The Brontini of the world: A generic review of the tribe

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The Brontini of the world: A generic review of thetribe (Coleoptera: Silvanidae: Brontinae)Michael C. ThomasFlorida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, Gainesville, FL

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1INSECTA MUNDI, Vol. 17, No. 1-2, March-June, 2003

Introduction

Examining more than 3,000 specimens repre-senting all described species of Uleiota Latreille fromnumerous collections over the past decade convincedme that more than one genus was masquaradingunder that name and led to this study. Studies ofindividual genera will follow.

Thomas (1984) divided the Silvanidae into twosubfamilies based primarily on genitalic characters,including the genera with inverted male genitalia inUleiotinae (now Brontinae (Lawrence and Newton1995)) and the genera with non-inverted genitalia inSilvaninae. The Brontinae was further subdividedinto two tribes: the Brontini (Uleiotini sensu Thomas1984) and the Telephanini, based primarily on thecondition of the anterior coxal cavities (open in Bron-tini, closed in Telephanini). Pal et al. (1985) provideda taxonomic history of the higher categories of Sil-vanidae and divided the family into four subfamilies:Uleiotinae (= Uleiotini sensu Thomas 1984), Crypta-morphinae and Psammoecinae (together = Telephani-ni sensu Thomas 1984), and Silvaninae.

Brontini Erichson 1845

Description:Form: Elongate, fusiform to ovate, mostly elon-

gate, parallel-sided; size small to medium, 5mm to15mm in length.

Surface sculpture and pubescence: Mod-erately to densely punctate; punctures of two types:simple in non-incrusted species, subtending an erector suberect simple seta, or broad, shallow, and ocel-late in incrusted species, subtending an erect orsuberect modified seta (truncate and broadened api-cally or thick and strongly curved). An integumentalincrustation occurs in Uleiota, most Macrohyliota,Brontopriscus, and Brontoliota. Microsculpture ispresent in non-incrusted species, absent in mostincrusted species. Pubescence is moderate, usuallyshort and inconspicuous, occasionally long.

Head:Head capsule: Transverse, somewhat tri-

angular in shape; frontoclypeal suture usually notobvious, but apparently indicated laterally by a shortline or by an abrupt elevation change; apex of clypeussubtruncate, usually deeply emarginate over anten-nal insertions; frontal region bounded laterally by

The Brontini of the world: A generic review of the tribe(Coleoptera: Silvanidae: Brontinae)

Michael C. ThomasFlorida State Collection of Arthropods

Florida Department of Agriculture and Consumer ServicesP.O. Box 147100

Gainesville, FL 32614-7100 U.S.A.

Abstract: The genera of the tribe Brontini (Silvanidae: Brontinae) are reviewed. The tribe is considered here tobe composed of 12 genera, Uleiota Latreille, Brontopriscus Sharp, and Dendrophagus Schönherr, plus nine newgenera: Australodendrophagus, Australohyliota, Brontoliota, Dendrophagella, Macrohyliota, Megahyliota,Microhyliota, Parahyliota, and Protodendrophagus. Aplatamus Grouvelle is removed from the Brontini and placedin the Telephanini. Four new species are described: Protodendrophagus antipodes Thomas; Brontoliotaindivisipennis Thomas; Brontoliota intermedius Thomas; and Brontoliota monteithi Thomas.

Described species are assigned to genera with the following new combinations resulting:Australodendrophagus australis (Erichson); Australohyliota chilensis (Blanchard); Australohyliota macleayi(Olliff); Denrophagella capito (Pascoe); Macrohyliota truncatipennis (Heller); Macrohyliota bicolor Arrow;Macrohyliota gracilicornis (Arrow); Macrohyliota lucius (Pascoe); Macrohyliota militaris (Erichson); Macrohyliotaspinicollis (Gory); Megahyliota feae (Grouvelle); Microhyliota integricollis (Fairmaire); Parahyliota africanusGrouvelle; Parahyliota alticola (Pal, Sen Gupta, and Crowson); Parahyliota atratus (Grouvelle); Parahyliotabrevicollis (Arrow); Parahyliota cinamomeus (Fairmaire); Parahyliota costicollis (Reitter); Parahyliota fallax(Grouvelle); Parahyliota indicus (Arrow); Parahyliota pallidus (Arrow); Parahyliota puberulus (Reitter); Parahyliotaserratus (Smith); Parahyliota serricollis (Candeze); Parahyliota siamensis (Arrow).

Two new synonymies are proposed: Uleiota crenicollis Grouvelle (=Uleiota costicollis Grouvelle) and Uleiotatexana Dajoz (=Uleiota dubius (Fabricius)). Uleiota truncatus Motschulsky, formerly treated as a subspecies ofU. dubius (Fabricius), is elevated to a full species, new status.

2 Volume 17, No. 1-2, March-June, 2003, INSECTA MUNDI

longitudinal grooves in some genera (frontal regionsubject to sexual dimorphism in Parahyliota); tem-ples slightly to markedly produced; head usuallytransversely impressed behind temples and abruptlyconstricted to form a neck; gular region simple orsemicircularly impressed; gular sutures present, wide-ly separated, divergent posteriorly.

Mouthparts: Mandibles (Figs.3-12 ) stout,laterally expanded or carinate, with two apical teethand a dorso-basally located, small to large mycangiumbounded anterolaterally with a tubercle; prosthecaand mola present; males of some species of severalgenera with an erect dorsal mandibular horn (Figs. 1,7), the function of which is unknown. Labrum short,transverse, apparently immoveable. Mentum trans-verse (Fig. 13). Labial palpomeres (Fig. 13) subequalin length, not strongly expanded (except Dendropha-gus); terminal palpomere simple; galea broadly rounded

and densely setose; lacinia (Fig. 14) with two apicalteeth and one subapical tooth. Labial palps withapical palpomere (Fig. 13) moderately to stronglysecuriform(except simple in Dendrophagus and in anintermediate form in Dendrophagella); ligula (Fig.13) corneous and emarginate apically.

Antennae: Filiform, elongate, sometimesexceeding body length; terminal antennomeres notforming a distinct club; scape elongate to very elon-gate, usually at least as long as head; pedicel usuallymuch shorter than antennomere III; antennae highlymodified in males of some species of Parahyliota, thefunction of such modified antennae is unknown.

Eyes: Moderate to large, flattened to hemi-spherical; conspicuously setose in a few species ofMacrohyliota.

Figures 1-2. Uleiota dubius. 1) Dorsal view, habitus of male; 2) Ventral view, habitus of female.

21


Figures 3-11. Uleiotini, mandibles, dorsal view: 3) Dendrophagus cygnaei; 4) Parahyliota siamensis; 5) Australohyliota macleayi;6) Dendrophagella capito; 7) Macrohyliota gracilicornis; 8) Macrohyliota militaris; 9) Australohyliota chilensis; 10)Australodendrophagus australis; 11) Macrohyliota bicolor.

11

3 4 5

6 7 8

9 10


Thorax:Pronotum: Quadrate to transverse, lateral

margins denticulate to strongly toothed (except Den-drophagus); disk sometimes bounded by longitudinalcarinae or costae; present in almost all members ofthe tribe is what appears to be a glandular pore locatedin a laterobasal excavation at the hind angles of thepronotum.

Prosternum: Anterior coxal cavities moder-ately to broadly separated, open posteriorly; intercox-al process broad; protrochantin not exposed.

Scutellum: Moderate, rounded or angulateposteriorly, usually without distinct transverse cari-na or groove.

Mesosternum: Meets metasternum in analmost straight line, suture simple; mesocoxal cavi-ties widely separated, open laterally; closed bymesepimeron only, mesepisternum does not contrib-ute to closure; mesotrochantin exposed.

Metasternum: Transverse (very short andtransverse in Brontopriscus), with median longitudi-nal line for half or more of length; metendosternite asin Fig. ; Pal et al. (1985) reported “...a pair of smallglandular cavity-like structure [sic] present at apicalmargin of metathorax...”. I have been unable to locatesuch structures. However, in many Brontini themetasternum immediately behind the mesocoxae isexcavate to a greater or lesser degree.

Elytra and hind wings: Elytra striatopunc-tate, disk usually with six or seven rows of puncturesplus a scutellary striole (absent in Parahyliota, Bron-topriscus, and Uleiota); intervals flat to costate;humeral carina present or absent; lateral marginexplanate to a greater or lesser degree, sometimesdenticulate; epipleura usually complete to apex, mod-erate to very broad; elytra are fused in Brontopriscusand Brontoliota, in which all known species lack hind

Figures 12-14. Uleiota dubius, female. 12) right mandible; 13) mouthparts; 14) lacinia.

12 14

13


wings. Hind wing venation well-developed, remark-ably consistent throughout the tribe.

Legs: Usually short, occasionally elongate;femora stout; tibiae short, usually simple, but occa-sionally with basal spine or carina; tibial spurspresent, short, subequal; tarsomeres not lobed (tar-someres are ventrally emarginate in some Parahylio-ta, but fleshy pads are not present (Fig. 17)); tarsalformula 5-5-5 in both sexes (except in Uleiota, inwhich tarsomere I is fused with II making the tarsalformula 4-4-4 (Fig. 16)); tarsi of two forms: (Den-drophagus-type) tarsomere I and IV very short, II-IIImuch longer (Fig. 15); (Uleiota-type) tarsomeres I-IVall short and subequal in length (Fig. 16-17); tarsalclaws simple.

Abdomen: With five subequal sterna, punctate,without sexual modifications; intercoxal process nar-row to broad; femoral lines closed (open in someParahyliota).

Genitalia: Male genitalia of normal, invertedcucujoid type. Parameres located on the ventral as-pect of the median lobe, usually elongate and articu-lated (e.g., Fig. 55-57), reduced to tooth-like processesin Parahyliota (Fig. 59); internal sac often withcomplex armature and usually with a flagellum.

Discussion of selected characters

Integumental incrustation: An opaque, brown,somewhat granular substance (Fig. 18) coats theintegument of all species of Uleiota, Brontopriscus,Brontoliota, Megahyliota, and most species of Macro-hyliota. Such an incrustation does not appear to occuranywhere else in the Silvanidae, except in the silva-nine Silvanoprus (Halstead 1993). It often has bits ofdebris imbedded in it and usually completely obscuresthe surface. It is not soluble in alcohol, ethyl acetate,or water, but can be removed in a detergent solutionusing pins to peel it off the surface. An ultrasoniccleaner can speed the process, but can destroy thespecimen if care is not used. The incrustation occursgenerally on the entire exterior surface except forantennomeres II-XI, eyes, mouthparts, and tarsi. Theamount of incrustation varies among individuals andit may be that the incrustation grows heavier as theindividual ages. The surface coating most probablyfunctions as camouflage, but may also be a deterent topredators.

I have examined with a scanning electron micro-scope representatives of two species in which individ-uals are incrusted: Uleiota dubius and Macrohyliota

Figure 18. Specimen of Brontoliota monteithi showing moderateincrustation and phoretic uropodine mites that are commonlyassociated with beetles of this tribe.

Figures 15-17. Tarsi of Brontini. 15) Dendrophagus cygnaei; 16)Uleiota dubius; 17) Parahyliota costacollis.

17

15

16


n. sp. On the elytra of U. dubius are alternating rowsof tubercles and punctures (Fig. 22). Each subtends aseta, which arises dorsally from the tubercles (Fig.24) but is situated at the lateral edge of the punctures

(Fig. 23). The tubercles also have one or two openingsnear the base of the seta that appear to be glandularin nature (Fig. 24). The setae situated at the edge ofthe punctures are inclined over the puncture and are

a

22 23 24

Figures 22-24. Uleiota dubius: 22) Enlargement of elytral striae; 23) Enlargement of strial puncture; showing modified seta; 24) Enlargementof tubercle on elytral interval, showing modified seta and pore.

20 21

Figures 19-21. Macrohyliota n.sp.: 19) base of elytron; 20) enlargement of area enclosed in rectangle of a, showing modified setae,puncture, pores and microsetae; 21) enlargement of area enclosed in rectangle of b, showing modified seta.

19

2020


Figures 25-28. 25) Australodendrophagus australis; 26) Australohyliota chilensis; 27) Australohyliota macleayi; 28) Brontoliota indivisipennis.

25 26

27 28


Figures 29-32. 29) Brontoliota intermedius; 30) Brontoliota monteithi; 31) Brontopriscus pleuralis; 32) Dendrophagella capito.

29 30

31 32


Figures 33-36. 33) Dendrophagus cygnaei; 34) Macrohyliota gracilicornis; 35) Megahyliota feae; 36) Microhyliota integricollis.

33 34

35 36


angulate apically (Fig. 23). Setae of both the tuberclesand punctures are somewhat longitudinally grooved.In Macrohyliota n. sp. the setae are even more highlymodified, being either long, strongly grooved, andapically truncate, or short, thickened, fluted struc-tures that barely resemble setae (Fig. 20). Additional-ly, microspines are located between the setae. Situat-ed at the base of each seta are openings in theintegument that may be glandular (Fig. 21). All ofthese modifications can be interpreted as structuresto distribute and anchor the substance that forms thebase of the incrustation. Such extensive modificationsalso suggest that the incrustation performs a functionthat is important to the survival of the beetles or totheir reproductive success.

Head: Well-defined temples and posteriorly constrict-ed head capsule are character states of the family asa whole. Longitudinal grooves on the dorsal surface ofthe head are found in both the Brontini and Telepha-nini, but not Silvaninae. A frontoclypeal suture oc-curs in a few Brontini and some but not all Telepha-nini; it does not occur in the Silvaninae. Male sexualcharacters on the frons occur only in the Brontini(Parahyliota). An elongate antennal scape is presentin all Brontinae; absent in all Silvaninae. Male sexual

characters occur in the flagellum of the antennae inBrontini (Parahyliota) and a few Telephanini (Tele-phanus), but are not known to occur in Silvaninae.Male sexual characters do not seem to occur on thehindlegs of Brontini, but they do occur there in someTelephanini (Telephanus and Cryptamorpha) andSilvaninae.

Mouthparts: A dorsal mycangium on the mandiblesis found in the Brontinae but not the Silvaninae; itsabsence there is inferred to be derived. Male mandib-ular horns occur only in certain Brontini.

Thorax: Lateral teeth or spines on the pronotum arefound throughout the family. Anterior coxal cavitiesare open posteriorly in Brontini only. A well-markedtransverse groove or carina paralleling the hindmargin of the scutellum is present in some Telepha-nini. In the Brontini, only Dendrophagella capito hassuch a structure conspicuously developed; however,an inconspicuous groove is present in some otherbrontine genera; it is absent in all Silvaninae. Coxalcavities are moderately or widely separated in allBrontini; narrowly separated in all Telephanini (ex-cept Aplatamus, which was assigned to Uleiotini byThomas (1984) and is here moved to the Telephanini)

Figures 37-38. 37) Parahyliota costicollis; 38) Protodendrophagus antipodes.

37 38


and most Silvaninae. Dendrophagus-type tarsi occurin both tribes of the Brontinae; Uleiota-type tarsi onlyin Brontini; lobed tarsi occur only in Telephanini andSilvaninae. Hindwing venation is very similar in allBrontinae with well-developed venation; hindwingvenation is reduced in some Telephanini and mostSilvaninae.

Elytra: A scutellary striole is present in most Bron-tini (except Parahyliota, Brontopriscus, and Uleiota)and some Telephanini, but is absent in all Silvaninae.

Male genitalia: The aedeagus is inverted (paramereslie on dorsal aspect of median lobe) in all Brontinae,the aedeagus is not inverted (parameres lie on ventralaspect of median lobe) in all Silvaninae. Complexarmature of the internal sac and a flagellum occurthroughout the family. Members of Macrohyliota,Brontopriscus, Dendrophagella, and Brontoliota pos-sess a median lobe that is emarginate distally. Themedian lobe in Uleiota, Megahyliota, Parahyliota,and Dendrophagus is entire and acute distally.Australodendrophagus and Microhyliota possess high-ly modified male genitalia that do not closely resembleeither group. The parameres in Uleiota and Mega-hyliota are rather broad and widely divergent apical-ly; the parameres in other genera are more narrowlydivergent apically and often nearly parallel-sided, orare otherwise modified.

Biology

Very little has been published on the habits andhabitats of this group of beetles, except that they occurunder loose bark where they feed on fungi of variouskinds. Savely (1939) reported finding only fungalhyphae and spores in the gut of Uleiota dubius.Crowson and Ellis (1969) studied the morphology andbiology of Dendrophagus crenatus and concluded thatit was clearly fungivorus and that the adult mandib-ular pits functioned as spore-transport mechanismsand were truly mycangia. Cekalovic and Quezada(1972) described the larva of Australohyliota chilen-sis and reported on the occurrence of adults and larvaeunder bark of Nothofagus. Pal et al. (1985) describedand illustrated the larva of Parahyliota indicus.

Distribution

Members of the Brontini are found on all conti-nents, but are absent from the American tropics (twospecies occur in Chile) and are poorly represented insub-Saharan Africa, where only two species of

Parahyliota (one described and one undescribed) oc-cur. The tribe is represented by the greatest numberof species and genera from Madagascar east to thePhilippines and the Solomon Islands, and both Aus-tralia and New Zealand possess diverse and endemicfaunas. The genera Uleiota and Dendrophagus arerepresented in the Holarctic with a total of fourspecies in the Nearctic, and two each in Europe andJapan.

Australodendrophagus Thomas, new genusFigs. 10, 25, 39

Type species: Dendrophagus australis Erichson, bypresent designation and monotypy.

Figure 39. Median lobe and tegmen: Australodendrophagusaustralis.


Diagnosis: The combination of head with shortlongitudinal lines (Fig. 25); subquadrate pronotumwith finely denticulate lateral margins and a short,broad tooth at the anterior angles; presence of ascutellary striole; and male genital structure is diag-nostic for this genus.

Description: With characteristics of Brontinae:Brontini as described above, plus: frontoclypeal su-ture absent, mandibles carinate dorsally, but notexpanded laterally; eyes large, hemisphaerical, morethan 1/3 length of head capsule; temple short, about0.4 length of eye;antennal scape longer than head.Pronotum more or less quadrate, laterally minutelydenticulate; anterior angles with short, broad tooth;posterior angles obtuse. Elytra with scutellary strioleand six rows of punctures, laterally costate. Proster-

nal process wider than a coxal cavity, slightly round-ed apically. Male genitalia as in Fig. 39.

Discussion: In general facies, adults of the singlemember of this genus most resemble members ofParahyliota, but differ in many details.

Etymology: Combination of the Latin adjective “aus-tralis” plus the genus name Dendrophagus, meaning“southern Dendrophagus.” It is masculine.

Included species: Australodendrophagus australis(Erichson), new combination; Australia.

Australohyliota Thomas, new genusFigs. 5, 9, 26-27, 40-41

Type species: Uleiota macleayi Olliff, by presentdesignation.

Diagnosis: The shape of the pronotum (Fig. 26-27),with short, broad anterolateral teeth, the compara-tively long pubescence of the dorsal surface andantennae, and the structure of male genitalia (Fig.40-41) characterize the members of this genus.

Description: With characteristics of Brontinae:Brontini as described above, plus: Surface withoutincrustation. Dorsal pubescence long, conspicuous.Head with frontoclypeal suture represented by slightchange in elevation; with longitudinal lines or not;temples present, short to long, rounded; basal lineimpressed; eyes large, convex to somewhat flattened;antennal scape longer to much longer than head;mandibles rounded, slightly expanded laterally. Pro-thorax more or less quadrate; laterally denticulate,teeth coarse to fine and inconspicuous; anterior angleswith longer, sharper teeth, directed somewhat dorsal-ly; posterior angles right to obtuse. Elytra withscutellary striole; with six punctate striae; laterallycarinate or not. Legs moderate to long, femora slen-der; tarsi “Dendrophagus-type”. Prosternal processas broad as or narrower than coxal cavity; roundedapically. Male genitalia (Fig. 40-41) with basal strutof aedeagus short and broadly spatulate; flagellumrecurved basally.

Distribution: Australia, Chile.

Discussion: The two species assigned to this genusseem to be rather isolated in the tribe, but may be

Figure 40-41. Median lobe and tegmen: 40) Australohyliotamacleayi; 41) A. chilensis.

40

41


closest to Brontopriscus and Protodendrophagus,based on the structure of the male genitalia.

Etymology: Combination of the Latin adjective “aus-tralis” plus the genus name Hyliota, meaning “south-ern Hyliota.” It is masculine.

Included species: Australohyliota chilensis (Blan-chard), new combination; Australohyliota ma-cleayi (Olliff), new combination.

Brontoliota Thomas, new genusFigs. 18, 28-30, 42-52

Type species: Brontoliota monteithi Thomas, bypresent designation.

Diagnosis: Adults of this genus (Figs. 28-30) aresimilar to those of Brontopriscus in their lack of hindwings, fused elytra, and short metasternum, butdiffer in their more elongate, narrow body form,possession of a scutellary striole, and structure of themale genitalia (Fig. 51-52).

Description: With characteristics of Brontinae:Brontini as described above, plus: Body long andslender; parallel-sided or fusiform. Dorsal surfacewith incrustation (Fig. 18). Head without frontocly-peal suture, but frontoclypeal region depressed belowgeneral level of head; longitudinal lines short,obscure;dorsal surface of head with five raised, moredensely setose areas, as follows: above antennal inser-tions, behind eyes, and medially; mandibles slightlyexpanded laterally; eyes small, very convex, set onshort stalks (Figs. 42-44); temple very long, angulateto rounded, with a deep longitudinal sulcus dividing

4442 43

Figures 42-44. Brontoliota, heads, dorsal view: 42) B. indivisipennis; 43) B. intermedia; 44) B. monteithi.

4745 46

Figures 45-47. Brontoliota, pronota: 45) B. indivisipennis; 46) B. intermedia; 47) B. monteithi.


it; basal line not or barely impressed; scape “flowerbud vase” shaped, about as long as head; antennaerather thicker than usual for genus. Pronotum quad-rate or fusiform; armed laterally with blunt teeth orspines, anterolateral angle with longest spine; poste-rior angle acute to obtuse; anterior and posteriormargins produced to form distinct neck-like regions,which are raised and transversely wrinkled. Elytradorsally flat, fused; basally constricted and toothedanterior to humerus; with six to eight rows of punc-tures plus a scutellary striole; alternate intervalsraised, intervals continuous basally but interruptedapically to form a series of elongate tubercles; raisedintervals more densely setose than rest of elytral disk;elytra laterally explanate, margin armed for entirelength; epipleura wide; hind wings absent. Legsrather long; femora moderately stout; tarsi “Den-drophagus-type”. Prosternal process wider than acoxal cavity, expanded and rounded apically. Malegenitalia (Fig. 51-52) very similar to those found inMacrohyliota, but with very much longer basal strutand flagellum.

Distribution: Queensland, Australia, with one spe-cies extending south into New South Wales.

Discussion: The affinities of these remarkable bee-tles are not clear. Their resemblance to Bronto-priscus, especially to B. sinuatus, may be because ofsimilar adaptations to winglessness or, perhaps, tosimilar habits or habitat. Dr. Geoffrey Monteith, whocollected most of the known specimens of this genus,

wrote (in. litt.) about their habitat: “I work with thebug family Aradidae which has a lot of bizarre wing-less forms which live on the outside of dead logs andsticks lying on the ground in rainforest, especially onthe moist underside which is partly in contact withthe soil and/or leaf litter. They are sedentary andhighly camouflaged, so I spend a lot of time picking upwood from the ground and peering intently at it untilmy critters swim into focus. So this is the situationwhere I find your beetles also. They sit perfectly still,flattened against the damp, mouldy bark surface.Rarely do you find one under loose bark, where theirmore normal winged relatives live.”

Etymology: The genus name is an arbitrary combi-nation of letters from Brontes and Hyliota. It ismasculine.

Included species: Brontoliota indivisipennis Tho-mas, new species; Brontoliota intermedius Thomas,new species; Brontoliota monteithi Thomas, newspecies.

Brontoliota indivisipennis Thomas, new speciesFigs. 28,42,45,48

Diagnosis: The shorter, more compact body (Fig.28), and entire elytral apices (Fig. 48) are diagnosticfor adults of this species.

Figures 48-50. Elytral apices of species of Brontoliota: 48) B. indivisipennis; 49) B. intermedius; 50) B. monteithi.

5048 49


Description: (male holotype): With characteristicsof Brontinae: Brontini: Brontoliota as described above,plus: Length, 8.3mm. Dark testaceous, tarsi andelytral margins paler, antennal flagellum darker.Head and pronotum with scattered, variably-sizedocellate punctures, smaller and less distinct than inB. monteithi, most dense medially and posteriorly,but irregularly placed; each subtending a thick,

curved, suberect setae; surface between punctureswithout microsculpture. Pronotum wider than long(1:1.2 length/width), more or less rectangular inshape, widest behind middle; with three low, longitu-dinal costae, situated medially and sublaterally, mostpronounced apically and basally; costae more denselysetose that remainder of pronotum. Elytra 2.4x longerthan wide, widest at about apical third; laterally

Figures 51-52. Brontoliota spp. 51) B. monteithi, median lobe and internal sac; 52) B. intermedius, tegmen, median lobe, basal partof internal sac, and sperm duct.

51 52


armed with irregular, short denticles, apices con-jointly rounded (Fig. 48). Male genitalia with apex ofmedian lobe broadly emarginate, as in Fig. 52.

Variation: Paratypes range in length from 5.6mm -7.3mm; elytral length/width proportions range from2.1 - 2.4; female antennae are a little shorter andstouter proportionally than those of the males, butthis is best seen in a series of specimens.

Etymology: The species name is from the Latin“indivisus,” meaning undivided, and “penna,” mean-ing feather or wing, referring to the entire elytralapices.

Types: Holotype male, deposited in the QueenslandMuseum with accession number QMT.108611, withlabel data as follows: “Cunninghams Gap, 17.xii.1965S.E.Q. B. Cantrell”. Paratypes (41) with label data inAppendix 1.

Brontoliota intermedius Thomas, new speciesFigs. 29, 43, 46, 49, 52

Diagnosis: Adults of this species resemble those of B.monteithi rather closely, but are less elongate (Fig.29); the elytral apices are not as prolonged (Fig. 49);there are few or no punctures on the head andpronotum; and the apex of median lobe is broadlyemarginate (Fig. 52).

Description: (male holotype): With characteristicsof Brontinae: Brontini: Brontoliota as described above,plus: Length, 7.1 mm. Dark testaceous, legs, mouth-parts, and elytral margins paler. Head and pronotumappearing impunctate, with thick, curved, suberectsetae which seem to arise directly from cuticle; sur-face without microsculpture. Pronotum quadrate (1:1length/width), more or less rectangular in shape, withtwo low, longitudinal costae sublaterally; costae moredensely setose that remainder of pronotum. Elytra2.3x longer than wide, widest just behind humerus;laterally armed with irregular, short denticles, apicesbifurcate (Fig. 49). Male genitalia with apex of medianlobe broadly emarginate, as in Fig. 52.


Etymology: The specific epithet refers to the inter-mediate position of this species.

Types: Holotype male, deposited in the QueenslandMuseum with accession number QMT.108612, withlabel data as follows: “Paluma Dam, N. Qld. 27.xii.1963.G. Monteith”. Paratypes (25) with label data in Appen-dix 1.

Brontoliota monteithi Thomas, new speciesFigs. 18, 30, 44, 47, 50, 51

Diagnosis: Adults (Fig. 18, 30) of this species aremore elongate than those of B. intermedius; theelytral apices are more prolonged (Fig. 50); head andpronotum punctate; apex of median lobe narrowlyemarginate (Fig.51). Additionally, the temples aremore rounded than in either of the other two Bronto-liota species (Fig. 44).

Description (male holotype): With characteristics ofBrontinae: Brontini: Brontoliota as described above,plus: Length, 8.6 mm. Dark testaceous, legs andelytral margins paler, antennal flagellum darker.Head and pronotum with scattered, variably-sizedocellate punctures, most dense medially but irregu-larly placed; each subtending a thick, curved, suberectsetae; surface between punctures without microscu-lpture. Pronotum just barely longer than wide (1:1.1width/length), distinctly fusiform in shape, widestbehind middle and strongly narrowed anteriorly andposteriorly; with low, longitudinal carinae parallelinglateral margin at about middle third; low, longitudi-nal costae situated medially to lateral costae, mostpronounced apically and basally, and a low, longitudi-nal median costa; costae more densely setose thanremainder of pronotum. Elytra 2.9x longer than wide,widest at about apical third; laterally armed withirregular, short denticles, apices strongly bifurcate(Fig. 50). Male genitalia with apex of median lobenarrowly emarginate, as in Fig. 51.


Etymology: I take pleasure in naming this speciesafter Dr. Geoffrey Monteith, who collected most of thespecimens known of this genus, and who gave un-


stintingly from his knowledge of their habits andhabitat.

Types: Holotype male, deposited in the QueenslandMuseum with accession number QMT.108613, withlabel data as follows: “Bellenden Ker Range, NQ 1kmS. of Cable Tower 6 Oct. 17-Nov. 5, 1981, 500mEARTHWATCH/QLD MUSEUM Pyrethrum knock-down”. Paratypes (112) with label data in Appendix 1.

Key to the species of Brontoliota

1. Elytral apices conjointly rounded (Fig. 48) ...........Brontoliota indivisipennis Thomas, new species

1'. Elytra apices divaricate (Figs. 49-50) ................. 2

2(1'). Head and pronotum impunctate, or nearly so; bodyless elongate (Fig. 29), elytra 2.5x or less longer

than wide; apex of median lobe broadly emargin-ate (Fig. 52) ........................................................... Brontoliota intermedius Thomas, new species

2'. Head and pronotum with large, conspicuous ocel-late punctures; body more elongate (Fig. 30),elytra more than 2.5x longer than wide; apex ofmedian lobe narrowly emarginate (Fig. 51) ............ Brontoliota monteithi Thomas, new species

Brontopriscus SharpFig. 31, 53

Brontopriscus Sharp 1886: 391

Type species: Brontes pleuralis Sharp, by originaldesignation. (Although Sharp (1886) did not explicitlystate the type species of Brontopriscus, he made thefollowing statement: “When I described Brontes pleu-ralis I stated it might be made the type of a new genus,and the discovery of a second species with the pecu-liarities still more exaggerated renders it advisablethat this should now be done.” The phrase “...thisshould now be done” clearly refers to making B.pleuralis the type of Brontopriscus and that this wasSharp’s intent.)

Diagnosis: The very wide epipleura, fused elytra,lack of an obvious scutellary striole, and absence ofhind wings are diagnostic for the members of thisgenus.

Figure 54. Median lobe: Dendrophagella capito.

Figure 53. Median lobe and tegmen: Brontopriscus pleuralis.


Description: With characteristics of Brontinae:Brontini as described above, plus: Dorsal surfacewith incrustation. Head without frontoclypeal su-ture; longitudinal lines short, obscure or absent;mandibles slightly expanded laterally; eyes moderate,convex; temple long, rounded; basal line impressed ornot; scape normal or “flower bud vase” shaped, longerthan head. Pronotum more or less quadrate; laterallywith blunt teeth, anterior and posterior angles acute.Elytra dorsally flat; fused, with six rows of punctures;without an obvious scutellary striole; laterally broad-ly explanate, margin minutely denticulate or not;epipleura very wide; hind wings absent. Legs ratherlong; femora moderate; tarsi “Dendrophagus-type”.Prosternal process wider than a coxal cavity, expand-ed and rounded apically. Male genitalia as in Fig. 53.

Distribution: New Zealand.

Discussion: Although the affinities of Brontopriscusseem to lay with Brontoliota, the male genitalia in thetwo genera are quite different. One or two puncturesoccur in the area of the elytron of B. pleuralis wherethe scutellary striole would be located, and they mayrepresent remnants of that. The elytra of B. sinuatusare abruptly constricted at the base, forming a char-acteristic “waist.” The narrowing of the elytra theremay have been the reason for the loss of the scutellarystriole in that species. They do not share the derivedtarsal characters of Uleiota or Parahyliota. The“flower bud vase”-shaped antennal scape seems to beunique within the Brontini with one of two Bronto-priscus species and all known species of Brontoliotapossessing it, but a similarly shaped antennal scapeoccurs in a few Telephanus (Telephanini). Sharp(1886) did not mention the flightlessness of thesebeetles.

Included species: Brontopriscus pleuralis (Sharp);Brontopriscus sinuatus Sharp.

Dendrophagella Thomas, new genusFigs. 6, 32, 54

Type species: Dendrophagus capito Pascoe, bypresent designation and monotypy.

Diagnosis: The combination of the pubescent dorsalsurface, elytral color pattern, long frontal lines, andDendrophagus-type tarsi is diagnostic for members ofthis genus.

Description: With characteristics of Brontinae:Brontini as described above, plus: Surface withoutincrustation, distinctly pubescent. Head with fronto-clypeal suture represented by a vague, broad, trans-verse impression; with long longitudinal lines; tem-ples present, short, rounded; basal line not impressed;eyes moderate, convex; antennal scape shorter thanhead; mandibles rounded, slightly carinate dorsally.Prothorax cordate, armed posterolaterally with sever-al denticles and at the anterior angles with a short,broad tooth. Elytra with scutellary striole; with sixpunctate striae; laterally vaguely costate. Legs mod-erately long, femora moderate; tarsi “Dendrophagus-type”. Prosternal process a little broader than a coxalcavity; rounded and weakly expanded apically. Malegenitalia (Fig. 54) characteristic.


Figure 55. Median lobe and tegmen: Dendrophagus longicornis.


Discussion: Dendrophagella capito shares the longfrontal lines and male genitalic characters with mem-bers of Dendrophagus, but differs from members ofthis genus in its distinctly pubescent dorsum, colorpattern, and shape of the pronotum. I have seen a veryfew specimens which were nearly uniformly piceousin color, but most specimens resemble the individualpictured in Fig. 32.

Etymology: The genus name is the feminine dimin-utive of Dendrophagus. The species epithet, capito,remains masculine as it is a noun in apposition.

Included species: Dendrophagella capito (Pascoe),new combination.

Dendrophagus SchönherrFigs. 3, 15, 33, 55

Dendrophagus Schönherr 1809: 50Hyliota Reitter, 1879: 80 (sensu Arrow 1901: 593, in

part)

Type species: Cucujus crenatus Paykull, by mono-typy.

Diagnosis: The combination of simple lateral mar-gins of the pronotum and obsolete anterior angles,long frontal lines (Fig. 33), and Dendrophagus-typetarsi is diagnostic for members of this genus.

Description: With characteristics of Brontinae:Brontini as described above, plus: Surface withoutincrustation, nearly glabrous. Head with frontocly-peal suture represented by a vague, broad, transverseimpression; with long longitudinal lines; templesabsent or present, rounded; basal line impressed; eyesmoderate, flattened to convex; antennal scape shorterto much longer than head; mandibles rounded, slight-ly carinate dorsally, with a short dorsal tooth in malesof one species. Prothorax quadrate, with obsoleteanterior and posterior angles; lateral margin sinuateand unarmed. Elytra with scutellary striole; with sixpunctate striae; laterally vaguely costate. Legs moder-ately long, femora moderate; tarsi “Dendrophagus-type”. Prosternal process a little broader than a coxalcavity; rounded and weakly expanded apically. Malegenitalia (Fig. 55) characteristic.

Distribution: Holarctic.

Discussion: The presence of dorsal mandibularhorns in one species of this genus (the Japanese D.longicornis Reitter) does not appear to have beendocumented previously.

Included species: Dendrophagus crenatus (Paykull);Dendrophagus cygnaei Mannerheim; Dendrophaguslongicornis Reitter.

Macrohyliota Thomas, new genusFigs. 7-8, 11, 19-21, 34, 56

Type species: Hyliota gracilicornis Arrow, by presentdesignation.

Diagnosis: These are relatively large, loosely jointedbeetles (Fig. 34). Most individuals are covered with abrown, granular incrustation that obscures the sur-

Figure 56. Median lobe and tegmen: Macrohyliota spinicollis.


face sculpture to a greater or lesser extent (individu-als of two species included here lack the incrustation).Individuals of most of the species have a tooth orcarina on the mesotibia. They have “Dendrophagus-type” tarsi, and in most of the species there are smallmandibular horns in the males.

Description: With characteristics of Brontinae:Brontini as described above, plus: Dorsal surface withincrustation in most species. Head without frontocly-peal suture; longitudinal lines short, obscure, orabsent; mandibles slightly expanded laterally; withdorsally directed tooth in males of some species; eyesmoderate, convex; temple moderate, rounded, absentin some species; basal line impressed or not; scapemuch longer than head. Pronotum more or lessquadrate; laterally with spines, anterolateral anglewith longest spine; posterior angle acute. Scutellumsemicircular. Elytra laterally carinate or not; discwith six rows of punctures plus a scutellary striole.Legs rather long for the tribe; femur moderatelystout; tibia carinate or toothed basally in most spe-cies; tarsi “Dendrophagus-type”. Prosternal processabout as wide as a coxal cavity, expanded androunded apically. Male genitalia (Fig. 56) with longbasal strut; long flagellum, and reduced armature ofinternal sac.

Distribution: Asia, Australia.

Discussion: Two species (M. bicolor and M. milita-ris) lack the incrustation found on the other species ofthe genus and may represent a different lineagedeserving of generic rank. However, except for thelack of an incrustation and the concomitant modifica-tions to surface sculpture and pubescence, M. bicoloris very close to M. lucius, a typical incrusted species.There are undescribed species of this genus fromAustralia, New Guinea, the Solomon Islands, and thePhilippines.

Etymology: The genus name is a combination of“macro,” the Greek word for “long”, and “hyliota,” theGreek word for “forester”. It is masculine.

Included species: Macrohyliota bicolor (Arrow),new combination; Macrohyliota gracilicornis (Ar-row), new combination; Macrohyliota lucius (Pas-coe), new combination; Macrohyliota militaris(Erichson), new combination; Macrohyliota spini-collis (Gory), new combination; Macrohyliota trun-catipennis (Heller), new combination.

Megahyliota Thomas, new genusFigs. 35, 57

Type species: Uleiota feae Grouvelle, by presentdesignation and monotypy.

Figure 57. Median lobe and tegmen: Megahyliota feae (Grouvelle).


Diagnosis: The combination of scutellary striole, sixstriae, lack of a mandibular tooth in males, and“Uleiota-type” tarsi are diagnostic for the members ofthis genus.

Description: With characteristics of Brontinae:Brontini as described above, plus: Surface withincrustation. Head with frontoclypeal suture repre-sented by an abrupt change in elevation; with shortlongitudinal lines; temples present, short, angulate;basal line impressed; eyes small, convex; antennalscape longer than head; mandibles laterally expand-ed, not armed in males. Prothorax transverse; later-ally armed with denticles; anterior angles with long-er, sharper bifid teeth; posterior angles right to acute.Elytra with scutellary striole; with six punctatestriae; humeral carina present; laterally broadlyexplanate with margin denticulate. Legs short, fem-ora moderately stout; tarsi “Uleiota-type”, tarsalformula 5-5-5. Prosternal process much broader thancoxal cavity; expanded and rounded apically. Inter-coxal process of first visible abdominal sternum nar-rowly rounded. Male genitalia (Fig. 57) with longbasal strut and long flagellum present.

Distribution: Southeast Asia.

Discussion: The members of this genus greatlyresemble those of Uleiota but differ in several charac-ters to the extent that treating them as a separate

genus seems necessary. In addition to the differencesmentioned above; the alternate elytral intervals inMegahyliota are costate and densely covered withshort, erect but strongly curved setae. In Uleiota,there is a single row of setae on each interval. Thereis at least one undescribed species in this genus.

Etymology: The genus name is a combination of“mega,” the Greek word for “large”, and “hyliota,” theGreek word for “forester”. It is masculine.

Included species: Megahyliota feae (Grouvelle),new combination.

Microhyliota Thomas, new genusFigs. 36, 58

Type species: Uleiota integricollis Fairmaire, bypresent designation and monotypy.

Diagnosis: The evenly rounded, denticulate prono-tal margins, small body size, and elytral color pattern(Fig. 36) make adults of the single known member ofthis genus easily recognizable.

Description: With characteristics of Brontinae:Brontini as described above, plus: Surface withoutincrustation, distinctly pubescent. Head with fronto-clypeal suture represented by an abrupt change inelevation; with long longitudinal lines; temples present,long, rounded; basal line impressed; eyes large, con-vex; antennal scape much longer than head; mandi-bles rounded, slightly expanded laterally. Prothoraxdistinctly rounded, with obsolete anterior and posteri-or angles; laterally denticulate. Elytra with scutel-lary striole; with six punctate striae; laterally vague-ly costate. Legs moderately long, femora slender;tarsi “Dendrophagus-type”. Prosternal process a lit-tle broader than a coxal cavity; rounded and expandedapically. Male genitalia (Fig. 58) characteristic.

Distribution: Chile.

Discussion: The species upon which this genus iserected appears to be rarely collected as I have seenonly three specimens. It seems closest to the membersof Dendrophagus in its narrow, delicate body form,long frontal lines, and median lobe without a basalstrut, but differs in other respects, such as the nearlyround prothorax, long filiform antennae, and compli-cated armature of the internal sac.

Figure 58. Median lobe and tegmen: Microhyliota integricollis.


Etymology: The genus name is a combination of“micros,” the Greek word for “small”, and “hyliota,”the Greek word for “forester”. It is masculine.

Included species: Microhyliota integricollis (Fair-maire), new combination.

Parahyliota Thomas, new genusFigs. 4, 17, 37, 59

Type species: Uleiota costicollis Reitter, by presentdesignation.

Diagnosis: The absence of a scutellary striole isunique to this genus, Brontopriscus, and Uleiotawithin the Brontini. Members of Parahyliota also arecharacterized by the presence of sexual modificationsto the antennae and frons in males of some species, the“Uleiota-type” tarsi, and the reduction of the parameresin the male genitalia.

Description: With characteristics of Brontinae:Brontini as described above, plus: Surface withoutincrustation. Head without frontoclypeal suture; withshort longitudinal lines, males of some species haveelaborate modifications to frons consisting of carinae,grooves, setae-lined excavations, etc.; temples present,short; basal line impressed; eyes moderate, convex;antennal scape longer than head, modified in males ofsome species, as are the pedicel and some basalflagellomeres; mandibles laterally expanded, withdorsal tooth in males of at least one species. Prothoraxmore or less quadrate; laterally armed with denticles;anterior angles with longer, sharper teeth; posteriorangles right to acute; disk of pronotum in somespecies bordered by longitudinal carinae. Elytra with-out scutellary striole; with six punctate striae, plusrows of secondary punctures between striae; laterallycostate or carinate. Legs moderate, femora moderate-ly stout; tarsi “Uleiota-type” but some species havewhat appears to be an “asymmetrically” bilobed tar-somere III. Prosternal process much broader thancoxal cavity; rounded apically; disk of prosternum andmesosternum flat in some species and laterally cari-nate; open coxal lines on first visible abdominalsternum present in some. Male genitalia (Fig. 59)with complex armature of internal sac and parameresreduced to tooth-like structures.

Distribution: Asia, Africa

Discussion: This is the largest brontine genus, with13 described species and several undescribed. It rang-es from Africa east through Madagascar to SoutheastAsia and the Philippines. It is the only brontine genusrepresented in Africa and Madagascar. It is also theonly brontine genus exhibiting sexual modificationsto the frons and antennae. Some of the species are verydifficult to distinguish without recourse to the malegenitalia. There are a number of undescribed species.

Figure 59. Median lobe and tegmen: Parahyliota costicollis.


In the course of this study it was determined thatUleiota crenicollis Grouvelle 1913:57, described fromTaiwan, is a junior synonym of Uleiota costicollisReitter1876: 44, described from Burma, new synon-ymy.

Etymology: The genus name is a combination of“para,” the Greek word for “near”, and “hyliota,” theGreek word for “forester”. It is masculine.

Included species: Parahyliota africanus Grouvelle,new combination, western Africa; Parahyliotaalticola (Pal, Sen Gupta, and Crowson), new combi-nation, India; Parahyliota atratus (Grouvelle), newcombination, Madagascar; Parahyliota brevicollis(Arrow), new combination, Madagascar; Parahylio-ta cinamomeus (Fairmaire), new combination,Madagascar; Parahyliota costicollis (Reitter), newcombination, SE Asia; Parahyliota fallax (Grou-velle), new combination, SE Asia; Parahyliotaindicus (Arrow), new combination, India; Parahylio-ta pallidus (Arrow), new combination; Madagas-car; Parahyliota puberulus (Reitter), new combina-tion, SE Asia; Parahyliota serratus (Smith), newcombination, Philippines; Parahyliota serricollis(Candeze), new combination, Sri Lanka; Parahylio-ta siamensis (Arrow), new combination, SE Asia.

Protodendrophagus Thomas, new genusFigs. 38, 60

Type species: Protodendrophagus antipodes Tho-mas, by present designation and monotypy.

Diagnosis: The moderate-sized, loosely jointed, wing-less adults (Fig. 38) of the only known species mostclosely resemble members of Dendrophagus and Den-drophagella in general appearance, but can be distin-guished from both by the presence of a distinctfrontoclypeal suture, flat eyes, shape of pronotum,and winglessness.

Description: With characteristics of Brontinae:Brontini as described above, plus: Dorsal surfacewithout incrustation. Head with frontoclypeal suturemarked by an impunctate, transverse, curved groove,deeper medially than laterally; mandibles carinatedorso-laterally, but not expanded laterally; eyes large,about 1/3 length of head capsule, but flat; templerounded, longer than eye; basal transverse impres-sion absent; antennae rather short for tribe, attainingapical third of elytra in males, midpoint in female.

Pronotum broad, strongly rounded laterally; postero-lateral angle obsolete, so that lateral margin flowsuninterupted into basal margin; anterolateral angleabsent (male) or represented by a small, acute denti-cle located below and behind the anterior margin(female). Scutellum semicircular, medially impunc-tate and slightly tumid. Elytra evenly rounded tolateral margin, which is narrowly but sharply explan-ate, explanate margin continues unbroken anteriorlyover the humerus to base; disc with six rows ofpunctures plus a scutellary striole, punctures smalland deep; elytra not fused. Hindwings absent. Legsrather long for genus, femora slender; tarsi “Den-drophagus-type”. Prosternal process narrower than acoxal cavity, strongly rounded apically. The structureof the male genitalia (Fig. 60) is most similar to thatfound in Brontopriscus.


Figure 60. Median lobe and tegmen: Protodendrophagusantipodes.


Discussion: This genus is represented by the singlespecies described below.

Etymology: The genus name is a combination of“protos,” the Greek word for “first”, and the genusname Dendrophagus. It is masculine.

Included species: Protodendrophagus antipodesThomas, new species.

Protodendrophagus antipodes Thomas, new speciesFigs. 38, 60

Description (male): With characteristics of Brontin-ae: Brontini: Protodendrophagus as described above,plus: Length, 8.7mm. Dorsal surface dark brown,mouthparts, antennal flagellum, tarsi, and elytrapaler. Head and pronotum with scattered short, pale,recumbent pubescence; elytra with shorter, denserpubescence, almost forming a pile on the surface;punctation consisting of umbilicate punctures, small-er than an eye facet and separated by about twodiameters (low tubercles on which punctures aresituated are contiguous); punctation less dense medi-ally on head but rather uniform on pronotum. Templelong, rounded gradually to basal constriction; scapeabout as long as head. Pronotum 1.1x wider than long,anterior angle armed with low, oblique ridge lateral-ly. Anterior tarsomeres II-III dilated. Male genitaliaas in fig. 60.

Variation: Females differ from male as follows:Punctation of head and pronotum similar but simpleand much less dense medially; pubescence similar,but less dense on elytra. Temple shorter and moreabruptly rounded; antennal scape shorter than head.Pronotum more transverse, 1.3x wider than long,with anterior angle armed with a short, acute tooth.Anterior tarsomeres II-III not dilated. Length, 8.7-9.3mm.

Etymology: The species name is a noun in apposi-tion, referring to the Antipodes, a region encompass-ing Australia and New Zealand.

Types: Holotype male, in NZAC, with following labeldata: “Nelson Lakes Nat. Pk. NN Mt. Cedric 5000' O.Calvert.”/”under rock”: Allotype female, in NZAC,with following label data: “Barton Saddle Nelson,4500' 10-I-61 J.I. Townsend.”/”R.M. Bull Collection”/”[NZAC label]”.

Paratypes, all females, 1, “Iron Hill Co. 66 V.Nelson 9 Mar 68 1500m J.S. Dugdale”/”crest ridgeunder rocks”/”[NZAC label]”; 1, “Mt. Domett 1450-1600m 30 Nov. 71"/”J.S. Dugdale”/”Brontini N. gen.N.sp. Det. J.S. Watt 1983"/”[NZAC label]” (NZAC).One additional female specimen, not designated aparatype because of its poor condition: “Mt. Domett35-3000 ft NN Nov. 71"/”[NZAC label]” (FSCA).

Uleiota LatreilleFigs. 1-2, 12-14, 16, 22-24, 61-62

Uleiota Latreille 1796: 46Brontes Fabricius 1801: 97Hyliota Reitter 1880: 80Hyleota Seidlitz 1888: 55

Type species: Pal et al. (1985: 213) stated thatLatreille (1796: 46) proposed Uleiota for Cerambyxflavipes (Fabricius). This is untrue. Latreille (1796:46) listed Cerambyx and Cucujus under Uleiota, butincluded no species. Arrow’s (1901: 595) statementthat “The type of the genus is H. planata, L...” seemsto be the first designation of a type species. Arrow’sconcept of the genus was much broader than thatadopted here.

Diagnosis: The combination of broad body shape,mandibular horns in the male, lack of a scutellarystriole and five discal striae on each elytron, andUleiota-type tarsi (Fig. 16) with only four tarsomeresis diagnostic for this genus.

Description (male holotype): With characteristics ofBrontinae: Brontini as described above, plus: Surfacewith incrustation. Head with frontoclypeal suturerepresented by an abrupt change in elevation; withshort longitudinal lines; temples present, short, an-gulate; basal line impressed; eyes small, convex;antennal scape longer than head; mandibles in maleswith long, curved dorsal spine; in females laterallyexpanded. Prothorax transverse; laterally armed withdenticles; anterior angles in most with longer, sharp-er bifid or trifid teeth; posterior angles right to acute.Elytra without scutellary striole; with five punctatestriae; humeral carina present; laterally explanateand margin denticulate. Legs short, femora moder-ately stout; tarsi “Uleiota-like”, tarsal formula 4-4-4.Prosternal process much broader than coxal cavity;expanded and rounded apically. Intercoxal process offirst visible abdominal sternum narrowly rounded.


Male genitalia (Fig. 61-62) with short basal strut andflagellum present.

Distribution: Holarctic.

Discussion: Both Uleiota Latreille and BrontesFabricius have been used extensively as the genus-group name. Brontes was used more commonly in the19th Century; Uleiota was accepted by Hetschko(1930) as the valid name in his world catalog of thegroup and has been used most commonly since that

time. Latreille (1796) first used the genus-group nameUleiota and accompanied it with a brief description,but included no species. Fabricius (1801) first usedthe genus-group name Brontes, accompanied by amore detailed description and with five listed species.Under the second listed species, flavipes Fabricius(now considered a synonym of planatus Linnaeus),Fabricius (1801) listed Uleiota Latreille as a syn-onym. According to Article 12.1 of the InternationalCode of Zoological Nomenclature (ICZN 1999), genus-group names proposed before 1931 had to have had a

Figures 61-62. median lobe and tegmen: 61) Uleiota truncata; 62) Uleiota dubius.

61 62


description provided or a valid indication. Since La-treille’s 1796 use of Uleiota meets the first require-ment, Uleiota Latreille 1796 is valid and clearly haspriority over Brontes Fabricius 1801.

The genus-group name Hyliota was first used byReitter (1880) without comment and subsequently byseveral authors, most notably Arrow in his 1901review of the genus. The late George Steyskal (pers.comm.) explained that Hyliota is a more accuratetransliteration of the Greek word for “forester,” but itis an unjustified emendation.

Uleiota truncatus has been treated consistentlyas a subspecies of U. dubius. However, the structureof the male genitalia (Figs. 61-62) and pronotum (thelong bifid spines are missing in U. truncatus) sup-ports the elevation of U. truncatus to species status,and it is so treated here.

Uleiota texana Dajoz (1989: 198) is here synony-mized under U. dubius, new synonymy. AlthoughDajoz (1989: 198) claimed the unique type to be amale, there is nothing inconsistent with a female U.dubius in Dajoz' description and illustration. I havebeen unable to borrow the type from the author, butit is clear from the key to species he provided that hewas not familiar with the characters separating theNearctic species.

Included species: Uleiota algericus Maran, Uleiotaarboreus (Reitter); Uleiota debilis (LeConte); Uleiotadubius (Fabricius); Uleiota planatus (Linnaeus); Ul-eiota truncatus Motschulsky, new status.

Checklist of the described speciesof the tribe Brontini

Australodendrophagus australis (Erichson), newcombination; Australia

Australohyliota macleayi (Olliff), new combination;Australia

Australohyliota chilensis (Blanchard), new combination;Chile, Argentina

Brontoliota indivisipennis Thomas, new species, AustraliaBrontoliota intermedius Thomas, new species, AustraliaBrontoliota monteithi Thomas, new species, Australia

Brontopriscus pleuralis (Sharp); New ZealandBrontopriscus sinuatus Sharp; New Zealand

Dendrophagella capito (Pascoe), New Zealand

Dendrophagus crenatus (Paykull); PalaearcticDendrophagus cygnaei Mannerheim; NearcticDendrophagus longicornis Reitter; Japan

Macrohyliota truncatipennis (Heller), new combination;SE Asia

Macrohyliota spinicollis (Gory), new combination; SEAsia

Macrohyliota gracilicornis (Arrow), new combination; SEAsia, New Guinea

Macrohyliota bicolor Arrow, new combination; AustraliaMacrohyliota lucius (Pascoe), new combination; AustraliaMacrohyliota militaris (Erichson), new combination;

Australia

Megahyliota feae (Grouvelle), new combination; Thailand

Microhyliota integricollis (Fairmaire), new combination;Chile

Parahyliota africanus (Grouvelle), new combination,western Africa

Parahyliota alticola (Pal, Sen Gupta, and Crowson), newcombination, India

Parahyliota atratus (Grouvelle), new combination,Madagascar

Parahyliota brevicollis (Arrow), new combination,Madagascar

Parahyliota cinamomeus (Fairmaire), new combination,Madagascar

Parahyliota costicollis (Reitter), new combination, SEAsia

Parahyliota fallax (Grouvelle), new combination, SEAsia

Parahyliota indicus (Arrow), new combination, IndiaParahyliota pallidus (Arrow), new combination;

MadagascarParahyliota puberulus (Reitter), new combination, SE

AsiaParahyliota serratus (Smith), new combination,

PhilippinesParahyliota serricollis (Candeze), new combination, Sri

LankaParahyliota siamensis (Arrow), new combination, SE

Asia

Uleiota algericus Maran, AlgeriaUleiota arboreus (Reitter); Japan, SiberiaUleiota debilis (LeConte); northeastern North AmericaUleiota dubius (Fabricius); northeastern North AmericaUleiota planatus (Linnaeus); PalaearcticUleiota truncatus Motschulsky, new status; western North

America

Key to the genera of the tribe Brontiniof the world

1. Tarsi of “Dendrophagus”-type (see Fig. 15) ........ 41'. Tarsi of “Uleiota”-type (see Fig. 16-17) .............. 2


a b c

2(1'). Scutellary striole absent; body with or withoutincrustation ....................................................... 3

2'. Scutellary striole present; body with incrustation(SE Asia) ...... Megahyliota Thomas, new genus

3(2'). Body not incrusted; parameres reduced to tooth-like processes (Fig. 59) (Asia, Madagascar, andAfrica) ............ Parahyliota Thomas, new genus

3'. Body incrusted; parameres not reduced (Holarctic)..................................................Uleiota Latreille

4(1). Head with long frontal lines (Fig. 33) ................. 54'. Frontal lines, if present, short and inconspicuous

............................................................................ 7

5(4) Pronotum with anterior and posterior angles obso-lete, pronotum almost circular in outline, mar-gins strongly curved, with minute denticles (Fig.36); elytra with mottled color pattern (Fig. 36);male genitalia as in Fig. 58. (Chile) ......................................... Microhyliota Thomas, new genus

5'. Not as above ......................................................... 6

6(5') Anterior pronotal angles obsolete (Fig. 33); puncta-tion of head simple; elytra without a distinctcolor pattern (Holarctic) ......................................................................... Dendrophagus Schönherr

6'. Anterior pronotal angles with a short, broad tooth(Fig. 32); head longitudinally strigose; elytrawith a distinct color pattern (New Zealand) ....................... Dendrophagella Thomas, new genus

7(4'). Elytra fused; hind wings absent; elytral epipleuravery wide (New Zealand, Australia) ................ 8

7'. Elytra not fused; hind wings present or absent;elytral epipleura at most moderately wide .... 9

8(7). Form broad; scutellary striole absent (New Zealand)............................................ Brontopriscus Sharp

8'. Form narrow; scutellary striole present (Austra-lia) ................... Brontoliota Thomas, new genus

9(7'). Body with incrustation; anterior pronotal angleswith long tooth, margins usually with conspicu-ous teeth (Asia, Australia) ....................................................... Macrohyliota Thomas, new genus

9'. Body without incrustation; anterior pronotal an-gles with at most a short tooth; margins with orwithout conspicuous teeth ..............................10

10(9').Hind wings present; anterior pronotal angles withshort, broad tooth (Fig. 25), margins minutelydenticulate; male genitalia as in Fig. 39. (Aus-tralia) .......................................................................... Australodendrophagus Thomas, new genus

10'. Hind wings absent; anterior pronotal angle repre-sented by a low, oblique costa or a small, acutedenticle (Fig. 38); male genitalia as in Fig. 60.

(New Zealand) ................................................................. Protodendrophagus Thomas, new genus

Collection codons

Collection codons used are from Arnett et al.(1993) and are as follows:

NMPC — National Museum (Natural History), Pra-gue

NZAC — New Zealand Arthropod Collection, Auck-land

QMBA — Queensland Museum, South BrisbaneUQIC — Insect Collection, University of Queen-

sland, Saint Lucia

Acknowledgments

I thank the curators of the many collections wholoaned specimens that made this study possible.Geoffrey Monteith provided many specimens of Bron-toliota, as well as his observations on biology anddistribution. Charles Porter and the late GeorgeSteyskal offered advice on nomenclatural problems.Paul Skelley helped with the scanning electron micro-scope; Lyle Buss tutored me on the AutoMontage®system at the University of Florida Entomology andNematology Department. David G.H. Halstead, JohnL. Lawrence, and Paul E. Skelley read and criticizedthe manuscript. This is Entomology Contribution No.959, Bureau of Entomology, Nematology and PlantPathology, Florida Department of Agriculture andConsumer Services.

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31INSECTA MUNDI, Vol. 17, No. 1-2, March-June, 2003A

ppen

dix.

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QIC

4

Documents

The Brontini of the world: A generic review of the tribe