Revision of the Pepila fuscomaculata species-group and description of four new species from...

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Australian Journal of Entomology (2003) 42, 313–326 Revision of the Pepila fuscomaculata species-group and description of four new species from Australia (Coleoptera: Chrysomelidae: Alticinae) Maurizio Biondi* and Paola D’Alessandro Dipartimento di Scienze Ambientali, University of L’Aquila, 67010 Coppito (AQ), Italy. Abstract The Pepila fuscomaculata species-group from Australia, including P. fuscomaculata (Baly 1877), P. hypocrita (Blackburn 1896), P. submetallescens (Baly 1877), P. tumbyensis (Blackburn 1896), P. blackburni sp. n., P. brittoni sp. n., P. nikitini sp. n. and P. reidi sp. n., is analysed. A key to all the species is presented. Line drawings of male and female genitalia of all species are included. Brief ecological and zoogeographical data are also reported. Key words Pepila, Australia, taxonomy, new species. INTRODUCTION The Alticinae (Coleoptera: Chrysomelidae) is a subfamily composed of minute-to-medium sized beetles whose enlarged hind femora and renowned jumping ability have earned the common name of ‘flea beetles’. They are highly specialised phytophagous insects. Both the adult and larval stages feed on stems, leaves or roots, more rarely on flowers, in almost all higher plant families. The world fauna of this subfamily is represented by approximately 7–8000 species. The flea beetle genus Pepila Weise 1923, endemic to the Australian region, was described to accommodate the species Chaetocnema megalopoides Baly 1877. Recently (Biondi 2002; Biondi & D’Alessandro in press a, b, c), the taxonomic identity of this genus was redefined and 46 species are now attributed to it, included those ones described in this paper. Pepila is part of a natural group of genera (Biondi 2002) including: Chaetocnema Stephens 1831, widespread in all the zoogeographical regions, with over 300 species; Terpno- chlorus Fairmaire 1904, known from the Afrotropical (two species) and Neotropical (one species) regions; Carcharodis Weise 1910, occurring in the Afrotropical region, with at least four species; Collartaltica Bechyné 1959, occurring in the Afrotropical region, with at least three species; Biodon- tocnema Biondi 2000, known from the south-west Africa, with only one species; Seychellaltica Biondi 2002, endemic to the Seychelles, where there are at least four species (Biondi 2002). Phylogenetic relationships of Pepila and the above genera are discussed in Biondi (2002). Pepila seems to include species generally associated with well-defined ecological situations, represented mainly by different types of forests and more rarely by moist environ- ments or other habitats. Notwithstanding the absence of information for many species, it is possible to make some comments on the ecology of Pepila. This flea beetle genus is well represented in the most part of the Australian biomes, with only an apparent absence in the western desert areas (Biondi & D’Alessandro in press c). With regard to host plants, the data available for this flea beetle genus are very scarce. The plants most frequently used by the Pepila species are mainly trees and shrubs of the Myrtaceae (Eucalyptus, Melaleuca, Angophora and Cal- listemon) and Fabaceae (Acacia). However, some species are also associated with Polygonaceae (Muhlenbeckia), and Casuarinaceae (Allocasuarina and Casuarina) (Biondi & D’Alessandro in press c). It is interesting to note that Pepila species are almost exclusively associated with shrubby and arboreous plants, while the other related genera (in particular Chaetocnema) are generally associated with herbaceous plants. On the basis of the collecting dates of the material studied, the main activity period is between August to December, with a peak occurring in October corresponding to the late- spring; a subsequent increase in activity also occurs in May, during autumn. This fact could be interpreted in two ways: either that the species are bivoltine and/or that some species have an exclusive late-summer or autumn phenology. Those species that occur in the northern areas of Australia, which is characterised by a monsoon dominated tropical climate, appear to have a winter phenology (Biondi & D’Alessandro in press c). In this contribution to the field, a natural group of species related to P. fuscomaculata (Baly) is analysed. This group includes the following eight species, of which four are new to science: P. fuscomaculata (Baly 1877), P. hypocrita (Black- burn 1896), P. submetallescens (Baly 1877), P. tumbyensis (Blackburn 1896), P. blackburni sp. n., P. brittoni sp. n., P. nikitini sp. n. and P. reidi sp. n. MATERIALS AND METHODS Material examined consisted of preserved dry specimens provided to us courtesy of the following institutions: ANIC, *Author to whom correspondence should be addressed (email: [email protected]).

Transcript of Revision of the Pepila fuscomaculata species-group and description of four new species from...

Australian Journal of Entomology

(2003)

42

, 313–326

Revision of the

Pepila fuscomaculata

species-group and description of four new species from Australia (Coleoptera: Chrysomelidae: Alticinae)

Maurizio Biondi* and Paola D’Alessandro

Dipartimento di Scienze Ambientali, University of L’Aquila, 67010 Coppito (AQ), Italy.

Abstract

The

Pepila fuscomaculata

species-group from Australia, including

P. fuscomaculata

(Baly 1877),

P. hypocrita

(Blackburn 1896),

P. submetallescens

(Baly 1877),

P. tumbyensis

(Blackburn 1896),

P. blackburni

sp. n.,

P. brittoni

sp. n.,

P. nikitini

sp. n. and

P. reidi

sp. n., is analysed. A key to all thespecies is presented. Line drawings of male and female genitalia of all species are included. Briefecological and zoogeographical data are also reported.

Key words

Pepila

, Australia, taxonomy, new species.

INTRODUCTION

The Alticinae (Coleoptera: Chrysomelidae) is a subfamilycomposed of minute-to-medium sized beetles whoseenlarged hind femora and renowned jumping ability haveearned the common name of ‘flea beetles’. They are highlyspecialised phytophagous insects. Both the adult and larvalstages feed on stems, leaves or roots, more rarely on flowers,in almost all higher plant families. The world fauna of thissubfamily is represented by approximately 7–8000 species.

The flea beetle genus

Pepila

Weise 1923, endemic to theAustralian region, was described to accommodate the species

Chaetocnema megalopoides

Baly 1877. Recently (Biondi2002; Biondi & D’Alessandro in press a, b, c), the taxonomicidentity of this genus was redefined and 46 species are nowattributed to it, included those ones described in this paper.

Pepila

is part of a natural group of genera (Biondi 2002)including:

Chaetocnema

Stephens 1831, widespread in allthe zoogeographical regions, with over 300 species;

Terpno-chlorus

Fairmaire 1904, known from the Afrotropical (twospecies) and Neotropical (one species) regions;

Carcharodis

Weise 1910, occurring in the Afrotropical region, with atleast four species;

Collartaltica

Bechyné 1959, occurring inthe Afrotropical region, with at least three species;

Biodon-tocnema

Biondi 2000, known from the south-west Africa,with only one species;

Seychellaltica

Biondi 2002, endemicto the Seychelles, where there are at least four species(Biondi 2002). Phylogenetic relationships of

Pepila

and theabove genera are discussed in Biondi (2002).

Pepila

seems to include species generally associated withwell-defined ecological situations, represented mainly bydifferent types of forests and more rarely by moist environ-ments or other habitats. Notwithstanding the absence ofinformation for many species, it is possible to make somecomments on the ecology of

Pepila

. This flea beetle genus is

well represented in the most part of the Australian biomes,with only an apparent absence in the western desert areas(Biondi & D’Alessandro in press c).

With regard to host plants, the data available for this fleabeetle genus are very scarce. The plants most frequently usedby the

Pepila

species are mainly trees and shrubs of theMyrtaceae (

Eucalyptus

,

Melaleuca

,

Angophora

and

Cal-listemon

) and Fabaceae (

Acacia

). However, some speciesare also associated with Polygonaceae (

Muhlenbeckia

), andCasuarinaceae (

Allocasuarina

and

Casuarina

) (Biondi &D’Alessandro in press c). It is interesting to note that

Pepila

species are almost exclusively associated with shrubby andarboreous plants, while the other related genera (in particular

Chaetocnema

) are generally associated with herbaceousplants.

On the basis of the collecting dates of the material studied,the main activity period is between August to December,with a peak occurring in October corresponding to the late-spring; a subsequent increase in activity also occurs in May,during autumn. This fact could be interpreted in two ways:either that the species are bivoltine and/or that some specieshave an exclusive late-summer or autumn phenology. Thosespecies that occur in the northern areas of Australia, which ischaracterised by a monsoon dominated tropical climate,appear to have a winter phenology (Biondi & D’Alessandroin press c).

In this contribution to the field, a natural group of speciesrelated to

P. fuscomaculata

(Baly) is analysed. This groupincludes the following eight species, of which four are new toscience:

P. fuscomaculata

(Baly 1877),

P. hypocrita

(Black-burn 1896),

P. submetallescens

(Baly 1877),

P. tumbyensis

(Blackburn 1896),

P. blackburni

sp. n.,

P. brittoni

sp. n.,

P. nikitini

sp. n. and

P. reidi

sp. n.

MATERIALS AND METHODS

Material examined consisted of preserved dry specimensprovided to us courtesy of the following institutions: ANIC,

*Author to whom correspondence should be addressed (email:[email protected]).

314

M Biondi and P D’Alessandro

Australia National Insect Collection, Canberra, Australia;BMNH, The Natural History Museum, London, UK; SAM,South Australia Museum, Adelaide, Australia. The speci-mens were examined using WILD MZ8 and MZ12.5 binocu-lar microscopes (Wild, Heerbrugg, Switzerland). The scanningelectron micrographs were taken on a PHILIPS SEM XL30CP (Royal Philips Electronics, Eindoven, The Netherlands).

For the description of the methods used for preparinggenitalia, we refer to Döberl (1986), Duckett (1995) andKonstantinov (1998).

Abbreviations used in the text are: LAED, length ofmedian lobe of aedeagus; LAN, length of antennae; LB, totallength of body; LE, length of elytra; LP, length of pronotum;LSP, length of spermatheca; WE, width of elytra; WP, widthof pronotum.

SYSTEMATICS

Pepila

has the following diagnostic characters that allow it tobe separated from the closely related genus

Chaetocnema

Stephens 1831 (cf., Biondi 2002; Biondi & D’Alessandro inpress a): (a) hind tibiae dorsally channelled in proximal half(cf., Figs 2,3); (b) frontal sulci short or, if elongate, veryweakly and finely impressed in distal part and not clearlyvisible around upper margin of eyes (cf., Figs 4,5); (c)pronotum subrectangular or subtrapezoidal, narrower at base,and lacking bevelled anterior angles (in

Chaetocnema

ante-rior angles are generally bevelled); (d) ungual segment ofhind tarsi generally swollen (cf., Figs 2,3); (e) vertex of headalways distinctly and uniformly punctured (in

Chaetocnema

vertex is often not or only partially punctured); (f) labrumgenerally very short; (g) elytral punctation arranged inregular rows; (h) third hind tarsomere small, not or veryslightly heart-shaped.

A natural group of species related to

P. fuscomaculata

(Baly), here analysed, is characterised by the followingsynapomorphies: (1) frontal sulci elongate, finely impressedtowards ocular margin (Figs 6–13); (2) ungual segment ofhind legs distinctly swollen (Fig. 2); (3) median lobe ofaedeagus (Figs 14–19) with evident ventral sulcus, character-ised by presence of narrow longitudinal median carina [malesare unknown for

P. hypocrita

(Blackburn) and

P. reidi

sp.n.]; (4) spermatheca (Figs 20–27) with basal part proximallyclearly swollen, often with evident longitudinal torsion;(5) apical part of spermatheca shorter than collum, oftensubglobose, and with elongate appendix; (6) ductus sperma-thecae moderately or strongly elongate, generally more orless arcuate. Furthermore they also share the followingcharacter states: (7) body moderately elongate, convex(Fig. 1); (8) frontal tubercles not distinct (Figs 6–13);(9) frontal carina moderately raised, apically widely rounded;(10) eyes elliptical, moderately protuberant; (11) labrumshort; (12) pronotum clearly transverse (WP/LP > 1.50), withlateral and basal margin finely bordered, anterior angles withevident dentiform prominence, apically rounded, and poste-rior angles obtuse; (13) pronotal punctation dense and

strongly impressed; (14) elytral punctation strongly and veryregularly impressed, including sutural row; (15) humeralcallus moderately prominent; (16) macropterous wings; (17)hind tibiae dorsally widely but not deeply channelled, withlateral emargination prominent and acute (Fig. 2); (18) tibialsocket moderately wide; (19) apical spur of hind tibiae short,robust, inserted on inner side of tibial apex (Fig. 2); (20) firsthind tarsomere approximately as long as second and thirdtogether, subcylindrical, narrow.

Key to species of the

P. fuscomaculata

species-group

In view of the strong similarity among the species included inthis taxonomic group, providing a key to their identificationis not easy. At present, the only characters that allow sureidentification of these species are those related to the shapeof the spermatheca and, for the species where the male isknown, also of the median lobe of the aedeagus. In

Pepila

,the male can be generally distinguished from the female bythe wider dilation of the first anterior and middle tarsomeresand by the smaller size.

Nevertheless, in the key below, chromatic, sculptural andmorphometric characters are also considered because theycan sometimes be useful for the identification of thesespecies.1 Dorsal integument dark brown often with evident

metallic reflection. Basal part of spermatheca(Fig. 22) with longitudinal torsion; ductus nottwisted around basal part. Median lobe of aedeagus(Fig. 16) a little elongate, in ventral view slightlynarrower in middle part; apex widely rounded, withshort and apically rounded median small tooth; inlateral view curved in basal fourth and slightlysinuous in apical part ........

P. submetallescens

(Baly)– Head and pronotum reddish brown, elytra yellowish.

Basal part of spermatheca without longitudinaltorsion (Figs 20,23 and 25–27), otherwise ductuspartially twisted around basal part (Figs 21 and 24).................................................................................... 2

2(1) Basal part of spermatheca (Figs 20,23 and 25–27)without evident longitudinal torsion; ductus nottwisted around basal part ......................................... 3

– Basal part of spermatheca (Figs 21 and 24) withevident longitudinal torsion; ductus at least partiallytwisted around basal part ......................................... 7

3(2) Spermatheca (Figs 23,25 and 27) with widelyarcuate ductus ......................................................... 4

– Spermatheca (Figs 20 and 26) without or with veryweakly arcuate ductus ............................................. 6

4(3) Median lobe of aedeagus (Fig. 18) in ventral viewclearly larger in distal half; ventral sulcus wellimpressed; in lateral view strongly curved at baseand in apical third. Basal part of spermatheca(Fig. 27) subpyriform; ductus moderately arcuateshorter than length of spermatheca .......................................................................................

P. brittoni

sp. n.

Pepila fuscomaculata

species-group

315

– Median lobe of aedeagus (Figs 15 and 19) in ventralview narrower in distal half; ventral sulcus weaklyimpressed; in lateral view almost straight or sinuousonly in apical third. Basal part of spermatheca(Figs 23 and 25) subreniform; ductus strongly arcuateas long as length of spermatheca.............................. 5

5(4) Size larger (LB > 2.40 mm). Distal antennomeresslightly brownish. Scutellum reddish brown. Elytralinterstriae clearly convex. First anterior and middletarsomere distinctly dilated in male. Median lobe ofaedeagus (Fig. 15) in ventral view apically acute butwithout prominent median small tooth; in lateralview widely sinuous in apical third. Basal part ofspermatheca (Fig. 23) distally clearly bordered;apical part generally a little elongate, subglobose ...............................................

P. tumbyensis

(Blackburn)– Size smaller (LB < = 2.40 mm). Distal anten-

nomeres clearly blackened. Scutellum piceous.Elytral interstriae almost flat. First anterior andmiddle tarsomere weakly dilated in male. Medianlobe of aedeagus (Fig. 19) in ventral view apicallywith strongly prominent median small tooth; inlateral view basally straight and apically slightlycurved in ventral direction. Basal part of sperma-theca (Fig. 25) distally weakly bordered; apicalpart generally more elongate, not subglobose ...................................................................

P. nikitini

sp. n.6(3) Head and pronotum with densely impressed puncta-

tion, subrugose (Fig. 6). Elytra often with dark spotsat base and in middle. Hind femora only partiallyobscured. Spermatheca (Fig. 20) larger (LSP > =0.19 mm) with basal part strongly swollen in basalhalf; ductus basally slightly arcuate; distal part withvery elongate appendix. Median lobe of aedeagus(Fig. 14) elongate, in ventral view laterally subpar-allel; apical part lanceolate with prominent mediansmall tooth apically rounded; in lateral view straightin apical part and clearly curved in basal third ...........

............................................. P. fuscomaculata

(Baly)– Head and pronotum with sparsely impressed punc-

tation, not subrugose (Fig. 13). Elytra yellowishwithout dark spots. Hind femora blackened. Sperma-theca (Fig. 26) smaller (LSP < 0.19 mm) with basalpart elongate, weakly swollen in basal half; ductusnot arcuate, adherent to basal part; distal part withmoderately elongate appendix. Male unknown ........................................................................

P. reidi

sp. n.7(2) Head reddish brown with more densely and strongly

impressed punctation on clearly wrinkled surface(Fig. 7). Distal antennomeres slightly obscured.Scutellum and elytral suture reddish brown. Humeralcallus not obscured. Hind femora reddish brown.Basal part of spermatheca (Fig. 21) with stronglongitudinal torsion; ductus very elongate twisted atleast twice around basal part; collum little longerthan apical part. Male unknown .................................

............................................ P. hypocrita

(Blackburn)

– Head piceous with more sparsely and weaklyimpressed punctation on microreticulate surface(Fig. 10). Distal antennomeres strongly blackenedbut basally and apically yellowish. Scutellum andelytral suture piceous. Humeral callus obscured.Hind femora dorsally clearly blackened. Basal partof spermatheca (Fig. 24) with less evident longi-tudinal torsion than in

P. hypocrita

(Fig. 21); ductusshorter than in

P. hypocrita

(Fig. 21) and twistedonly partially around basal part; collum approxi-mately twice as long as apical part. Median lobe ofaedeagus (Fig. 17) a little elongate, in ventral viewslightly wider in distal half; apical part widelyrounded, with evident subtriangular median smalltooth, apically subtruncate; in lateral view almostregularly curved but in apical third with evidenthump on inner side and apically clearly bent indorsal direction ..........................

P. blackburni

sp. n.

Pepila fuscomaculata

(Baly) (Figs 1,6,14,20 and 28)

Chaetocnema fuscomaculata

Baly 1877: 174; Bryant 1923:141.

Plectroscelis fuscomaculata

(Baly): Blackburn 1896: 58, 63.

Pepila fuscomaculata

(Baly): Biondi & D’Alessandro (inpress a, c).

Type.

Western Australia.

Lectotype male (designated byM. Biondi in Biondi & D’Alessandro in press a) J. Baly Coll.(BMNH).

Other material examined.

Western Australia.

1, NovaHollandia Occid., De Boulay, J.L. Fry Coll. 1905.100, 38351(BMNH); 3, Geraldton, 28.46S 114.36E, A.M. Lea (SAM);1, Swan River, 31.45S 116.42E, A.M. Lea (SAM).

NorthernTerritory.

3, Plenty Highway, 268 km ENE of Alice Springs,22.47S 136.18E, 14.x.1978, M.S. Upton (ANIC).

Queens-land.

2, Darr River, 31 km NW by N of Longreach, 23.13S144.04E, 22.x.1975, M.S. Upton (ANIC); 1, Conardoo,Fletcher, 9 miles S of Stanthorpe, 28.46S 151.51E,20.xi.1968, E.B. Britton & W. Misko (ANIC).

South Aus-tralia.

1, Yelpawaralinna Water Hole on Warburton Creek,27.07S 138.42E, 23.xi.1993, at light, J.A. Forrest & D. Hirst(SAM); 1, Coward Springs, 29.04S 136.33E, at light,9.xi.1966, G.F. Gross (SAM); 1, Old Billa Kalina Home-stead, 29.27S 136.29E, 14.v.1981, J.A. Forrest (SAM); 11,Margaret River Overflow, 3 km W Curdimurka, 29.33S137.32E, 14.v.1981, J.A. Forrest (SAM); 1, DavenportSprings, 8 km SE Alberrie Creek, 29.37S 137.50E,13.v.1981, J.A. Forrest (SAM); 10, 4.8 km SE of Marree,29.37S 138.05E, 15.ix.1972, light trap, Z. Liepa. (ANIC); 8,same data but 21.ix.1972 (ANIC); 2, Frome River crossing ofBirdsville Track nr. Marree, 29.38S 138.05E, 25.x.1966, atlight, G.F. Gross (SAM); 1, Windy Creek, 22 km N ofBeltana, 30.35S 138.25E, 14.ix.1972, Z. Liepa (ANIC); 4,3 km NNE Yacka, 33.27S 138.31E, 18.xii.1985, on

Acacia

,C. Reid (ANIC); 2, Gawler Town, 34.36S 138.44E, J. Baly(BMNH).

New South Wales.

1, Narrabri, Valley of Namoi

316

M Biondi and P D’Alessandro

River, 31.01S 150.17E, 19–20.ix.1960, B.M. 1961–77,M.I. Nikitin (BMNH); 3, Fowlers Gap Research Station,31.05S 141.42E, 29.xi.-2.xii.1981, taken at light, J.C.Cardale (ANIC).

Description.

Male

(

n

= 10; mean and standard deviation)

:

LB, 2.27

±

0.19 mm; LAN, 1.61

±

0.08 mm; LP, 0.47

±

0.04 mm; WP, 0.80

±

0.06 mm; LE, 1.63

±

0.13 mm; WE,1.10

±

0.08 mm; LAED, 0.83

±

0.05 mm; LAN/LB, 0.71

±

0.04; WP/LP, 1.72

±

0.08; LE/LP, 3.50

±

0.18; LE/LAED,1.96

±

0.06.

Female

(

n

= 10)

:

LB, 2.43

±

0.28 mm; LAN,1.50

±

0.13 mm; LP, 0.49

±

6 0.06 mm; WP, 0.85

±

0.09 mm; LE, 1.80

±

0.20 mm; WE, 1.17

±

0.14 mm; LSP,0.20

±

0.02 mm; LAN/LB, 0.62

±

0.03; WP/LP, 1.75

±

0.04;LE/LP, 3.71

±

0.15; LE/LSP, 9.15

±

0.53.Species variable in sculpture and colour. Head and pro-

notum dark brown or reddish brown; elytra paler, often withblackish spots mainly evident near base and middle;maximum pronotal width near apical third; maximum elytralwidth near middle. Head densely and generally deeply punc-tured in interocular space; vertex proximally not puncturedbut with clearly shagreened surface; interantennal spacemore or less wide but always narrower than length of firstantennomere; labrum short, reddish brown or piceous; maxil-lary palpi generally reddish brown; antennae in male approx-imately as long as three-quarters of body length, shorter infemale; antennomeres 1–4 generally pale, 5–11 graduallybrownish; length of each antennal segment proportional tothe numerical sequences 173:92:69:87:95:110:85:89:98:94:184 (male) and 102:50:46:63:81:70:74:73:80:68:108(female) (1 = 0.002 mm). Pronotum transverse slightlynarrower at base, laterally weakly rounded; punctationdensely and strongly impressed on smooth surface. Elytralaterally weakly rounded; punctation strongly impressed on

smooth surface, with sutural row interrupted at basal fourth;apical declivous strongly punctate; interstriae generallyclearly convex. Scutellum small, half-roundish, with finelygranulate surface. Legs yellowish, but with hind femoraobscured; ungual first anterior and middle tarsomere in malelittle more dilated than in female. Ventral part with proster-num reddish, metasternum and abdomen blackened. Laststernite in both sexes very weakly punctured on finelygranulate surface; pubescence elongate, yellowish; particu-lar preapical impressions absent. Median lobe of aedeagus(Fig. 14) elongate, with almost smooth surface; in ventralview laterally subparallel; apical part lanceolate with prom-inent median small tooth apically rounded; ventral sulcuswide, clearly impressed, with longitudinal median carinaelongate; dorsal sulcus wide, interrupted at basal third;dorsal ligula wide distally subtrapezoidal; in lateral viewmedian lobe straight in apical part and clearly curved inbasal third. Spermatheca (Fig. 20) subreniform, with basalpart strongly swollen in proximal half; apical part shortwith elongate appendix; ductus short, uncoiled, weaklyarcuate.

Distribution. Western Australia, Northern Territory, SouthAustralia, New South Wales (Fig. 28).Ecology. This species seems to be associated with moistenvironments; it was collected on Acacia sp. (familyFabaceae).

Pepila hypocrita (Blackburn) (Figs 7,21 and 29)

Plectroscelis hypocrita Blackburn 1896: 58, 63.Pepila hypocrita (Blackburn): Biondi & D’Alessandro inpress a, c).

Figs 1–5. Morphologicalcharacters of the genera Pepila

Weise and ChaetocnemaStephens: (1) habitus of Pepilafuscomaculata (Baly); (2) hind

right tibia of P. brittoni sp. n.;(3) hind right tibia of C. conducta

(Motschulsky 1838); (4) distaltract of the frontal sulci of

C. conducta (4); (5) distal tractof front sulci of P. brittoni sp. n.

Pepila fuscomaculata species-group 317

Type. Western Australia. Lectotype female (designated byM. Biondi in Biondi & D’Alessandro in press a), T. Black-burn Coll., T-6094P (BMNH).Description. This description is based on the lectotype. Maleunknown. Female: LB, 1.80 mm; LAN, 1.00 mm; LP,0.40 mm; WP, 0.67 mm; LE, 1.39 mm; WE, 0.92 mm; LSP,0.17 mm; LAN/LB, 0.55; WP/LP, 1.69; LE/LP, 3.47; LE/LSP, 7.90.

Dorsal integument reddish brown, slightly paler on elytra;maximum pronotal width near basal third; maximum elytral

width near basal third. Head with little densely and moder-ately impressed punctation on clearly wrinkled surface; inter-antennal space approximately as wide as length of firstantennomere; labrum reddish brown; maxillary palpi pale;antennae in female approximately as long as half bodylength; antennomeres 1–5 pale, 6–11 gradually and veryweakly brownish; length of each antennal segment propor-tional to the numerical sequence 88:53:38:39:48:55:51:54:55:54:73 (female) (1 = 0.002 mm). Pronotum subrectangular,laterally weakly rounded; punctation strongly impressed

Figs 6–13. Head of the Pepila fuscomaculata species-group: (6) P. fuscomaculata (Baly);(7) P. hypocrita (Blackburn);(8) P. submetallescens (Baly);(9) P. tumbyensis (Blackburn);(10) P. blackburni sp. n.;(11) P. brittoni sp. n.;(12) P. nikitini sp. n.;(13) P. reidi sp. n.

318 M Biondi and P D’Alessandro

on smooth surface. Elytra laterally widely rounded; puncta-tion impressed on smooth surface, with sutural row inter-rupted at basal fourth; apical declivous clearly punctate;interstriae subconvex. Scutellum small, half-roundish, withfinely granulate surface. Legs pale but with hind femorareddish brown, dorsally clearly obscured. Ventral partyellowish with metasternum reddish brown. Last sternitevery weakly punctured on finely granulate surface; pubes-cence moderately elongate, yellowish; particular preapicalimpressions absent. Basal part of spermatheca (Fig. 21) withevident longitudinal torsion and larger in proximal half;apical part short but with elongate appendix; ductus veryelongate, at least twice twisted around basal part.Distribution. Western Australia (Fig. 29).Ecology. No information is available on the ecology of thisspecies.

Pepila submetallescens (Baly) (Figs 8,16,22 and 29)

Chaetocnema submetallescens Baly 1877: 175.Plectroscelis longior Blackburn 1896: 58–59 (synonymizedby Biondi & D’Alessandro in press a).Pepila submetallescens (Baly): Biondi & D’Alessandro inpress a, c.Types. South Australia. Lectotype male (designated by M.Biondi in Biondi & D’Alessandro in press a), Gawler Town,J. Baly Coll. (BMNH). Paralectotypes: 1 male, 1 female,same data as lectotype (BMNH).Other material examined. South Australia. 3, Port Augusta,32.29S 137.46E, iii.1998, defoliating Eucalyptus kruseana,B. Barton (SAM); 3, Maitland, 32.45S 151.35E, 13.viii.1990,CA900144, defoliating exotic eucalypts, H. Longbottom

Figs 14–19. Lateral, ventraland dorsal view of the median

lobe of the aedeagus of thePepila fuscomaculata

species-group:(14) P. fuscomaculata (Baly),

Australia, South Australia, nearMarree; (15) P. tumbyensis(Blackburn), Australia, South

Australia, Eyre’s Peninsula,near Tumby, lectotype;

(16) P. submetallescens (Baly),Australia, South Australia,

Gawler Town, lectotype;(17) P. blackburni sp. n.,

Australia, South Australia,Port Lincoln, holotype;

(18) P. brittoni sp. n., Australia,Queensland, Darr River,

holotype; (19) P. nikitini sp.n., Australia, South Australia,

Old Alton DownsHomestead, holotype.

Pepila fuscomaculata species-group 319

(SAM); 1, Port Lincoln, 34.43S 135.51E, A.M. Lea (SAM);3, same locality, T. Blackburn [paralectotypes of Plectros-celis longior Blackburn (designated by M. Biondi in Biondi& D’Alessandro in press a)] (SAM); 1, near Port Lincoln,34.43S 135.51E, T. Blackburn Coll. 1910–236, T-471 [lecto-type female of Plectroscelis longior Blackburn (designated byM. Biondi in Biondi & D’Alessandro in press a)] (BMNH); 6,Murray Bridge, 35.07S 139.15E, A.M. Lea (SAM); 1, 1.5 kmSE Kybybolite, SE Box Bulloak survey (BBG01101),36.52.46S 140.56.16E, pitfalls, 4–8.xii.1995 (SAM).

Description. Male (n = 5; mean and standard deviation): LB,2.13 ± 0.14 mm; LAN, 1.52 ± 0.09 mm; LP, 0.47 ± 0.03 mm;WP, 0.82 ± 0.05 mm; LE, 1.55 ± 0.09 mm; WE, 1.07 ±0.08 mm; LAED, 0.89 ± 0.04 mm; LAN/LB, 0.71 ± 0.01;WP/LP, 1.74 ± 0.07; LE/LP, 3.28 ± 0.09; LE/LAED,1.74 ± 0.04. Female (n = 5): LB, 2.31 ± 0.21 mm; LAN,1.44 ± 0.06 mm; LP, 0.49 ± 0.04 mm; WP, 0.86 ± 0.08 mm;LE, 1.68 ± 0.17 mm; WE, 1.14 ± 0.12 mm; LSP, 0.21 ±0.01 mm; LAN/LB, 0.63 ± 0.06; WP/LP, 1.75 ± 0.06; LE/LP, 3.44 ± 0.21; LE/LSP, 7.84 ± 0.63.

Figs 20–27. Spermatheca of the Pepila fuscomaculata species-group:(20) P. fuscomaculata (Baly), Australia, South Australia, near Marree; (21) P. hypocrita (Blackburn), Australia, Western Australia, lectotype;(22) P. submetallescens (Baly), Australia, South Australia, Port Lincoln; (23) P. tumbyensis (Blackburn), Australia, New South Wales, The Rock;(24) P. blackburni sp. n., Australia, Western Australia, Moora, paratype;(25) P. nikitini sp. n., Australia, South Australia, Margaret River, paratype; (26) P. reidi sp. n., Australia, Western Australia, Banjup, paratype;(27) P. brittoni sp. n., Australia, Queensland, near Charleville, paratype.

320 M Biondi and P D’Alessandro

Fig. 28. Distribution of thePepila fuscomaculata

species-group:(�) P. fuscomaculata (Baly)

and (●) P. tumbyensis(Blackburn).

Fig. 29. Distribution of thePepila fuscomaculata

species-group: (?) P. hypocrita(Blackburn);

(�) P. submetallescens (Baly)and (�) P. blackburni sp. n.

Pepila fuscomaculata species-group 321

Dorsal integument generally dark brown with evident giltreflection, rarely reddish brown; maximum pronotal width atapical fourth in male and at basal third in female; maximumelytral width near middle. Head sparsely and little deeplypunctured; surface from weakly to clearly wrinkled; inter-antennal space narrower than length of first antennomere;labrum reddish brown; maxillary palpi reddish brown,sometimes piceous; antennae in male approximately as longas two-thirds body length, in female little longer than halfbody length; antennomeres 1–4 generally pale, 5–11 gradu-ally and distinctly brownish; length of each antennal segmentproportional to the numerical sequence 130:60:59:54:69:76:79:76:77:77:114 (male) and 111:62:47:55:74:65:68:75:69:65:105 (female) (1 = 0.002 mm). Pronotum basally slightlynarrower, laterally weakly rounded; punctation mediallydense and strongly impressed on generally smooth surface.Elytra laterally weakly rounded; punctation impressed onsmooth surface, with sutural row interrupted at basalfourth; apical declivous weakly punctate; interstriae sub-convex. Scutellum small, half-roundish, with finely granu-late surface. Legs with tibiae and tarsi pale but with femoramore or less obscured, especially hind ones; first anteriorand middle tarsomere dilated in male, clearly narrower infemale. Ventral part generally blackened with reddishbrown pygidium. Last sternite very sparsely and weaklypunctured on slightly shagreened surface; pubescenceelongate, yellowish, absent in middle part; male withevident apical depression, absent in female. Median lobe of

aedeagus (Fig. 16) thickset, with almost smooth surface; inventral view slightly narrower in middle part; apex widelyrounded, with short and apically rounded median smalltooth; ventral sulcus wide, moderately impressed, withlongitudinal median carina evident in basal half; dorsalsulcus wide, deeply impressed, interrupted at basal third;dorsal ligula interrupted at apical fifth, distally subrectan-gular; median lobe in lateral view curved in basal fourthand slightly sinuous in apical part. Basal part of spermath-eca (Fig. 22) with evident longitudinal torsion, clearlybordered at distal margin and swollen in proximal half;apical part short with evident appendix; ductus moderatelyelongate, uncoiled but slightly sinuous.Distribution. South Australia (Fig. 29).Ecology. This species was collected on leaves of Eucalyptuskruseana (family Myrtaceae).

Pepila tumbyensis (Blackburn) (Figs 9,15,23 and 28)

Plectroscelis tumbyensis Blackburn 1896: 58, 63.Chaetocnema tumbyensis (Blackburn): Bryant 1923: 141[erroneously synonymized with Chaetocnema fuscomaculataBaly 1877].Pepila tumbyensis (Blackburn): Biondi & D’Alessandro (inpress a, c).

Fig. 30. Distribution of the Pepila fuscomaculata species-group: (�) P. brittoni sp. n., (�) P. nikitini sp. n and (�) P. reidi sp. n.

322 M Biondi and P D’Alessandro

Types. South Australia. Lectotype male (designated by M.Biondi in Biondi & D’Alessandro in press a), Eyre’s Penin-sula, near Tumby, T. Blackburn Coll., T-1185 (BMNH).Paralectotypes: 1 female, same data as lectotype (BMNH);1 female, Australia, Jacoby Coll. 1909–28a, ex T. BlackburnColl., T-5811 (BMNH); 2 females, Port Lincoln, T. Black-burn Coll., T-1885 (SAM). Western Australia. 3 females,Nova Hollandia Occid., De Boulay, J.L. Fry Coll., T-1905.100 (BMNH).Other material examined. Western Australia. 5, Dongarra,29.15S 114.55E, 13–22.viii.1935, B.M. 1935–240, R.E.Turner (BMNH); 1, same data but 23.viii–5.ix.1935(BMNH). South Australia. 1, Morgan, 34.02S 139.39E,A.M. Lea (SAM); 2, Adelaide, 34.55S 138.35E (BMNH,SAM); 3, Gawler Town, 34.36S 138.44E, J. Baly Coll. (exsyntypes of Chaetocnema submetallescens Baly 1877)(BMNH). New South Wales. 3, Tamworth, 31.05S 150.55E,A.M. Lea (SAM); 1, Kinchega NP, 32.26S 142.18E,25.ix.1985, Eucalyptus largiflorens, C. Reid (ANIC); 1,44 km NW of Narrandera, 34.28S 146.11E, 2.iv.1975,Z. Liepa (ANIC); 1, Cootamundra, 34.38S 141.01E,A.M. Lea (SAM); 6, The Rock, 35.15S 147.06E, 14.ix.1968,M.S. Upton & J. Balderson (ANIC). Victoria. 2, Birchip,35.58S 142.53E (SAM).Description. Male (n = 10; mean and standard deviation):LB, 2.45 ± 0.03 mm; LAN, 1.72 ± 0.05 mm; LP, 0.51 ±0.01 mm; WP, 0.91 ± 0.02 mm; LE, 1.83 ± 0.04 mm; WE,1.26 ± 0.03 mm; LAED, 0.95 ± 0.02 mm; LAN/LB, 0.70 ±0.02; WP/LP, 1.78 ± 0.05; LE/LP, 3.59 ± 0.07; LE/LAED,1.93 ± 0.05. Female (n = 10): LB, 2.59 ± 0.15 mm; LAN,1.62 ± 0.06 mm; LP, 0.52 ± 0.03 mm; WP, 0.94 ± 0.05 mm;LE, 1.95 ± 0.09 mm; WE, 1.33 ± 0.06 mm; LSP, 0.22 ±0.01 mm; LAN/LB, 0.63 ± 0.02; WP/LP, 1.82 ± 0.04; LE/LP, 3.78 ± 0.10; LE/LSP, 8.99 ± 0.54.

Head and pronotum generally reddish brown; elytra paleusually with more or less evident brownish spots near baseand in middle; maximum pronotal width near middle;maximum elytral width near middle. Head densely but shal-lowly punctate, with finely wrinkled surface; interantennalspace approximately as wide as length of first antennomere;labrum reddish brown; maxillary palpi pale; antennae in bothsexes little longer than half body length; antennae sometimesentirely pale, more often with antennomeres 1–4 pale and5–11 gradually and clearly brownish; antennomeres 1–4generally pale, 5–11 gradually and distinctly brownish; lengthof each antennal segment proportional to the numericalsequence 113:63:53:67:83:78:84:80:91:61:110 (male) and 122:64:59:61:78:65:82:62:70:71:108 (female) (1 = 0.002 mm).Pronotum subrectangular, laterally weakly rounded; puncta-tion strongly and densely impressed on generally finelygranulate surface. Elytra laterally weakly rounded; puncta-tion impressed on finely and sparsely punctulated surface;sutural row interrupted at basal fourth; apical declivousclearly punctate; interstriae convex. Scutellum small, half-roundish, with finely granulate surface. Legs generallyentirely pale, sometimes with obscured femora; first anteriorand middle tarsomere in male weakly dilated, narrower in

female. Ventral part generally entirely reddish brown. Laststernite very weakly punctured on finely granulate surface;pubescence moderately elongate, yellowish; particularpreapical impressions absent. Median lobe of aedeagus(Fig. 15) elongate, with almost smooth surface; in ventralview laterally subparallel, with apical part lanceolate, withrounded apex; ventral sulcus wide, weakly impressed, longi-tudinal median carina weakly raised, more evident in basalhalf; dorsal sulcus wide, clearly impressed, interrupted atbasal third; dorsal ligula well developed, distally subtrapezoi-dal; in ventral view median lobe slightly arcuate, apicallydistinctly sinuous. Spermatheca (Fig. 23) subreniform, withbasal part strongly swollen in proximal half, with distalmargin doubly bordered; apical part short with mediallyelongate appendix; ductus moderately elongate, with evidenthalf coil.Distribution. Western Australia, South Australia, NewSouth Wales, Victoria (Fig. 28).Ecology. This species was collected on leaves of Eucalyptuslargiflorens (family Myrtaceae).

Pepila blackburni sp. n. (Figs 10,17,24 and 29)

Types. South Australia. Holotype male, Port Lincoln,34.43S 135.51E, T. Blackburn (ex syntype of Plectrosceliscrassipennis Blackburn) (SAM). Paratypes. South Australia.2 males, same data as holotype (ex syntypes of Plectrosceliscrassipennis Blackburn) (SAM); 1 female, same data butA.M. Lea (SAM); 1 male, 1 female, Lucindale, 36.58S140.21E, A.M. Lea (SAM); 1 female, SE Bool Lagoon,37.08S 140.42E, 23.iv. 1979, sweeping vegetation on island,G.F.Gross (SAM). Western Australia. 3 males, 2 females,15 km W Moora, 30.36S 115.50E, 20–23.viii.1987, Mela-leuca trees, C. Reid (ANIC).Diagnosis. In the particular shape of spermatheca, with basalpart with an evident longitudinal torsion and ductus elongateand twisted around the basal part, P. blackburni sp. n. showsan evident affinity mainly with P. hypocrita (Baly). How-ever, the new species can be easily distinguished on the basisof the diagnostic characters in the key above.Description. Holotype (male). Head and pronotum reddishbrown with weak metallic reflection, elytra yellowish withclearly brownish suture, scutellum piceous. Body littleelongate-elliptical, moderately convex (LB, 1.85 mm). Maxi-mum pronotal width near basal third 0.75 mm; maximumelytral width at basal fourth 0.97 mm. Head sparsely andweakly punctured but with clearly microreticulate surface;frontal tubercles not distinct; frontal sulci elongate, finelyimpressed towards ocular margin; interantennal space nar-row, approximately as wide as half length of first antenno-mere; frontal carina weakly raised, apically widely rounded;eyes elliptical, laterally clearly protuberant; labrum veryshort, reddish brown; maxillary palpi reddish brown;antennae little longer than length of body (LAN, 1.30 mm;LB/LAN, 0.70); antennomeres 1–6 pale and 5–11 clearlyblackened but each with reddish basal and apical part; length

Pepila fuscomaculata species-group 323

of each antennal segment proportional to the numericalsequence 81:44:62:47:43:48:53:50:43:72 (1 = 0.002 mm).Pronotum clearly transverse (LP, 0.45 mm; WP/LP, 1.67),laterally weakly rounded; lateral and basal margin finelybordered; anterior angles with weak dentiform prominenceapically rounded; posterior angles slightly obtuse; punctationdensely and strongly impressed on almost smooth surface.Elytra elongate (LE, 1.38 mm; LE/LP, 3.06), entirely cover-ing pygidium, laterally widely rounded; punctation arrangedin 9 regular rows (+ 1 sutural interrupted at basal fourth);large and strongly impressed punctuation on smooth surface;apical declivous more weakly punctured; interstriae sub-convex; humeral callus weakly prominent; macropterous.Scutellum small, subtriangular, with finely granulate surface.Legs entirely pale, but hind femora reddish brown, dorsallypartially blackened; hind tibiae dorsally widely but littledeeply channelled, with clearly prominent and acute lateralemargination; hind tibial socket moderately wide; hindapical spur short, robust, reddish, inserted on inner side oftibial apex; hind first tarsomere moderately elongate, sub-cylindrical, narrow; hind ungual segment moderately swol-len; first anterior and middle tarsomere weakly dilated.Ventral part reddish, with slightly darker prosternum andpaler last two sternites. Last sternite sparsely and weaklypunctured on finely granulate surface; pubescence yellowish;particular preapical impressions absent. Median lobe ofaedeagus (Fig. 17) little elongate (LAED, 0.65 mm; LE/LAED, 2.10), with almost smooth surface; in ventral viewslightly wider in distal half; apical part widely rounded,with evident subtriangular median small tooth, apicallysubtruncate; ventral sulcus wide, more deeply impressed inapical third, with longitudinal median carina clearly visiblein basal two-thirds; dorsal sulcus evident in distal half;dorsal ligula wide, apically subtrapezoidal; median lobe inlateral view almost regularly curved but in apical third withevident hump on inner side and apically clearly bent indorsal direction.Paratypes. Male (n = 5; mean and standard deviation): LB,1.83 ± 0.04 mm; LAN, 1.26 ± 0.03 mm; LP, 0.42 ± 0.02 mm;WP, 0.74 ± 0.01 mm; LE, 1.36 ± 0.03 mm; WE, 0.96 ±0.03 mm; LAED, 0.64 ± 0.01 mm; LAN/LB, 0.69 ± 0.02;WP/LP, 1.74 ± 0.06; LE/LP, 3.20 ± 0.09; LE/LAED,2.12 ± 0.02. Female (n = 5): LB, 1.86 ± 0.22 mm; LAN,1.32 ± 0.11 mm; LP, 0.41 ± 0.04 mm; WP, 0.74 ± 0.08 mm;LE, 1.40 ± 0.15 mm; WE, 0.95 ± 0.09 mm; LSP, 0.18 ±0.01 mm; LAN/LB, 0.61 ± 0.03; WP/LP, 1.78 ± 0.06; LE/LP, 3.37 ± 0.15; LE/LSP, 7.65 ± 0.37. Paratypes in sculptureand in colour very similar to the holotype. Female with firstanterior and middle tarsomere distinctly narrower; sizesimilar to male; length of each antennal segment proportionalto the numerical sequence 66:49:30:35:37:45:44:42:51:44:62 (1 = 0.002 mm). Spermatheca (Fig. 24) with basal partclearly swollen in proximal half, with an evident longitudinaltorsion; collum elongate and distal part short with evidentappendix; ductus moderately elongate running around basalpart.Distribution. Western Australia, South Australia (Fig. 29).

Etymology. This species is named after Thomas Blackburn,its first collector, who contributed significantly to our know-ledge of beetles in the Australian Region.Ecology. This species was collected on leaves of Melaleucatrees (family Myrtaceae) and by sweeping herbaceousvegetation.

Pepila brittoni sp. n. (Figs 2,4,11,18,27 and 30)

Types. Queensland. Holotype male, 31 km NW by N ofLongreach, 23.13S 144.04E, Darr River, 22.x.1975, M.S.Upton (ANIC). Paratypes. Queensland. 9 females, same dataas holotype (ANIC); 1 male, 1 female, 31 km NW by N ofLongreach, 23.13S 144.04E, 10.v.1973, M.S. Upton (ANIC);1 male, 1 female, Nogoa River, Emerald, 23.31S 148.09E,9.v.1970, Z. Liepa (ANIC); 1 male, 1 female, 15 km S byW of Charleville, 26.32S 146.12E, 21.x.1975, M.S. Upton(ANIC); 1 female, 24 miles SW by W of Kingaroy, 26.44S151.31E, 21.xi.1968, E.B. Britton & W. Misko (ANIC). NewSouth Wales. 1 female, Narrabri, 30.20S 149.46E, 26.i.1960,B.M. 1960–370, by sweeping, around cultivated fields, M.I.Nikitin (BMNH); 1 female, same data but 15.iii.1960(BMNH); 1 female, Fowlers Gap Research Station, 31.05S141.42E, 29.xi–2.xii.1981, taken at light, J.C. Cardale(ANIC); 1 male, 27 km E of Cobar, 31.31S 146.06E,22.ix.1975, Z. Liepa (ANIC); 2 females, Bottle Bend,Murray River, 9 km S by E Trentham Cliffs, 34.16S142.18E, 13.v.1984, T.A. Weir (ANIC); 1 female, SturtHighway, 20 km SE of Trentham Cliffs, 34.24S 142.30E,13.v.1984, mallee, T.A. Weir (ANIC). Western Australia.1 male, 4 females, Millstream, Crystal Pool, 21.35S 117.04E,28.x.1970, at light, E.B. Britton (ANIC); 1 female, same databut 29.x.1970, at light, spinifex-eucalypt junction (ANIC); 1,same data but eucalypt-spinifex (ANIC); 1 female, same databut 31.x.1970, at light, eucalypt woodland (ANIC); 1 male,1 female, same data but 3.xi.1970, at light (ANIC); 1 female,13 km E by N of Newman, 23.15S 119.52E, 12.xi.1970, E.B.Britton (ANIC). Northern Territory. 1 male, Tindal, 14.31S132.22E, 1–20.xii.1967, light trap, W.J.M. Vestjens (ANIC);1 female, Delamere, 15.45.46S 131.39.5E, 6.vii.1968,M. Mendum (ANIC); 1 female, Goose Lagoon, 11 km SWby S of Borroloola, 16.10S 136.15E, 17.iv.1976, at light,J.E. Feehan (ANIC); 1 male, 2 females, McArthur River,48 km SW by S of Borroloola, 16.27S 136.05E, 29.x.1975,M.S. Upton (ANIC); 1 male, McArthur River, 80 km SW ofBorroloola, 16.39S 135.51E, 13.v.1973, M.S. Upton (ANIC);1 male, Fergusson River, 19.14S 131.50E, 7.viii.1968, M.Mendum (ANIC); 2 females, Plenty Highway, 268 km ENEof Alice Springs, 22.47S 136.18E, 14.x.1978, M.S. Upton(ANIC).Diagnosis. The shape of spermatheca places this new speciesnear P. tumbyensis (Blackburn) and P. nikitini sp. n., but onthe basis of the shape of median lobe of aedeagus it has arather isolated position within this species-group. The diag-nostic features useful for its identification are given in thekey above.

324 M Biondi and P D’Alessandro

Description. Holotype (male). Dorsal integument reddishbrown, paler on elytra, with weak metallic reflection. Bodymoderately elongate-elliptical, convex (LB, 2.10 mm).Maximum pronotal width at middle 0.80 mm; maximumelytral width at apical third 1.10 mm. Head with vertexdensely but not deeply punctured on finely shagreenedsurface; frontal tubercles not distinct; frontal sulci elongate,finely impressed towards ocular margin; interantennal spaceapproximately as wide as length of first antennomere; frontalcarina weakly raised, apically widely rounded; eyes ellipti-cal, laterally weakly protuberant; labrum and maxillary palpiyellowish; antennae approximately as long as two-thirdslength of body (LAN, 1.47 mm; LB/LAN, 0.70); antenno-meres 1–4 pale and 5–11 gradually brownish; length of eachantennal segment proportional to the numerical sequence78:45:48:50:61:46:55:54:46:51:90 (1 = 0.002 mm). Pronotumsubrectangular, clearly transverse (LP, 0.45 mm; WP/LP,1.78), laterally weakly rounded; lateral and basal marginfinely bordered; anterior angles with evident dentiformprominence apically rounded; posterior angles obtuse; punc-tation strongly and densely impressed on very finely reticu-late surface, almost smooth. Elytra elongate (LE, 1.68 mm;LE/LP, 3.72), entirely covering pygidium, laterally weaklyrounded; punctation arranged in 9 regular rows (+1 suturalinterrupted at basal third); large and strongly impressedpunctuation on smooth surface; apical declivous moreweakly punctured; interstriae subconvex; humeral callusmoderately prominent; macropterous. Scutellum small, sub-triangular, with finely granulate surface. Legs entirely pale,but hind femora dorsally partially blackened; hind tibiaedorsally widely but little deeply channeled, with stronglyprominent and acute lateral emargination; hind tibial socketmoderately wide; hind apical spur short, robust, reddish,inserted on inner side of tibial apex; hind first tarsomereelongate, subcylindrical, narrow; hind ungual segmentclearly swollen; first anterior and middle tarsomere veryweakly dilated. Ventral part entirely reddish. Last sternitevery weakly punctured on finely granulate surface; pubes-cence moderately elongate, yellowish; particular preapicalimpressions absent. Median lobe of aedeagus (Fig. 18) a littleelongate (LAED, 0.80 mm; LE/LAED, 2.10), with almostsmooth surface; in ventral view wider at distal third; apicalpart widely rounded, with moderately prominent mediansmall tooth, apically rounded; ventral sulcus wide, distinctlyimpressed, with longitudinal median carina evident in basaltwo-thirds; dorsal sulcus wide, interrupted at basal half;dorsal ligula wide, distally weakly raised but clearly mediallyincised; median lobe in lateral view apically clearly andwidely curved in ventral direction and dorsally with evidenthump at apical third.Paratypes. Male (n = 10; mean and standard deviation): LB,2.20 ± 0.20 mm; LAN, 1.54 ± 0.13 mm; LP, 0.45 ± 0.04 mm;WP, 0.81 ± 0.05 mm; LE, 1.69 ± 0.13 mm; WE, 1.13 ±0.09 mm; LAED, 0.77 ± 0.04 mm; LAN/LB, 0.70 ± 0.03;WP/LP, 1.78 ± 0.05; LE/LP, 3.72 ± 0.06; LE/LAED,2.18 ± 0.09. Female (n = 10): LB, 2.40 ± 0.30 mm; LAN,1.50 ± 0.17 mm; LP, 0.49 ± 0.05 mm; WP, 0.87 ± 0.11 mm;

LE, 1.86 ± 0.21 mm; WE, 1.23 ± 0.16 mm; LSP, 0.18 ±0.02 mm; LAN/LB, 0.63 ± 0.02; WP/LP, 1.78 ± 0.03; LE/LP, 3.81 ± 0.06; LE/LSP, 10.10 ± 0.66. Paratypes in sculp-ture and in colour very similar to the holotype. Female withfirst tarsomere of anterior and middle legs very weaklynarrower; size generally little bigger than male; length ofeach antennal segment generally proportional to numericalsequence 98:55:57:55:68:54:72:64:57:62:100 (1 = 0.002 mm).Spermatheca (Fig. 27) subpyriform, with basal part distinctlylarger in proximal half; apical part short, not globose withevident appendix; collum moderately elongate; ductus arcu-ate, not coiled.Distribution. Western Australia, Northern Territory, Queens-land, New South Wales (Fig. 30).Etymology. This species is named after Ev Britton, one of itscollectors.Ecology. Pepila brittoni was collected in different habitats,such as: Eucalyptus forest, spinifex–eucalypt interface andaround cultivated fields. No information is available on itshost plants.

Pepila nikitini sp. n. (Figs 12,19,25 and 30)

Types. South Australia. Holotype male, Old Alton DownsHomestead, 48 km SW by W of Birdsville, 26.07S 138.56E,19.ix.1972, J. Upton (ANIC). Paratypes. Northern Territory.1 male, 1 female, Chilla Well, 21.31S 130.59E, 5.xi.1968,A.O. Nicholls (ANIC). Queensland. 1 male, 9 miles N ofDayboro, 27.04S 152.46E, 23.v.1966, Z. Liepa (ANIC).South Australia. 4 males, 2 females, Old Alton Downs,Simpson Desert, 26.07S 138.56E, 19.ix.1972, Z. Liepa(ANIC); 1 female, near Lignum, Coongie Lake, 27.12S140.10E, 27–29.x.1995, pitfall, D. Hirst (SAM); 2 females,25 km N by E of Etadunna Homestead, Birdsville Track,28.40S 138.38E, 17.ix.1972, Z. Liepa (ANIC); 1 female,Margaret River Overflow, 3 km W Curdimurka, 29.33S137.32E, 14.v.1981, J.A. Forrest (SAM); 3 females, 4.8 kmSE of Marree, 29.37S 138.05E, 15.ix.1972, light trap,Z. Liepa (ANIC); 1 male, Murray Bridge, 35.08S 139.15E,A.M. Lea (SAM). New South Wales. 2 males, 2 females,Narrabri, 30.20S 149.46E, 26.i.1960, B.M. 1960–370, bysweeping, around cultivated fields, M.I. Nikitin (BMNH);1 female, same data but 27.i.1960, B.M. 1960–619 (BMNH);1 male, 1 female, 15.iii.1960, B.M. 1960–370, by sweepingaround cultivated fields (BMNH); 1 male, Fowlers GapResearch Station, 31.05S 141.42E, 29.xi-2.xii.1981, J.C.Cardale (ANIC); 1 male, same data but swept from well-watered lawn (ANIC); 1 male, 9 km NW of Wilcannia,31.30S 143.17E, 23.ix.1975, Z. Liepa (ANIC); 1 male,Kinchega NP, 32.26S 142.18E, 25.ix.1985, on Muehlen-beckia cunninghami, C. Reid (ANIC); 2 females, same databut 26.ix.1985, Acacia bivenosa flowers (ANIC); 1 male,Barham, 35.37S 144.07E, 23.iii.1960, B.M. 1960–370, bysweeping, banks of irrigation ditches, M.I. Nikitin (BMNH).Victoria. 1 male, Lake Culluleraine, W of Mildura, 34.11S142.06E, 26.xii.1966, Z. Liepa (ANIC).

Pepila fuscomaculata species-group 325

Diagnosis. This new species is closely related toP. tumbyensis, as shown by similar shape of median lobe ofaedeagus (Fig. 19) and spermatheca (Fig. 25), but it is easilydistinguishable by the characters in the key above.Description. Holotype (male). Dorsal integument withreddish brown head and pronotum with weak metallic reflec-tion; elytra yellowish. Body little elongate-elliptical, moder-ately convex (LB, 2.04 mm). Maximum pronotal width atmiddle 0.70 mm; maximum elytral width at apical third0.97 mm. Head densely but shallowly punctured; surfaceclearly wrinkled; frontal tubercles not distinct; frontal sulcielongate, finely impressed towards ocular margin; interanten-nal space little wider than length of first tarsomere; frontalcarina weakly raised, apically widely rounded; eyes ellip-tical, laterally weakly protuberant; labrum short, dark brown;maxillary palpi reddish brown; antennae approximately aslong as half body length (LAN, 1.45 mm; LB/LAN, 0.68);antennomeres 1–4 pale and 5–11 gradually brownish; lengthof each antennal segment proportional to the numericalsequence 77:39:41:40:47:50:50:55:54:52:99 (1 = 0.002 mm).Pronotum subrectangular, clearly transverse (LP, 0.40 mm;WP/LP, 1.75), laterally very weakly rounded; lateral andbasal margin finely bordered; anterior angles with evidentdentiform prominence apically rounded; posterior anglesobtuse; punctation strongly and densely impressed on veryfinely reticulate surface, almost smooth. Elytra elongate (LE,1.41 mm; LE/LP, 3.53), entirely covering pygidium, laterallyweakly rounded; punctation arranged in 9 regular rows(+1 sutural interrupted at basal fourth); large and moderatelyimpressed punctulation on smooth surface; apical declivousmore weakly punctured; interstriae almost flat, sparselypunctulated humeral callus moderately prominent; macrop-terous. Scutellum small, subtriangular, with finely granulatesurface. Legs entirely pale, but hind femora reddish browndorsally darker; hind tibiae dorsally widely but little deeplychannelled, with strongly prominent and acute lateral emar-gination; hind tibial socket moderately wide; hind apical spurshort, robust, yellowish, inserted on inner side of tibial apex;hind first tarsomere elongate, subcylindrical, narrow, approx-imately as long as second and third together; hind ungualsegment clearly swollen; first anterior and middle tarsomeredistinctly dilated. Ventral part with reddish prosternum;meso–, metasternum and abdomen blackened but withreddish last two sternites. Last sternite not punctured butfinely granulate, with sparse yellowish pubescence; particu-lar preapical impressions absent. Median lobe of aedeagus(Fig. 19) moderately elongate (LAED, 0.75 mm; LE/LAED,1.90), with almost smooth surface; in ventral view a littlewider in middle part; apical part rounded, with very promi-nent median small tooth; ventral sulcus wide, distinctlyimpressed, with longitudinal median carina evident in basalfourth; dorsal sulcus wide, visible in apical half; dorsal ligulawide, distally subtrapezoidal; median lobe in lateral viewalmost straight with apical part slightly bent in ventraldirection.Paratypes (mean and standard deviation). Male (n = 10): LB,1.95 ± 0.12 mm; LAN, 1.34 ± 0.10 mm; LP, 0.42 ± 0.03 mm;

WP, 0.71 ± 0.05 mm; LE, 1.46 ± 0.11 mm; WE, 0.99 ±0.06 mm; LAED, 0.78 ± 0.03 mm; LAN/LB, 0.68 ± 0.04;WP/LP, 1.70 ± 0.04; LE/LP, 3.50 ± 0.07; LE/LAED,1.88 ± 0.07. Female (n = 10): LB, 2.11 ± 0.19 mm; LAN,1.35 ± 0.10 mm; LP, 0.44 ± 0.04 mm; WP, 0.76 ± 0.07 mm;LE, 1.62 ± 0.15 mm; WE, 1.08 ± 0.10 mm; LSP, 0.19 ±0.01 mm; LAN/LB, 0.64 ± 0.03; WP/LP, 1.72 ± 0.04; LE/LP, 3.64 ± 0.10; LE/LSP, 8.28 ± 0.75. Paratypes in sculptureand in colour very similar to the holotype. Female with firstanterior and middle tarsomere slightly narrower; size littlebigger than male; length of each antennal segment propor-tional to the numerical sequence 85:50:37:41:47:52:51:53:48:44:73 (1 = 0.002 mm). Spermatheca (Fig. 25) subreni-form, with basal part strongly swollen in proximal half;apical part generally short but not globose, with elongateappendix; ductus elongate, widely arcuate.Distribution. Northern Territory, Queensland, South Aus-tralia, New South Wales, Victoria (Fig. 30).Etymology. This species is named after M.I. Nikitin, its firstcollector.Ecology. This species seems to be associated with moisthabitats and was collected on Muehlenbeckia cunninghami(family Polygonaceae) and Acacia bivenosa (familyFabaceae).

Pepila reidi sp. n. (Figs 13,26 and 30)

Types. Western Australia. Holotype female, 15 km WMoora, 30.36S 115.50E, 20–23.viii.1987, Melaleuca trees,C. Reid (ANIC). Paratypes. 1 female, 15 km W Moora,30.36S 115.50E, 20–23.viii.1987, Melaleuca trees, C. Reid(ANIC); 2 females, Nova Hollandia Occid., De Boulay, J. L.Fry Coll. 1905.100 (BMNH); 1 female, Banjup S Perth,Harper Road, small lake, 32.07S 115.53, 10.i.1986, on Euca-lyptus rudis, C. Reid (ANIC).Diagnosis. Pepila reidi sp. n. is related to P. fuscomaculata(Baly), but the particular shape of spermatheca, characterisedby an elongate basal part and the short ductus adherent tobasal part, places it in an isolated position within thistaxonomic group and allows easy identification (see keyabove).Description. Holotype (female). Dorsal integument reddishbrown with head and pronotum dark reddish brown, clearlypaler on elytra and scutellum piceous. Body moderatelyelongate-elliptical, moderately convex (LB, 2.18 mm).Maximum pronotal width at middle 0.80 mm; maximumelytral width near middle 1.12 mm. Head sparsely andweakly punctured; surface finely shagreened; frontal tuber-cles not distinct; frontal sulci elongate, finely but clearlyimpressed towards ocular margin; interantennal spaceapproximately as wide as half length of first antennomere;frontal carina weakly raised, apically widely rounded; eyeselliptical, laterally moderately protuberant; labrum short,yellowish; maxillary palpi reddish; antennae little longer thanhalf body length (LAN, 1.35 mm; LB/LAN, 0.62); antenno-meres 1–4 pale and 5–11 gradually brownish; length of eachantennal segment proportional to the numerical sequence

326 M Biondi and P D’Alessandro

90:41:37:38:48:37:45:53:49:50:73 (1 = 0.002 mm). Pronotumsubrectangular, clearly transverse (LP, 0.51 mm; WP/LP,1.56), laterally very weakly rounded; lateral and basal marginfinely bordered and blackened; anterior angles with evidentdentiform prominence apically rounded; posterior anglesobtuse; punctation strongly and densely impressed on smoothsurface. Elytra elongate (LE, 1.63 mm; LE/LP, 3.17), entirelycovering pygidium, laterally weakly rounded; punctationarranged in 9 regular rows (+ 1 sutural interrupted at basalfourth); large and strongly impressed punctures on smoothand sparsely punctulated surface; apical declivous withstrongly impressed punctation; interstriae convex; humeralcallus moderately prominent; macropterous. Scutellumsmall, half-roundish, with finely granulate surface. Legsentirely pale, but hind femora blackened and hind tibiaereddish; hind tibiae dorsally widely but little deeply chan-neled, with strongly prominent and acute lateral emargina-tion; hind tibial socket moderately wide; hind apical spurshort, robust, reddish, inserted on inner side of tibial apex;hind first tarsomere elongate, subcylindrical, narrow; hindungual segment clearly swollen; first anterior and middletarsomere very weakly dilated. Ventral part with reddishbrown prosternum and mesosternum; metasternum andabdomen strongly blackened but last sternite slightly paler.Last sternite sparsely and weakly punctate on almost smoothsurface; pubescence yellowish; particular preapical impres-sions absent. Spermatheca (Fig. 26) (LSP, 0.19 mm; LE/LSP,8.70) subreniform, with basal part elongate, swollen inproximal half; apical part with elongate appendix; collummoderately elongate; ductus adherent to basal part.Paratypes. Female (n = 4; mean and standard deviation): LB,2.00 ± 0.14 mm; LAN, 1.24 ± 0.09 mm; LP, 0.45 ± 0.05 mm;WP, 0.73 ± 0.06 mm; LE, 1.50 ± 0.11 mm; WE, 1.03 ±0.09 mm; LSP, 0.18 ± 0.01 mm; LAN/LB, 0.62 ± 0.02; WP/LP, 1.62 ± 0.04; LE/LP, 3.31 ± 0.09; LE/LSP, 8.40 ± 0.46.Male unknown.Distribution. Western Australia (Fig. 30).Etymology. This species is named after Chris Reid, distin-guished specialist of leaf beetles who collected most of thespecimens.Ecology. This species was collected on trees of Melaleucasp. and Eucalyptus rudis (family Myrtaceae).

ACKNOWLEDGEMENTS

We are very grateful to colleagues who enabled us to studythe valuable material preserved in their respective institu-tions: E. Matthews (SAM), S.L. Shute (BMNH) and T.A.Weir (ANIC).

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Accepted for publication 4 April 2003.