Australian Weevils (Coleoptera: Curculionoidea) II: Brentidae, Eurhynchidae, Apionidae and a Chapter on Immature Stages by Brenda May

By EC Zimmerman

This volume covers the remainder of the primitive weevils (Division Orthoceri), namely the families Brentidae, Eurhynchidae and Apionidae. It catalogues 43 genera and 173 species and features almost 2000 individual drawings and black-and-white photographs. These illustrations are augmented by 270 full-colour habitus photographs in Volumes V and VI. The volume also includes an important chapter on the Immature Stages of Australian Curculionoidea by Brenda May, New Zealand, which describes the larval and pupal stages of 158 species of Australian weevils and features overviews of larval characters and their nomenclature as well as of rearing and preservation techniques applicable to weevil larvae. More than half the drawings in the volume accompany this chapter.

• Language: English
• Publisher: CSIRO PUBLISHING
• Release date: Jun 1, 1994
• ISBN: 9780643105614

Book preview

CATALOGUE OF THE AUSTRALIAN

CURCULIONOIDEA

ILLUSTRATED AND ANNOTATED

CONTINUED

Division I

ORTHOCERI

Continued from Volume I

BRENTIDAE, EURHYNCHIDAE

AND APIONIDAE

Conspectus of the Families, Subfamilies, Tribes, Subtribes and Genera Treated in this Volume

Where letters follow numbers in the lists (136A, for example), they indicate that the names were added after the manuscript was completed. Future additions may be made in the same way.

7. BRENTIDAE, page 15

BRENTINAE, page 35

BRENTINI, page 35

AMORPHOCEPHALINA, page 37

103. Ankleineella Zimmerman, page 43

104. Cordus Schoenherr, page 53

105. Kleineella Strand, page 110

ARRHENODINA, page 115

106. Baryrhynchus Lacordaire, page 117

107. Caenorhychodes Kleine, page 121

108. Ectocnemus Pascoe, page 122

109. Schizoeupsalis Kleine, page 124

CYPHAGOGINI, page 127

CYPHAGOGINA, page 130

110. Allaeometrus Senna, page 132

111. Ancylobrentus Damoiseau, page 133

112. Callipareius Senna, page 133

113. Catagogus Kleine, page 135

114. Cyphagogus Parry, page 138

115. Isomorphus Kleine, page 147

116. Mesoderes Senna, page 148

117. Neosebus Senna, page 152

HOPLOPISTHIINA, page 153

118. Carcinopisthius Kolbe, page 153

STEREODERMINA, page 155

119. Cerobates Schoenherr, page 155

TRACHELIZINI, page 166

CEOCEPHALINA, page 167

120. Hormocerus Schoenherr, page 167

121. Uropteroides Kleine, page 177

ITHYSTENINA, page 178

122. Ithystenus Pascoe, page 178

123. Mesetia Blackburn, page 181

MICROTRACHELIZA, page 182

124. Arairorrhinus Senna, page 184

125. Higonodes Zimmerman, page 184

126. Microtrachelizus Senna, page 188

TRACHELIZINA, page 196

127. Euschizus Kleine, page 197

128. Miolispa Pascoe, page 201

129. Stenobrentus Damoiseau, page 207

130. Trachelizus Schoenherr, page 209

131. Tracheloschizus Damoiseau, page 210

CYLADINAE, page 216

132. Cylas Latreille, page 222

8. EURHYNCHIDAE, page 237

133. Aporhina Boisduval, page 241

134. Ctenaphides Pascoe, page 246

135. Eurhynchus Schoenherr, page 255

9. APIONIDAE, page 273

APIONINAE, page 281

APIONINI, page 282

136. Apion Herbst, page 282

136A. Coelocephalapion Wagner, page 312

137. Exapion Bedel, page 313

APLEMONINI, page 314

138. Perapion Wagner, page 315

NOTAPIONINI, page 317

139. Kelainapion Zimmerman, page 320

140. Leaoapion Zimmerman, page 325

141. Lissapion Zimmerman, page 328

142. Notapion Zimmerman, page 330

143. New genus A, page 335

RHINORHYNCHIDIINAE, page 337

144. Rhinorhynchidius Voss, page 337

MYRMACICELINAE, page 349

145. Myrmacicelus Chevrolat, page 353

Checklist of the Genera and Species

Treated in this Volume

(Qld.= Queensland, N.S.W.= New South Wales, Vic.= Victoria, Tas.= Tasmania, S.A.= South Australia, N.T.= Northern Territory; W.A.= Western Australia)

Order COLEOPTERA Aristotle, continued

Superfamily CURCULIONOIDEA Latreille, continued

Division I ORTHOCERI Schoenherr, continued

Family BRENTIDAE Schoenherr

Subfamily BRENTINAE Schoenherr

Tribe BRENTINI Schoenherr

Subtribe AMORPHOCEPHALINA Power

Genus ANKLEINEELLA Zimmerman

401. australis (Lacordaire)

Qld., N.S.W., N.T., W.A.

402. mniszechi (Power)

Qld.

403. subscripta (Damoiseau)

Qld., N.S.W.

404. sulcicollis (Pascoe)

Qld., N.S.W., Vic., S.A., N.T., W.A.

Genus CORDUS Schoenherr

405. acutipennis Power

Qld.

406. demarzi Damoiseau

Qld., N.T., W.A.

407. festus Damoiseau

S.A., W.A.

408. firmus Damoiseau

N.S.W., S.A., W.A.

409. ganglebaurei Senna

Qld., N.S.W., Vic., S.A., W.A.

410. hospes Schoenherr

Qld., N.S.W., Vic., Tas., S.A., N.T., W.A.

411. laevis Damoiseau

W.A.

412. occidentalis Damoiseau

W.A.

413. pascoei Power

Qld., N.S.W., Vic., S.A., W.A.

41e. plumipennis Damoiseau

N.T.

415. queenslandicus Senna

Qld., N.T.

416. recticornis Damoiseau

W.A.

417. rostralis Damoiseau

Qld.

418. schoenherri Power

Qld., N.S.W., Vic., S.A., N.T., W.A.

419. ultimus Damoiseau

Qld.

420. vermiculatus Damoiseau

Qld., N.T.

421. vicinus Senna

N.T., W.A.

422. new species 1

Qld., W.A.

423. new species 2

Qld.

424. new species 3

Qld.

425. new species 4

S.A.

426. new species 5

Qld.

427. new species 6

Qld.

427A. new species 7

Vic.

Genus KLEINEELLA Strand

428. barbata (Kleine)

Qld.

Subtribe ARRHENODINA Lacordaire

Genus BARYRHYNCHUS Lacordaire

Subgenus EUPSALOMINUS Kleine

429. lineicollis Power

Qld., widely distributed in Indonesia.

Genus CAENORHYCHODES Kleine

430. digramma (Boisduval)

Qld., N.S.W., India to Australia

Genus ECTOCEMUS Power

431. decemmaculatus (Montrouzier)

Qld., Indo- and Austromalaya

Genus SCHIZOEUPS Kleine

432. promissa (Pascoe)

Qld., N.S.W., Austromalaya

Tribe CYPHAGOGINI Kolbe

Subtribe CYPHAGOGINA Kolbe

Genus ALLAEOMETRUS Senna

433. breviceps Senna

Qld., India to Australia

Genus ANCYLOBRENTUS Damoiseau

434. australicola Damoiseau

Qld. (introduced?)

Genus CALLIPAREIUS Senna

435. planitarsus (Perroud and Montrouzier)

Qld., New Guinea, New Hebrides, New Caledonia

Genus CATAGOGUS Kleine

436. diorymerus (Lea)

Qld., N.S.W. (introduced?)

Genus CYPHAGOGUS Parry

437. bipunctatus Senna

Qld., N.S.W., Vic., Tas., New Guinea

438. crassitarsus Damoiseau

Qld. (introduced?)

439. delicatus Lea

Qld., N.S.W., Vic., Tas., New Guinea

440. modiglianii Senna

Qld., Indo- and Austromalaya

441. nigraustralis Damoiseau

Qld., New Guinea

442. odewahni Pascoe

Qld., Indomalaya

443. tristriatus Kleine

Qld., New Guinea?

Genus ISOMORPHUS Kleine

444. species 1

Qld. (introduced?)

445. species 2

N.T. (introduced?)

Genus MESODERES Senna

446. guttatus (Kleine)

Qld., Java to New Guinea

447. maculatus Senna

Qld., Indonesia

448. sexnotatus Senna

Qld., New Guinea

Genus NEOSEBUS Senna

449. Queensland species 1

Qld. (introduced?)

Subtribe HOPLOPISTHIINA Senna and Calabresi

Genus CARCINOPISTHIUS Kolbe

450. kolbei Senna

Qld., Indomalaya to New Guinea

Subtribe STEREODERMINA Sharp

Genus CEROBATES Schoenherr

Subgenus CEROBATES Schoenherr

451. laevipennis Senna

Qld., India to the Solomon Islands

452. sexsulcatus Motschulsky

Qld., Ceylon to New Guinea

453. tristriatus (Fabricius)

Qld., Ceylon to New Guinea

Subgenus IONTHOCERUS Lacordaire

454. ophthalmicus (Pascoe)

Qld., Lord Howe Island, Sumatra?

Tribe TRACHELIZINI Lacordaire

Subtribe CEOCEPHALINA Lacordaire

Genus HORMOCERUS Schoenherr

455. compressitarsis (Senna)

Qld., Indo- and Austromalaya

456. fossulatus Blackburn

Qld. (introduced?)

457. reticulatus (Fabricius)

Qld., Indo- and Austromalaya

Genus UROPTEROIDES Kleine

458. gestroi (Senna)

Qld., Indo- and Austromalaya

Subtribe ITHYSTENINA Lacordaire

Genus ITHYSTENUS Pascoe

459. hollandiae (Boisduval)

Qld., New Guinea and neighbouring islands

Genus MESETIA Blackburn

460. amoena Blackburn

N.S.W. (introduced?)

Subtribe MICROTRACHELIZINA Zimmerman

Genus ARAIORRHINUS Senna

461. howitii (Pascoe)

N.S.W., Vic., Lord Howe Island, Norfolk Island

Genus HIGONODES Zimmerman

462. novenarius (Damoiseau)

Qld., Aru Islands

Genus MICROTRACHELIZUS Senna

463. laevis Damoiseau

Qld. (introduced?)

464. species 2

Qld. (introduced?)

465. species 3

Qld. (introduced?)

466. species 4

Qld. (introduced?)

467. species 5

Qld. (introduced?)

468. species 6

Qld. (introduced?)

469. species 7

N.S.W. (introduced?)

Subtribe TRACHELIZINA Lacordaire

Genus EUSCHIZUS Kleine

470. alarius (Kleine)

Qld., N.S.W. (introduced)

471. dictatorius Kleine

N.S.W. (introduced)

472. internatus (Pascoe)

Qld., N.S.W., Norfolk Island (introduced)

Genus MIOLISPA Pascoe

473. australiana Senna

Qld., N.S.W., New Guinea?

474. novaguineensis (Guerin-Meneville)

Qld., Indonesia

Genus STENOBRENTUS Damoiseau

475. lineatus Damoiseau

Qld. (introduced?)

Genus TRACHELIZUS Schoenherr

476. bisulcatus (Fabricius)

Qld., Oriental Region to Melanesia

Genus TRACHELOSCHIZUS Damoiseau

477. altilis (Kleine)

Qld., New Guinea, Solomon Islands

478. angulaticeps (Senna)

Qld., Indo- and Austromalaya

479. dichrous (Lacordaire)

Qld., N.S.W. (introduced?)

480. species 4

Qld. (introduced?)

481. species 5

Qld. (introduced?)

482. species 6

Qld. (introduced?)

483. species 7

Qld. (introduced?)

Queensland Genus A

484. species 1

Qld. (introduced?)

Queensland Genus B

485. species 1

Qld. (introduced?)

Queensland Genus C

486. species 1

Qld. (introduced?)

Lord Howe Island Genus A

487. species 1

Lord Howe Island (introduced?)

Subfamily CYLADINAE Schoenherr

Genus CYLAS Latreille

488. formicarius (Fabricius)

Qld., N.S.W., N.T., Indomalaya and now Pan Tropical

Family EURHYNCHIDAE Lacordaire

Genus APORHINA Boisduval

489. australis Heller

Qld., Banks Island, Moa Island

490. splendida (Blackburn)

Qld.

Genus CTENAPHIDES Pascoe

491. maculatus (Pascoe)

W.A.

492. porcellus Pascoe

W.A.

493. new species 1

W.A.

Genus EURHYNCHUS Kirby

494. acanthopterus Boisduval

Qld., N.S.W., Vic., Tas.

495. laevior (Kirby)

N.S.W.

496. quadridens Erichson

Qld., N.S.W., Vic., Tas., S.A.

497. quadrituberculatus Boheman

Qld., N.S.W., Tas.

498. scabrior (Kirby)

N.S.W.

Family APIONIDAE Schoenherr

Subfamily APIONINAE Schoenherr

Tribe APIONINI Schoenherr

Genus APION Herbst

499. aemulum Lea

Qld., N.S.W., Vic., S.A., N.T.

500. amabile Lea

N.S.W.

501. argutulum Pascoe

Qld., N.S.W., N.T.?, W.A.?

502. astri Lea

S.A.

503. basiinflatum Lea

Qld.

504. binotatum Lea

W.A.

505. carpophagum Lea

W.A.

506. comosum Pascoe

W.A.

507. condensatum Lea

N.S.W.

508. congestum Lea

Qld., N.S.W.

509. congruum Lea

W.A.

510. convexipenne Lea

Qld., W.A., Cornwallis, Mabuiag and Thursday Islands

511. foveicolle Lea

W.A.

512. fuscosuturale Lea

W.A.

513. hoblerae Lea

Qld.

514. immundum Lea

N.S.W.

515. integricolle Lea

W.A.

516. macleayense Lea

Qld., N.S.W.

517. melvillense Lea

N.T.

518. microscopicum Lea

N.S.W., Vic., Tas.

519. nigrosuturale Lea

W.A., N.T.

520. nigroterminale Lea

N.S.W.

521. niveodispersum Lea

Qld.

522. parvocastaneum Lea

S.A.

523. philanthum Lea

W.A.

524. pictipes Lea

Qld.

525. pilistriatum Lea

Qld.

526. pudicum Lea

W.A.

527. pulicare Pascoe

W.A.

528. quadricolor Lea

Qld., N.T.

529. rivulare Lea

N.T.

530. solani Lea

N.S.W.

531. subopacum Lea

Tas.

532. tasmanicum Lea

Tas.

533. teretirostre Lea

W.A.

534. terraereginae Blackburn

Qld., N.S.W., N.T., W.A.

535. turbidum Lea

S.A.

536. vertebrale Lea

N.S.W., W.A.

537. new species 1

Qld.

538. new species 2

A.C.T.

539. new species 3

S.A.

540. new species 4

N.S.W., N.T.

541. new species 5

W.A.

542. new species 6

W.A.

543. new species 7

Qld.

544. new species 8

W.A.

545. new species 9

W.A.

546. new species 10

Qld.

547. new species 11

Qld.

548. new species 12

Qld.

Genus COELOCEPHALAPION Wagner

548A. aculeatum (Wagner)

Qld. (introduced from Mexico)

Genus EXAPION Bedel

549. ulicis (Forster)

N.S.W., Vic., Tas., S.A. (introduced from Europe)

Tribe APLEMONINI Kissinger

Genus PERAPION Wagner

550. antiquum (Gyllenhal)

Qld., N.S.W., Vic., S.A., W.A. (introduced from South Africa)

Tribe NOTAPIONINI Zimmerman

Genus KELAINAPION Zimmerman

551. illawarense (Lea)

N.S.W.

552. inflaticolle (Lea)

Qld.

553. stilbum (Lea)

Qld., N.S.W.

Genus LEAOAPION Zimmerman

554. agonis (Lea)

W.A., N.S.W., W.A.

Genus LISSAPION Zimmerman

555. albertisii (Pascoe)

Qld.

556. sculpticeps (Lea)

Qld.

557. varistriatum (Lea)

Qld., N.S.W.

Genus NOTAPION Zimmerman

Subgenus NOTAPION Zimmerman

558. mediopunctum (Lea)

Qld.

559. striatipenne (Lea)

N.S.W.

560. tenuistriatum (Lea)

N.S.W.

561. trilobicolle (Lea)

Qld.

562. varirostre (Lea)

Qld.

Subgenus GANOAPION Zimmerman

563. clavicorne (Lea)

Qld.

564. mednonstriatum (Lea)

Qld.

New Genus A

565. new species 1

Lord Howe Island

Subfamily RHINORHYNCHIDIINAE Zimmerman

Genus RHINORHYNCHIDIUS Voss

566. australasiae (Lea)

W.A.

567. meridionalis (Lea)

S.A.

568. new species 1

S.A.

569. new species 2

Tas.

570. new species 3

N.S.W.

Subfamily MYRMACICELINAE Zimmerman

Genus MYRMACICELUS Chevrolat

571. exsertus Pascoe

S.A., W.A.

572. formicarius Chevrolat

Qld., N.S.W.

573. puerulus Lea

W.A.

Note: Throughout the text SL means standard length (the length as seen in side view from apex of pronotum to apex of elytra), and TL means total length (the length in side view from apex of rostrum, whatever its position if it is in advance of the head, or from apex of head if that is farther in front, to apex of elytra as the specimen is mounted). On the illustrations, the sizes are usually given as total lengths as the specimens are viewed. See also the note on page 34 of Volume I.

7

Family

BRENTIDAE

Schoenherr

Brenthides Schoenherr, 1823:1137; 1826:7, 68; 1840:465; 1845:326. Westwood, 1836:334 (see for early literature listed). Labram and Imhoff, 1838, Figures (no page or figure numbers). Imhoff, 1856. Lacordaire, 1866:398 (classification of genera of the world). Lameere, 1900:369 (untenable conclusions regarding relationships).

Brenthidae, Gemminger and Harold, 1872:2703 (world list). Masters, 1872:269, 1886:508 (Australian lists). LeConte and Horn, 1876:323; 1883:531 (North American fauna and classification). Sharp, 1895:11 (Biologia Centrali Americana). Schoenfeldt, 1908:1 (Genera Insectorum, edition I); 1910:1 (Coleopterorum Catalogus, edition I). Schaeffer, 1915:52 (key to North American fauna). Bl tchley and Leng, 1916:18 (Northeastern American fauna). Kleine, 1921e:109 (myrmecophily); 1921f:38 (geographical distribution; faulty, consult with caution); 1922d (new taxonomy); 1926a: 1 (Indian catalogue); 1926b (Philippine fauna); 1926d (Indonesian fauna, with keys); 1927 (Coleopterorum Catalogus, edition II); 1927 (Genera Insectorum, edition II); 1928d:63 (locations of type specimens); 1930:195 (bibliography of works on Brentidae); 1931:149 (biology); 1938 (Genera Insectorum edition III); see main bibliography for a listing of many other works by Kleine. Beeson, 1925:98 (larvae). Tillyard, 1926:240 (Australia and New Zealand faunas). Böving and Craighead, 1931:15, 66 (larvae). Gardner, 1935:139 (larvae). Hoffmann, 1945:161 (French fauna: 1 species). Jeannel, 1949:974 (general discussion). Crowson, 1955:163 (higher classification). Haedo Rossi, 1961 (detailed account of Argentine fauna). Morimoto, 1962a:369, 370 (classification and morphology); 1962b:31 (classification and Japanese fauna); 1962c:177 (catalogue of Japanese fauna); see also under Brentidae, below. W. Schedl, 1970:97 (western Palaearctic fauna). Britton, 1970:518, 615 (Australian fauna). Kasap and Crowson, 1977:35 (comparative anatomy). Calder, 1989:1210 (digestive tract and nervous system); 1990:457, 469 (male and female reproductive systems).

Brentides, Latreille, 1825:389; 1828:589.

Brentidae, Blackwelder, 1947:771 (Neotropical catalogue). De Muizon, 1960 (detailed account of African fauna). Morimoto, 1967a:267 (classification and revision of Japanese fauna); 1976b:469 (revision of family taxonomy). Damoiseau, 1963a (nomenclature, revised taxonomy; 1967, major African faunal monograph); see listing of many other titles by Damoiseau in main bibliography in Volume III.

Brentoidea, Pierce, 1916:462.

The correct spelling of the family name is Brentidae. The type-genus is Brentus Fabricius, 1787:95; 1792:491; 1798:174; 1801:545. Illiger, 1805:161. Germar, 1817:340; 1824:XIV, 189. Schoenherr, 1823:1167, altered the spelling to Brenthus, an unjustified emendation, and he used the spelling Brenthides for the family-group name. Most of the early authors followed Schoenherr’s usage, but A family-group name based upon an unjustified emendation of a generic name is an unjustified original spelling and must be corrected …. (International Code 3Sd). It is illogical and confusing to have a family-group name that is not based upon the correct original spelling of its type-genus.

Some authors have adopted the mistaken practice of considering the first author who uses a family-group suffix (-ina, -ini, -inae, -idae) as the author of that name. Such usage could lead to as many authors’ names being used in a group as there are subdivisions above the genus. The Principle of Coordination applies. (A name established for a taxon at any rank in the family group is deemed to be simultaneously established with the same author and date for taxa based upon the same name-bearing type …. Article 36 of the Code.)

This is the first review of the Australian Brentidae as a whole. The family has been mostly ignored by Australian workers.

Distribution and Bionomics

Brentidae are almost unknown as fossils, and none is known fossil in Australia. Kleine, 1941:41, described a species of Xestocoryphus, which has several living African species, from African copal of undetermined age. No ancient fossil brentids have been found.

Brentidae are mostly tropical insects. Their greatest development is in the Indo-Pacific where there are more than 150 genera. New Guinea has a rich but poorly known fauna, and endemic species are found in diminishing numbers eastward to Fiji.

I consider the few species described by Kleine from Tahiti and the Marquesas to be introductions. Kleine, 1928a described Cyphagogus samoanus and Chalybdicus reverens from Samoa, and he gave a nonsense explanation for the reasons for their being in Samoa. Buxton, in an editor’s footnote to that paper was right to comment unfavourably upon Kleine’s remarks. I suggest that the names apply to introduced species. The Cyphagogus may be an introduced western Pacific species (probably modiglianii Senna), several of which are widely distributed, and the Chalybdicus may be a synonym of hahnei Kleine of the New Hebrides. Kleine, 1944:150, described Opisthenoxys samoanus from one specimen found in the old Fairmaire collection in the National Museum in Paris. This specimen is labelled as having come from Samoa, but I suspect that the species is not Samoan. It was incorrectly assigned to Opisthenoxys, and Damoiseau, 1964a:415, described Nesidiobrentus as a new genus to receive it.

Figures 1, 2 Outline sketches of dorsal and ventral sides of the brentid Ectocnemus decimmaculatus (Montrouzier), male, to show features used in taxonomy. The prothorax is bent downward to expose the scutellum which is normally concealed. See Figure 3 of Volume I for additional morphological details.

Kleine described Kleineella piceonitens from Tahiti, but the unique holotype was either a mislabelled specimen or a specimen accidentally introduced by commerce and not established on Tahiti. It was said to have been found at the port of Papeete. Many years ago, R.D. Pope, British Museum (Natural History), tried to find the type for me without success. It could not be found in the Zoological Museum, Berlin, in 1962, and it may have been destroyed with Kleine’s collection. I consider piceonitens Kleine to be a new synonym of the Australian Kleineella barbata (Kleine).

Kleine described Brentus efferatus from the Marquesas. It is extraordinary that anyone would consider the presence of a species in an otherwise entirely American genus not to be an introduction by man to the isolated Marquesas Islands whose fauna is unrelated to and so distinctive from that of distant South America. I found the species on Hibiscus tiliaceus near the coast of Tahiti in 1934, but I have not heretofore published the record. I submitted specimens to R.D. Pope, and he has found Kleine’s name efferatus to be a new synonym of the widely distributed, variable Latin American Brentus cylindrus (Fabricius). Kleine evidently shared the mistaken belief of some unskilled workers who have often considered specimens of a species found distant from their known homes to be, ipso facto, different species.

New Caledonia has few described species, and there is only one endemic species in New Zealand. The Ethiopian Region, including Madagascar, whose fauna is distinctive, has the second richest fauna of more than 80 genera. There are about 50 genera in the Neotropical Region. Haedo Rossi, in his 1961 monograph, recorded 13 genera and 39 species in Argentina (the species occur only in the tropical and subtropical north of the country). Excluding the southern islands of Japan which have been penetrated by about 13 Oriental genera, some of which are introductions by man, and southern China, the Palaearctic Region is mostly without representatives of the Brentidae. Only three species reach southern Europe, but none is endemic. Excluding four genera that reach the warm southern boundary of America north of Mexico, only one species is found, and it is the only species that reaches as far north as the southern boundary of Canada. George Horn, 1872:127, said no territory in the world of equal size is probably as poorly represented in this family.

Excluding a few genera and species that have been reported only from Australia, and which I suspect are not endemic, Australia has undoubted endemic species in only three genera of myrmecophiles. These are Cordus, Kleineella and Ankleineella, but none of those genera is endemic. The myrmecophilous Cordus appears now to be in the process of explosive speciation, and within Australia many species remain to be described. In view of the unbalanced endemic brentid fauna, one may ask: Where was Australia when the Brentidae were distributed? It would appear that when Australia was at a colder higher latitude it was not colonised, or if it had an ancient fauna that fauna was exterminated by cold. In geologically recent time, when close contact with Indonesia became established, the Indonesian fauna began to populate Australia. Because of the present aridity of most of Australia, however, the fauna of the adjacent wet tropics has penetrated naturally little more than the northern border, parts of Queensland and northern New South Wales whose warm rainforests now provide suitable habitats for many species. The myrmecophiles are an exception; they have been able to disperse throughout the continent in company with their ubiquitous, multitudinous hosts. The brentids are not represented by endemic forms, excepting mymecophiles, in the ancient lands of Western Australia. I know only three brentid species (two Cyphagogus and one widely distributed Cordus) to be established in Tasmania, and they are introduced species probably carried by man.

Many Brentidae are subject to great, often confusing, individual variation in size and form. It is common to find more than 100% variation in size within a species. The heads of the males of the many species whose heads are sexually dimorphic often display bewildering variation. The femoral dentation is also subject to great individual variation, and the teeth may be large, small, rudimentary or absent. Unlike most weevils whose male abdominal discs are depressed or concave, and whose sexes are thus easily distinguished, the abdomens of most brentids do not display such distinctive features. Some brentid females have the first two ventrites as flattened or as broadly concave as those of the males, and sexing may prove difficult.

Sexual dimorphism is common in the Brentidae, and it is so extreme in some species that correct association of the sexes may be difficult, or the sexes may be considered to represent separate species or even different genera. The sexes of other species are so similar that it is often difficult to distinguish the sexes from external examination.

Little is known about the bionomics of Brentidae in Australia. Excepting Cylas, I do not know of Australian life history accounts of any species. Excepting Cylas, nothing is known about the actual food of any of the species in Australia. The larvae of the woodboring species appear to obtain sustenance from the micro-organisms and fungi in the decaying wood. Where the larvae of the myrmecophilous species occur and what they eat remains a mystery (see further notes under Cordus, Ankleineella and Kleineella below).

Because so few Brentidae occur in Europe and North America, little was written concerning their biologies before the studies of Beeson and Gardner in India (see the bibliography). Kleine gave a summary of data on biology, 1938:2. C.V. Riley, 1874:114, noted that the North American Eupsalis minuta (Drury) (known in some literature as Platysystrophus minutus ) belongs "to a group the true position of which has long perplexed systematists. This perplexity is, in a great measure, due to the fact that its larval structure has remained unknown, and that another larva has been mistaken for it. Dr. Harris, as early as 1838 [Insects Injurious to Vegetation, page 67] gave a very full account of the insect, but the larva which he describes as belonging to it … differs very materially from the genuine larva …. Harris mistook the rather similar-appearing larva of a tenebrionid, whose larval habits resemble those of the brentid, for the species. Riley corrected the error and for the first time described and illustrated the larva of a brentid. Riley noted that the female bores holes through the bark or surface of the wood of the host (often in the stumps of oak trees) and inserts her eggs therein. He quotes details on the biology of the species from a letter from W.R. Howard, as follows: It requires about a day to make a puncture and deposit the egg. During the time the puncture is being made, the male stands guard, occasionally assisting the female in extracting her beak: this he does by stationing himself at a right angle with her body, and by pressing his heavy pro-sternum against the tip of her abdomen; her stout fore legs serving as a fulcrum, and long body as a lever. When the beak is extracted, the female uses her antennae for freeing [the mandibles] of bits of wood or dust, the antennae being furnished with stiff hairs and forming an excellent brush. Should a strange male approach, a heavy contest at once ensues, and continues until one or the other is thrown from the tree. The successful party then takes his station as guard. These contests sometimes last for hours, and are always repeated if the proper male is defeated, though not often if he is successful … with instructive illustrations, has published an account;[If the challenger is] successful in routing his rival he takes the same care of the female as did the vanquished individual."

Meads, 1976:171, 176, with instructive illustrations, has published an account of his observations on the habits of the large Lasiorhynchus barbicornis (Fabricius), the only endemic brentid in New Zealand. He found the species attacked only recently dead or dying parts of trees of various species. He reported that The female bores a hole 0.5 mm wide through the bark by inclining her rostrum at approximately 45° to the body line and twisting her head from side to side …. She withdraws her rostrum after boring about 0.3 mm and ejects the debris collected inside the rostrum and between the mandibles, rather in the way a cork borer is used. This procedure is continued until a hole 3–4 mm deep is made 45° to the surface of the tree. The antennae are laid along the rostrum with the extremities touching the tree while boring .... When the hole is completed the female reverses her position and lays into the hole. She then tamps the hole with bark scraped with her valvifer from around the hole. [I believe this is incorrect. It is the serrated caudal tergite that is used. E.C.Z.] The entire process takes approximately 30 minutes. Copulation normally takes place when the female is boring and precedes oviposition …. Meads reported that the adults fed upon secretions from decaying tree tissues. He also reported that:

Fighting occurs frequently when an extra male or males find a copulating pair. An attacking male first uses his downwardly curved mandibles and long rostrum with a raking action across the back of the copulating male in order to dislodge him. If this is not successful both contestants endeavour, rather awkwardly, to secure any part of their opponent’s leg with their mandibles. If this difficult manoeuvre is accomplished and a firm hold is achieved, the defeated male immediately becomes submissive and is lifted bodily off the tree and dropped. Some beetles have legs missing (mainly tibia or tarsi) and this may be attributed to such fights. Smaller males tend to retreat quickly when challenged and do not fight. The defeated males spend long periods hiding in crevices, holes or under loose bark.

In his account of his second residence at Dubbo in the Aru Islands, between Australia and New Guinea, in May and June, 1857, A.R. Wallace, 1906:385, made the following observation on some Brentidae:

Those curious little beetles, the Brenthidae, were very abundant in Aru. The females have a pointed rostrum, with which they bore deep holes in the bark of dead trees, often burying the rostrum up to the eyes. The males are larger, and have the rostrum dilated at the end, and sometimes terminating in a good-sized pair of jaws. I once saw two males fighting together; each had a fore leg laid across the neck of the other, and the rostrum bent quite in an attitude of defiance, and looking most ridiculous. Another time, two were fighting for a female, who stood close by busy at her boring. They pushed at each other with their rostra, and clawed and thumped, apparently in the greatest rage, although their coats of mail must have saved both from injury. The small one, however, soon ran away, acknowledging himself vanquished. In most Coleoptera the female is larger than the male, and it is therefore interesting, as bearing on the question of sexual selection, that in this case, as in the stag-beetles where the males fight together, they should be not only better armed, but also much larger than the females.

W.S. Eberhard, 1979:254, includes a discussion of fighting between male brentids in his paper on the functions of the horns of Coleoptera.

I have observed peculiar fungi of the family Laboulbeniaceae growing on an unidentified brentid from Lord Howe Island. I have seen many wet forest Hawaiian Carabidae carrying the fungi, but I have not heretofore seen such infestations on any Curculionoidea.

The most detailed studies of the biologies of Brentidae are those of C.F.C. Beeson who worked in the Indian forest service many years ago (and from whom, in Banbury, England in 1950, I purchased some of the literature that has assisted me in this study). In 1925:172, he reported that "observations made by the writer suggest that the Indian species (at any rate) in the larval stage are true wood-borers and not bark-borers, and that their habitat is normally in felled or fallen timber and not in living trees. The adult beetles occur gregariously between the bark and wood of trees that have been attacked by bark and sapwood-boring larvae (mainly Longicornia); they often hibernate, under dead bark in company with histerids, staphylinids and clavicornia, but may be found in most months of the year. The eggs appear to be deposited singly on the surface of the sapwood under the shelter of the wood dust and fibres in the excavations of surface-borers. In many genera the female bores a hole in the wood with the mandibles for the reception of the egg. The larva constructs its own gallery radially into the wood, and feeds like the Lymexylonidae and Platypodidae on sap or saprophytic fungi growing in the gallery. Pupation usually occurs in a cell enlarged along the axis of the tunnel, and emergence takes place via the site of oviposition …. This simple type of life-history may be modified by the utilisation of the internal tunnels of other wood-borers, with the construction of larval galleries at right angles to the wall of the tunnel, as, e.g., the case of Microtrachelizus in the tunnels of Haplocerambyx spinicornis …. A further modification occurs when the utilisation of the gallery of another wood-borer involves the ejection of the rightful occupant, e.g., in the case of Cyphagogus and the Platypodidae.

"Since the life-cycle of brenthids progresses concurrently with those of primary wood-borers, the family should be ranked among the borers of freshly felled or fallen timber, rather than among those groups that establish themselves only after the timber has lost a high proportion of its moisture and has begun to decay.

The myrmecophilous group of brenthids includes two types: symphilines and robbers. The former live in association with ants nesting in the soil as well as in plants, and are characterised by the possession of exudatory organs. The latter are usually without excretory pores but exhibit other anatomical modifications. Nothing is known of their larval habits.

There are four basic types of life habits of Australian Brentidae:

Many species have larvae that are dead wood-borers, and the females of some species bore holes in the wood, or through the bark into the wood and lay their eggs therein, whereas others may deposit their eggs on the disturbed surfaces of their hosts or in the borings of other Coleoptera. The adults of some species may be found, often in groups, beneath loose dead bark of trees.

Cylas may be considered to represent a separate group, because its larvae bore in living tissues of stems and tubers of sound sweetpotato and allied Convolvulaceae, the adults also feed on similar live tissue, and the species is of considerable economic importance.

Cyphagogus and many allied forms have especially modified bodies that enable them to enter the tunnels made by other woodborers, such as scolytid and platypodid ambrosia beetles. They may extirpate the original owners of the tunnels, and then lay their eggs in the host tunnels from which their own larvae bore short tunnels outward. The repeated reports in literature from other lands that these brentids are cannabilistic are, I believe, erroneous. See further comment below under Cyphagogus.

Myrmecophiles are represented in Australia by Cordus, Ankleineella and Kleineella. Species of these genera have remarkable modifications on the heads to disseminate glandular fluids that are evidently utilised by the ants, but nothing is known about the fluids or their connection with ants. Strangely, although the adults of these species are often found in large numbers in ants’ nests, the larvae remain unknown. The myrmecophilus species are mostly nude and they assume the castaneous colorations of the ants. See further commentary under the generic headings below.

Figure 3 (Opposite) Some examples of tunnels of Indian brentid larvae in wood. a, tunnels of Cyphagogus corporaali originating from a gallery of Platypus solidus in Butea frondosa; b, tunnels of Cyphagogus westwoodi radiating from a gallery of Crossotarsus sp. in Vatica lanceaefolia (note irregular pitting of the Crossotarsus gallery and the pupal cells of Cyphagogus at the ends of the larval tunnels); c, longitudinal section of a Cerobates tunnel showing the apex narrowed toward the outer surface of the sapwood; d, transverse section of the prepupal tunnel of Hoplocerambyx spinicomis in Shorea assamica showing origin and trend of tunnels of Microtrachelizus beneficus (note location of pupal cells); e, longitudinal section of the prepupal tunnel and pupal chamber of Hoplocerambyx spinicomis showing the location of the exit holes of Microtrachelizus beneficus; f, longitudinal section of tunnels of Cerobates tristriatus in Bombax malabaricum. (After Beeson, 1925.)

Masters, 1886:508, listed 16 species of Brentidae in Australia in the second edition of his catalogue of the Australian Coleoptera. Britton, 1970:518, in Insects of Australia, estimated that there were about 33 species in the country. In this text I report on 85 species, but I have seen other species whose identities I have not been able to determine or which are new species. Perhaps not more than about half the species of Brentidae in Australia are known.

Taxonomy

The suprageneric status of the brentids has been the subject of much difference of opinion among taxonomists, and in recent years the problems have received increased attention.

Crowson, 1955:163, without making a detailed morphological study, contributed some pertinent remarks without reaching a firm conclusion as to the family status, and he placed the brentids as a family adjacent to the Apionidae. He noted that "the articulated larval legs however remain as an anomalous feature, only found otherwise in the most primitive of Curculionoid larvae, i.e. the Nemonychids and Anthribids; assuming that the closest affinity of Brenthids is really to Apionids we shall be obliged to assume either that the loss of the larval legs has taken place independently in the ancestry of Attelabids, Apionids, Belids, and certain Anthribids (e.g. Bruchela) or that larval legs in Brenthids have been secondarily regained – in which case one might pertinently ask – why only in BRENTHIDAE?" Only some brentid larvae have legs, and the feature is variable within the family.

Morimoto, in his detailed, nine year modern study of the comparative morphology and taxonomy of the Curculionoidea (1962a), found certain similarities between the brentids and apionids. In 1962b:31, Morimoto merged the brentids, cyladids, apionids, nanophyids and ithycerides into one family, and unfortunately, he added tentatively the distinctive desmidophorids as a subfamily. After maturing his opinions, he published a revised classification (1976:469). In this latter study he treated the brentids as a family distinct from the Apionidae, he gave full family status to the Ithyceridae, and he correctly abandoned his former position on the Desmidophorinae as an included group. In his new proposal, Morimoto included the Australian Eurhynchus (= Eurhinus) and Cylas in a subfamily Eurhininae (recte Eurhynchinae) of the Brentidae, and he moved Cybebus and Myrmacicelus from the Cyladini, where Kissinger had placed them, to the Apioninae. I have given my opinions regarding Cybebus and Myrmacicelus elsewhere here under Apionidae, and they need not be repeated. I do not agree that the Eurhynchidae should be included in the Brentidae as Morimoto, 1976:469,has suggested, or in the Apionidae as Crowson, 1955:164, and Kissinger, 1968:10, have done. I believe that some workers may have been unduly influenced by the superficial similarity of formation of the antennae of Cylas and Eurhynchus and have concluded that this feature is indicative of family relationship. See my chapter on the Eurhynchidae for my conclusion regarding that group.

Those who believe that the Brentidae should be merged with the Apionidae have overlooked certain features in addition to the character differences indicated by Crowson and Morimoto, both of whom have not mentioned them. Many years ago I observed the unusual formation of the eyes of the Brentidae: if the eyes of any of a multitude of curculionids are examined, it will be noted that the ommatidia are clearly defined on the surface, and many eyes have a more or less blackberry like appearance. If, however, the eyes of a brentid are examined, it will be noted that they have a unusual appearance – the ommatidia are covered by a remarkable cyrstalline lens, and if they are examined under a scanning electron microscope their surfaces appear smooth or almost smooth. I thought my observation was original, but recently I have found that John LeConte, that acute observer and noted pioneer American coleopterist, had noticed this feature a century previously. LeConte, 1876:324, wrote: "The smooth eyes, the reticulations of which are seen only through the transparent integument and the form of the front tibiae, indicate a resemblance, though a remote one, to Rhyssodidae …." It is surprising that, to my knowledge, no author has given any further attention to this remarkable feature. It serves easily to separate the Brentidae, including Cylas, from the Apionidae and other Curculionoidea.

An interesting feature of the Brentidae, and one ignored in the literature, is the almost universal lack of true squamae (in contrast to hairs or setae) on the adults. They share this characteristic with the primitive weevils. Many Apionidae also lack squamae, but a multitude have squamiform setae. The Rhynchophoridae also lack true squamae. It is not until one advances up the taxonomic scale to the advanced weevils that wide, fluted or fanlike squamae become commonplace.

Antennal cleaners are well developed on the protibiae of most Brentidae (compare Belidae, also), but I have observed no similar structures on the Apionidae. The development of the antennal cleaners varies within the family, and it reaches its peak of distinctiveness in Cerobates in which species the inner margins of the protibiae are remarkably excised and the excision is clothed with stiff setae to form a concave brush (see Figures 174, 176).

Tibial spurs are characteristic of primitive Curculionoidea, and they are rarely found on only a few of the higher Curculionidae. I have observed none on the Apionidae. They are a feature of the Brentidae. On the specialised Cylas they have been lost from the protibiae, but they are present, although rudimentary and apparently variable on the meso- and metatibiae of Cylas formicarius. Cylas under the influence of its altered biology, evidently no longer requires them, and they are lost or in the process of being lost in that genus. Most workers have overlooked the tibial spurs of the Brentidae. The basic, and primitive, spur formula is 2-2-2, and there are variations upon this, often 1-2-2 or 1-1-1.

All Brentidae have simple tarsal claws (plesiomorphic); none is known with toothed claws. Most Apionidae have toothed tarsal claws (apomorphic). The ratio of toothed to simple claws in Australian Apion probably exceeds 25:1.

The scutelli on all of the Apionidae (excluding Myrmacicelus and the Nanophyidae on which they are also concealed) that I have examined are exposed on those species whose wings are developed for flight. The scutellum is reduced or concealed only on those species whose wings are non-functional or vestigial. The scutellum, however, is concealed on all of the volant Brentidae. Why should the scutelli be concealed on all winged brentids? This is an extraordinary and divergent characteristic. It is normal for flying Curculionoidea to have exposed scutelli. Another associated unusual condition is the reduction of the width of exposure of the metepisternum on the volant Brentidae.

Lacordaire, 1863:529, with his usual skillful observation, noted the concealed scutellum, the narrow metepisternum, the structure of the mesosternal pleura and the form of the elytra, whose humeri are effaced, and these features led him to the seemingly logical but false conclusion that Cylas species were apterous.

The internal sacs of the aedeagi of most Brentidae contain an unusual ()-shaped pair of sclerites (Figures 162a, 178a). The monotonous stability of this simple feature from genus to genus and tribe to tribe is noteworthy. The structures also occur in the Eurhynchidae (Figures 232a, 244a) and in Myrmacicelus (Figure 288; see also Kissinger, 1968, Figure 6f). Although some Australian Apioninae have paired sclerites in the internal sac, I have not observed the equivalent of the brentid form, and the architecture of the aedeagus differs. The armature of the internal sacs of the aedeagi of most Apionidae is more complicated (apomorphic). Alonso-Zarazaga, in a recent paper (1989:222) calls the ()-shaped sclerites frena, and in his 1990:28 monograph on the Palaearctic Apionidae, he notes that Frena are absent except in some primitive Tanaonini. Kissinger, 1968, Figure 8, illustrates them in Tanaos. Sanbourne, 1981:63, Figures 99–103, illustrates similar structures in the Ithyceridae. Somewhat similar sclerites occur also in the Nemonychidae (see illustrations in Kuschel, 1989 and in Volume I of Australian Weevils).

A flagellum is present in the aedeagus of most (all?) Brentidae. They are present or absent in the Eurhynchidae, present or absent in the Belidae, and present in the Ithyceridae and Nemonychidae. I have not seen a flagellum in any Anthribidae. Few apionids have flagelli. Kissinger, 1968, Plate 119, a, b, illustrated a species with a flagellum without stating that it is a flagellum, and a flagellum in his figures of Myrmacicelus. In this text I illustrate the well-developed flagellum of Lissapion varistriatum and of Myrmacicelus. Alonso-Zarazaga, 1990:28, said There is no flagellum in Palaearctic Apioninae …. The nanophyids have well-developed flagelli.

If one observes an alert brentid of many species, including Cylas, it will be noted that the front end is often elevated and the body assumes an inclined position (Plates 4, 67). This feature may be observed on many dead museum specimens. Compare, for example, the accompanying illustrations of species of Cordus. This differs markedly from the Apion I have observed.

In sum, I believe that the Brentidae are a distinctive group worthy of being entitled to full family status. The characters they share with some other groups indicate affinity but not identity. Cylas, whose larvae have legs, as do many Brentidae, is neither a typical brentid nor a typical apionid, but its assignment to a distinctive subfamily of the Brentidae more accurately expresses its affinities than associating it with the Apionidae. If it were not to be considered to belong to the Brentidae, then, in the light of its uniqueness, it may be best to consider it to represent a separate family, the Cyladidae, annectent between the Brentidae and the Apionidae. Lacordaire, 1863:529, included Cylas and Myrmacicelus in his Cylades, a group to which he gave equal status to that of the Belidae, Eurhynchidae and Apionidae. LeConte, 1876:327, treated Cylas and Myrmacicelus (misspelled Myrmecacelus) as members of one of two subfamilies (Cyladidae = Cyladinae) of the Brentidae.

In 1977, in an obscure journal, which evidently is not held in any Australian institution, Kasap and Crowson, his professor, published a paper entitled A Comparative Study of the Internal Anatomy and Abdominal Structures of Curculionoidea in which, inter alia, they reported upon two Brentidae. Such a study is particularly relevant today, but, unfortunately, this paper contains a number of faults, falls short of expectations, and it does not meet the high standards demanded of others by Crowson. Kasap and Crowson report that they studied the North American Eupsalis minuta (Drury) and the Indo-Australian Cerobates australasiae Fairmaire. Eupsalis minuta is correctly known as Arrhenodes minutus, (see Kuntzen, 1937:190); from 1917, when Kleine erected a new genus for it, until 1937, the species was known as Platysystrophus minutus), and Cerobates australasiae is an old synonym formerly applied to the female of Cerobates ophthalmicus (Pascoe). Although Kasap and Crowson examined only the two named brentids, they make some wide-ranging generalisations regarding the Brentidae. I do not have available living or fixed specimens of the species they studied, and I can now comment only upon some of their statements, some of which are erroneous and others require critical examination and confirmation before being accepted. The paper serves as a stimulant to further study, but it must be used with caution.

On page 38, Kasap and Crowson note that the two brentids studied lack proventriculi; Arrhenodes has no mid-gut regenerative crypts which are present on Cerobates; Arrhenodes has four Malpighian tubules grouped 2 by 2 at both ends; anteriorly they open ventrally to the end of the mid gut and posteriorly join 2 by 2 into 2 common ducts before attaching to the colon.

On page 44 they note that "In Eupsalis [Arrhenodes]… the structure of the vasa efferentia, vesiculae seminales and the opening of the accessory glands to the vasa efferentia instead of vesiculae seminales are very different from those of the others. If such differences occur in also other brenthids, then this family can be distinguished from other Curculionoidea by these structures." These details merit detailed investigation.

On page 55 they say "We discovered a pair of round glands underneath tergite 7 of Eupsalis and Cerobates (Brenthidae). Each gland discharges via a small duct into a small hole in the tergite 7. In both species there is an internal ridge in the posterior region of the elytra, which help to delimit a special space beyond the tips of the folded wings between the most posterior part of the elytra and tergite 7. It is probable that the secretion of these glands is discharged and stored in this space. These details have been based upon incorrect observation and appear to be false. I have examined specimens of both species, and I have found no such structures opening through tergite 7. Furthermore, should there be such remarkable structures discharging a fluid into the cavity between the internal apices and the abdomen, such a fluid would be lost whenever the insect opened its wings or flew! The internal ridge" mentioned is the subcostal plica whose apex characteristically turns inward toward the elytral suture on the Brentidae, thus leaving a hiatus between the ends of the retracted wings and the elytral apices.

On page 43, they report that on Arrhenodes (Eupsalis) "the accessory glands open to the vasa efferentia, thus presenting an unusual situation as they open to the vesiculae seminales in other Curculionoidea. The vesiculae seminales are soft and membranous-looking whereas they are usually heavily musculated in the others. So it may be that in Eupsalis the function of the true vesiculae seminales is taken over by the vasa efferentia, which may therefore be analogous structures to the vesiculae seminales of the other species studied. And on page 44, they conclude that the structure of the vasa efferentia, vesiculae seminales and the opening of the accessory glands to the vasa efferentia instead of vesiculae seminales are very different from those of the others. If such differences occur in also other brenthids, then this family can be distinguished from other Curculionoidea by these structures."

On page 47 they report that "Brenthidae, Apionidae (except for Antliarrhinus with 3) and Curculionidae (including Platypodidae and Scolytinae) can be distinguished in having two ovarioles per ovary, whereas there are at least four ovarioles per ovary in the other groups studied."

Kasap and Crowson found "The tergal muscles of Eupsalis are very unlike those of other genera studied; there are only [tergal longitudinal muscles] in tergite 1 and tergites 5–7 …. And on page 64 they note that of the groups studied, Nemonychidae, Belidae and Bruchela (Anthribidae) have a complete set of sternal muscles. In Curculionidae only the sternite 3 and in Attelabidae at least the sternite 3 have no muscles, whereas Araecerus (Anthribidae), Brenthidae and Apionidae studied have no muscles in the sternites 3–6."

They conclude, on page 68, that "In most characters studied by us, Eupsalis and Cerobates agreed with Apionidae, but in the abdominal nerve chord ... they showed an additional free ganglion more or less as in Nemonychidae. In larval Brenthidae, a comparable feature is the frequent presence of short but distinct thoracic legs, otherwise practically confined to Anthribidae in Curculionoidea – however, short but distinct leg rudiments have also been reported in some Apion." It would have been helpful had references to the latter been given. Could they be referring to Emden’s 1938:17 mention of pedal discs? Pedal discs are not the same as distinct legs.

For many years the study of the Brentidae of the World was dominated by Richard Kleine who died in 1949. Kleine’s work was mostly poor, much of what he wrote is rubbish that should never have been published, and he left a legacy of extraordinary confusion. To compound the difficulties, his large collection was destroyed at Stettin-Pomern during World War II. For further comment on the work of Kleine, see Damoiseau, 1963a.

A new approach to brentid taxonomy was made by Damoiseau (1962–1968), but his early tragic death brought his work to a premature end before he could mature his ideas. His work does, however, contain some unusual faults, especially an inadequate treatment of genital study. My research for this chapter suffered a severe blow by Damoiseau’s loss, because he had been assisting me with some identifications and aiding in solving problems. There is now no widely experienced authority on the family in Europe, where most of the relevant material for the study of the Australian fauna is stored, to whom one can appeal for guidance.

The Australian Brentidae have received little attention from Australian entomologists. The family is poorly represented in most Australian collections, there is little Australian literature on the fauna, no keys have been prepared for it, and it has been difficult for Australians to make or to obtain identifications. To compound the difficulties, most of the Australian Brentid fauna is geologically recent immigrant or introduced through commerce, and, because of the lack in Australia of comparative collections, especially from Indonesia, Australians have had to rely upon European institutions, especially the British Museum, for identifications. For many years Papua New Guinea was under Australian administration, but the Australian authorities did nothing regarding entomological surveys. Because of the lack of representative collections from the New Guinea area and islands to the northwest of our continent, the Australian National Insect Collection has suffered an irreparable loss which has placed Australian workers under severe handicap. Because of these problems, and because I have been unable to visit collections overseas to name them or to have them identified by European workers, I have been unable completely to record in this text various additional genera and species that I have seen.

The taxonomy of the Brentidae remains unsatisfactory, and it is not greatly changed since the system proposed more than 100 years ago by the master coleopterist, Lacordaire. It was Lacordaire, 1866:399, who first laid a foundation for the classification of the Brentidae of the world. He established the following taxonomy:

Family BRENTHIDAE

Tribe BRENTHIDES Vrais

Group Taphrodérides (including present Cyphagogini)

"        Ischnomérides

"        Héphébocérides

"        Trachélizides

"        Arrhénodides

"        Bélophérides

"        Bélorhynchides

"        Eutrachélides

"        Brenthides vrais

"        Céocéphalides

"        Némocéphalides

"        Ithysténides

Tribe ULOCERIDES

Lacordaire’s taxQnomic system has been modified in relatively minor ways as many new genera have been discovered and described, but much of what has been written between the time of Lacordaire more than 100 years ago and the recent works of Damoiseau has been based mostly upon superficial study. Even today various morphological and anatomical features of the Brentidae remain ignored or inadequately studied by taxonomists. Little attention appears to have been given to the features of the buccal cavity since Lacordaire’s 1866:400 discussion. Both the early writers, Olivier and Latreille, mentioned the subject, and Westwood, 1839:334, wrote The structure of the mouth is very curious, and has not yet been properly examined. The fact that the Brentidae display both adelognathus and phanerognathus conditions usually has been overlooked or ignored. LeConte, 1876:323, noted that the mouthparts of Cylas (which is phanerognathus) differed from the limited number of Brentidae that he had examined. Such features define major subdivisions of the Curculionidae. Kleine examined the genitalia of many species, and he wrote a separate paper entitled Meine Praeparationsmethode des Kopulationsorganes (1917c), but his methods were crude and his numerous illustrations mostly misleading or worthless. Damoiseau made a more careful and detailed study of the male genitalia, but his illustrations were made mostly from grossly distorted specimens flattened on slides. I have seen some of his slide mounts and have found them rendered worthless by deterioration of the mountant. The female genital systems