell & Bell, 1978: Fiji, New Hebrides