the south american weevil genus rhyephenes

1. New York Emomol. Soc. 104( 1- 2): 1- 20, 1996
THE SOUTH AMERICAN WEEVIL GENUS RHYEPHENES
(COLEOPTERA: CURCULIONIDAE; CRYPTORHYNCIDNAE)
J UAN J . M OR RONE
Laborat ori o de Sistem atica y Biologia Evolutiva (LAS BE), Mu seo de La Plata,
Paseo del Bosque, 1900 La Plata, Argentina
Abstract. - T he ge nus Rhyephelles Schoenhe rr, e nde mic to the Ce ntra l C hil ean and Subantarctic biogeographic provinces of southern So uth A me rica, is eas ily recog ni zed by its bl ack
body w ith shape rese mbling a spider. This ge nus includes seven species: Rhyephenes c:/mhratus
R. Phili ppi, R. gayi (Guerin), R. gOl/real/i (Gay & Solie r), R. humeralis (Gue rin), R. ImeraUs
(Guerin), R. maillei (Gay & Solier), and R. sql/all/iger F. Philippi (rei nstated he rei n fro m synony my w ith R. gay;). A key, redescri ptions, habitu s photographs. and illu strations of the spec ies
are prov ided, and the ir geographical d istri bution is mapped. A cladi sti c analys is usin g 2 1 characters fro m external mo rpho logy and ma le a nd fe male genitalia produced four cladograms (CI
= 0.55, RI = 0.51. length 43 steps), which after success ive we ig hting we re reduced to one
c1adogra m (C I = 0.82, RI = 0.85, length III ste ps). In the cladogram. the fo llowi ng phylogenet ic seque nce results: (R. squall/iger. ( R. ImeraUs. « R . clalhratus. R . gOllreaui), (R. gayi.
(R. humeralis. R. maillei))))). Biogeographic patterns ex hibited by these species indi cate a
seq ue nce fro m central Chile to southern Ch ile and Argentina.
The weev il genus Rhyephenes (Curculionidae: Cryptorhynchinae) was described
by Schoenherr (1 837). This taxon of uncertain placement is one of the most characterislic weevil genera from central Chile, wilh two species extending also to southern Chile and Argentina. They are popularly known in Spanish as " arafiitas" (little
spiders), because of their typical habitus (see Figs. 1- 14). Sixteen species-group
names proposed by se veral authors (Guerin, 1830 ; Erichson, 1834 ; Gy llenhal, 1837 ;
G ay & Solie r, 1839 ; Blanchard, 1853; R. Philippi, 1859; F. Philippi , 1899; Fiedler,
1942) were assigned to Rhyephenes. This nomenclatural pro liferation is mainly due
to the intraspecific variability of the species of Rhy ephe n es, which led authors to
describe as valid spec ies mere geographical variant s. Recent syno nymies by Kuschel
(1 99 3) and Morrone (1 994) reduced the number o f species to six . I suspected, ho wever, that the synonymy o f R. squamiger F. Philippi and R. gay; (Guerin) by Kuschel
(199 3) was incorrect . A cladistic anal ys is of its species can help ellucidate which
are the vaJid species of Rhyephe n es.
MAT ER IAL AND M ETIiODS
The specimens examined in this study are from the following colJections: AMNH ,
American Mu seum of Natural History, New York, USA (Lee Herman); BMNH, The
Natural Hi story Museum, London, Great Britain (Christopher Lyal); CWOB, Charles
W. O ' Brien private collection, TalJahassee. USA (Charles O ' Brien); lADLZA, Inslituto de Investigaciones de las Zonas Aridas, Mendoza. Argentina (Sergio Roig-Jufient); lPCN, Instituto Patag6nico de Ciencias Naturales. San Martin de los Andes,
Argentina (Mario Gentili); MACN, Museo Argentino de C iencias Naturales Bernardino Rivadavia, Buenos Aires, Argentina (Axel Bachmann); MHNS , Museo Na-
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104( 1- 2)
Table I. Data matrix and li st o f characters used in the clad isti c analysis of Rhyephe"es.
plesiomorphic: 1. 2. 3 = apomorph ic.
o=
outgroup
R. clatlirafl/s
R. gay;
R. gOllreau;
R. humeral;s
R. lateralis
R. maille;
R. sqllamiger
00000 00000
11001 11001
10010 11000
0000 1 11001
2 1110 IIIII
22000 10001
2 1010 IIIII
000 10 00101
00000
21 100
2 1111
0 11 0 1
21 101
3 1000
01000
10000
00000
00010
11100
0 111 0
12101
0 100 1
11100
00100
0
0
I
I
0
I. Tubercles on male rostrum: (0) absent: ( I) present at si des: (2) present on dorsu m and
sides.
2. Punc lUres on female rostrum: (0) bro ad. abu ndant: ( I) finc. Icss abundant: (2) fine . s parse.
3. Prothorax : (0) sligh tl y g lobose: (I) strongly globose. prolUberant.
4. Relati ve width of the prothorax : (0) as wide as elytra: ( I) w ide r than e lytra .
5. Prothoracic anterior impress ion: (0) absent: ( I) present.
6 . Pro tho rac ic punctures: (0) small . sha llow: ( I) large. dcep.
7. Elytral basal area : (0) slig htl y imprcssed: ( I ) strong ly impressed .
8. Elytral tubercles: (0) rounded: ( I) subconical.
9. Elytral tubercles o n interval s: (0) small , similar to those o n striae: ( I) large. more developed
than thosc on striae .
10. Elytral tubercles o n interval s and striac: (0) c learly separated: ( I) part iall y fused.
1 I. Elytral humeral stripes o f white scales: (0) abse nt; ( I) two short stripes: (2) three short
stripes: (3) three stripes. two sho rt and o ne lo nger.
12. Body vestiturc : (0) clothed with scales: ( I) lacki ng scales.
J 3. Acdeagus in latera l view: (0) slender: ( I) robust.
14. Ostiolar sc1eritcs of aedeagus: (0) rounded: ( I) acute.
15. Parameres: (0) long. narrow: (I) sho rt. broad .
16. Femal e sternu m 8 plate : (0) as lo ng as w ide: ( I) longer tha n widc.
17 . Re lative le ngth o f female sternum 8 p late: (0) < 0.40 X length of manubrium : (I) 0.41 0 .61 X length of ma nubrium : (2) > 0 .61 X length of manubrium.
18. Setae on sides of fe mal e sternum 8 plate: (0) absent ; (I) present.
19 . Arms of female ste rnum 8 plate : (0 ) narrow: (I) broad .
20. Spermalhecal nodulus: (0) developed: ( I) not deve loped .
21 . Spermathecal ramu s: (0) developed : (I) not developed.
cional de Historia Natural. Santiago, Chile (Mario Elgueta): and MLP, Museo de La
Plata. La Plata. Argentina (Juan Schnack).
Drawings were made with a camera lucida attached to a stereoscopic microscope.
Full data o f type spec imens are cited enclosing information from each label with
square brackets with each line separated by a slash.
Characters were derived from the external morphology , and male and female genitalia. The classification of the subfamily CrYPlOrhynchinae is in some disarray (LyaJ.
1993) and there are no keys avai lable to the South American genera (O'Srien. 1984 ).
Furthennore, the distinctness of Rhyephenes makes it presently not possible to identify its sister group, so an uncooted ingroup analysis was performed, and the root of
the cladogram was determined a posteriori using an hypothetical outgroup, following
1996
THE WEEVIL GENUS RHYEPHENES
3
the procedure described by Nixon and Carpenter (I 993). In order to construct thi s
out grouP. several South American genera were examined and the revisi on of the
New Zealand Cryplorhynchinae (Lyal. 1993) was consulted. Table 1 contains the
data matrix and the list of characters analyzed (all multi state characters were treated
as a.dcliti.v~). Analysis was carried out with Hennig86 1.5 (Farris, 1988), applying
the ImplicIt enumeration opt jon and the successive weighting procedure. CI and Rl
were calculated excluding autapomorphies. CLADOS 1. J (Nixon, 1992) was employed for examination of character distributions and for rooting the cladogram after
the unrooted analysis.
RHYEPHI£N£S SCHOENHERR,
1837
RhyepIJenes Schoenherr, 1837:312 (type species R. incas GyJlcnhal [=R. humeralis
(G uerin)], by original designation): Gay and Solier, 1839:24 (key): F. Philippi.
1899:3 (revision); Fiedler, 1942:274 (key).
PhysotltorLlS Gay and Solier, 1839:22 (type species P. moille; Gay and Solier. by
original designaLion); Solier. 1839:L (=Rhyephenes) .
Rllyephenes is easily recognized by its black color and habitus. which resembles
a spider (Figs. 1- 14).
Biology: Adult Rhyephenes are usually seen on woody plants, fe mal es ovipos it under bark of Lhe host plants, and larvae are endophytic, excavating galleries and
feeding on live wood (Angulo, 1970; Elguet3. 1993). Table 2 includes a li st of the
known plants where species of Rhyephenes have been collected. fro m personal obse rvaLi ons in the field, spec imen labels, and relevant literature (Fiedler, 1942: Havrylenko & Winterhalter, 1949; So lervicens & Elgueta, 1989; Barriga et aI .• 1993;
Elgueta, 1993; Morrone & Roig-Jufient , 1995). These plants cannot be certainly
assumed to be host plants.
Distribution : The species of Rhyephenes occur in the Central Chilean and Subantarctic provinces of the Andean subregion of southern South America (Figs. 50- 53).
KEY TO SPECIES OF RHY£PHEN£S
I. Elytra with 2- 3 humeral stripes of white scales (Figs. t , 3. 7. 9. 13)
.. 2
Elytra lack..ing humeral stripes of white scales (Figs, 5. II ) , , , , , , , , , , , , ... , ..... 6
2. Prothorax strongly g lobose. prolUbcr.uu (Fig . 7): c lytra with tubercles o n inter val s more
developed than those o n striae . . . . . . . , . . . . . . . . . . . . . . . . R. !mmeralis (Figs. 7-8)
Prothorax slightly globose: elytra with tubercles o n interval s sim ilar to those on striae
.. .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Elylra with basal area slig htl y impressed : aedeugus s le nder in lateral view (Figs. 24.
28)
............... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........
.. 4
Elytra w ilh basal area strongly impressed : aedcagus robust in latera l view (Figs. 16.
18) . . . . . . . . . . . .
.. .. . . . . . . .. . . .. . . .
. . . .. . .
. ......... 5
4 . Body clothed with scales (Fi g. 13): male rostrum lac king tubercles : fe male rostrum with
broad. abundant punctures: prothornx wider than clytra. and with small . shallow punctures: clytra with subconical tubercles and two shon humeral SLOpeS (Fig. 13)
. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . .
R. squamiger (Figs. 13- 14)
Body lacking scales (Fig. 9): male rostrum w ith tubercles: female rostrum with fin e.
sparse punctures: prothorax as wide as e lytra. and with large. deep punctures; e lytra
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JOU RNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1 - 2)
4
6
8
Figs. 1- 8.
Rhyt!phelles .!Ipp., habitus. 1.3,5,7. dorsal view; 2. 4. 6, 8, lateral view. I , 2.
R. cfarhratus: 3, 4. R. gay;: 5, 6, R. gOll rellui: 7. 8. H. !lIIl1/eralis.
1996
THE WEEVIL GENUS RHYEPHEN ES
9
II
12
Figs . 9 - 14. Rhyephenes s pp ., ha bitus. 9, II. 13. dorsal view; 10, 12. 14, lateral view. 9,
10. R. la teralis ; II , 12. R. ma illei ; 13, 14. R. squamiger.
with ro unded tubercles and two s ho rt and one lo nger hume ral s tripes ( Fig. 9)
R. lareralis (Figs. 9- 10)
5. Protho rax as w ide as elytra and wit.h ante rio r impressio n ; e lytra with tu bercles partiall y
fu sed; fe male rostrum with fin e, less a bundant punctures
R. c/at " ra lu .~ (Fi gs . J- 2)
Protho rax wi der than elytra and lacking ante rior impression : e lytra w ith tubercles clearly
separated : fe male rostru m w ith broad, abundant punctures
R. gay; (Figs . 3-4)
6. Prothorax wide r than e lytra and lacking ante rior impression ; e lytra w ith basal area
5
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOC I ETY Vol. 104( 1- 2)
strong ly impressed . and subconi ca l tubercles; aedeagus slender in latera~ vi, cw: male 'J
. . . .
R. maille, ( F igs. 11 - 1-)
Prolho rax as wide as e lytra and wi th a nterior impression: c lytra with basal area slig ht ly
impressed . and rounded tu bercles: aedeagus robust in latera l view; mal e rostrum .Iac ~ing
tubercles.
. .... . . . . . . . . . . . . . . . . .. .......... . . R. g{)uream (FIgs. 5- 6)
rostrum with tubercles
Rhyephenes clathratus R. Philippi . 1859
(Figs. 1, 2, I S, 16,29,36.43,50)
Rhyephenes clathratus R. Philippi , 1859:666; Fairmaire. 1860:251 (= R. goureaui);
F. Philippi, 1899:84 (reinslaled) .
Rhyephenes clathraws was cons idered a synonym of R. gOllreaui by Fairmaire
( 1860). Accordi ng to thi s analysi s, they are sister taxa; the lalter is distingui shed by
the e lytra lacking the humeral stripes o f whit e scales and by characters of the female
gen italia.
Redescription: Habitus (Figs. 1, 2). Body clothed with scales. Prothorax sLi ghtl y
g lobose, as wide as elytra ; di sc with anterior impression, and large and deep punctures . Elytra with basal area strongly impressed ; with rounded tubercles; tubercles
on intervals small, similar to those on striae, and partially fused to them ; humeri
with three short stripes of white scales. Male. Rostrum with tubercles at sides. Aedeagus (Figs. IS. 16) robust in lateraJ view; osti olar sclerites ro unded; parameres
long, narrow. Female. Rostrum with fine punctures. Sternum 8 (Fig. 29) with plate
as long as wide , and sides lacking setae; arms broad. Hemi sternite (Fig. 36). Spermalheca (Fig. 43) with nodulus and ramus developed . Length 7.8- 13.7 rum.
Type material: Lectotype male (here designated: [LECTOTYPElRhyephenesl
cla thratus/R . Philippi] [Rhyeph elleslclathratus/R . Phil./det. G. Ku sc hell 1990J
(MH NS).
Other material examined: C HILE. Without more precise data: 23 ( 1 AMNH , 3
BMNH, 19 MHNS). Caulin: Caulin, L. E. Peii a. I (CWOB). C uric6: La Mo nlaii a,
8- IX- 1970. A . Eglilis, 3 (MHNS). Limarf: Las Trancas, 12-1- 1978, D. Jackson. I
(MHNS). Malleco: Mall eco. XlI - 1986, G. Perez de Arce. I (MHNS). Talca: Constituci 6n. pi no insigne, 25-lX - 1970, E. Holsten, 2 (MHNS ); Los Cipreses, 1,000 m,
14-1- 1968, L. E . Pena, 2 (MHNS).
Rhyephenes gay; (G uerin. 1830)
(Figs. 3. 4 , 17. 18.30.37,44,50)
Tylodes gay ; G uerin, 1830: 125.
Rhyephenes cacic IIs Gy llenhaJ , 1837:3 15 .
Rhyephelles gay;; Sc hoenherr, 1844:402 ( = R. cacicus ): Lacordaire, 1866: 108 (reinstated).
This species occurs abundantly in centra] Chile. It is identified by the acute osti o lar
sc1erite o f the aedeagus.
Redescription: Habi tus (Figs. 3, 4). Body clothed wi th scales. Prothorax sl ighll y
globose, wider than e lytra ; disc lacki ng anterior impress ion, and large and deep
punclUres. Elytra with basal area strongly impressed; w ith rounded tubercles; tuber-
1996
THE WEEVI L GENUS RHY£PH£N£S
7
Table 2. Known pi am associations of the species of Rhyephelles (plant families and species
arranged alphabetically).
- Asteraeeae
sp.
Ambro.~ ia
R. gay;
Bahia ambrosioides
Bocc:hari.f sp.
R. squamiger
R. sqllamiger
R. squomiger
R. gay; and R. IlUmeralis
- Bt!rberidaceae
Berberis TOlIIndifolio
R. IlIlmeruJis
iJlIcc/Ulri s COIIC(lI '{l
Baccharis rhomboidulis
- Boraginaceae
Heliotropium .~ te"op")'llum
R. squami[:er
- Bromeliaceae
I~uya
chi/ensis
R. squumiger
-Celas traceae
M ay tenus boaria
R. maille;
-Fabaceae
Adesm ia mic rophylla
Anorlhrophyllum sp.
Lupinlls sp.
Selina cumming;;
Sen na mU/lig lallt/u/osa
SelllUi sp.
R. gay;
R. gay ;
R. humertllis
R. gay;
R. squomiger
R. gay;
-Juglandaceae
lug /ailS r egia
R. humerali.f
- Lauraceae
Per.fell a meric ana
R. squamiger
-Nothofagaceae
NOlhofag lls amar-elica
N. dombe),;
N . nilida
N. pumilio
NOlhofilgus sp.
R. moille;
R. IlIImeralis and R. moille;
R. moille;
R. moille;
R. IlIImeralis and R. moille ;
- Pinaceae
Pinus rad;ata
R. cJlIlhralll.f. R. Iwmeralis. and R. moille;
- Rosaceae
Prllnus pers;ca
Quillaja saponar;a
R. humera l;s
R. maillei
cles on interval s small , similar to those on striae. and clearly separated from them ;
humeri with three short stripes of whi te scales. Male. Rostrum with tubercles at
s ides. Aedeagus (Figs. 17, 18) robust in lateral view; ostiolar sclerites acute; parameres short. broad. Female. Rostrum with broad . abundant punctures. Sternum 8
(Fig. 30) with plate longer than wide. and sides with setae; arms narro w. Hemisternite
(Fig. 37). Spermatheca (Fig. 44) with nodulus developed and ramus not deve loped.
Leng th 7.5- 15 .5 nun .
Material examined: CHILE. Without more precise data: 102 (9 AMNH, 4 BMNH ,
89 MHNS). 8io-8eo: Fundo Maria Ester, 15 Ian W Victoria, 14-1- 1989, M . Ram irez,
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY VoL 104( 1-2)
15
I6
17
n~
-j
\8
\9
20
21
22
~
~A
2J
24
2S
26
27
28
Figs. 15- 28. Rhyephene.~ spp .. aedeagus. 15. 17. 19. 2\. 23, 25. 27, dorsal view; 16. 18.
20,22,24.26.28. lateral view. 15, 16, R. clarhrarus: 17. 18. R. gay;; 19,20. R. gOllreaui; 21,
22. R. Iwmerali!;: 23, 24, R. lateralis; 25. 26, R. maille;; 27. 28. R. sqllamiger.
2 (MACN). Cachapoal: Tennas de Cauquenes, 11-1-1953, 2 (MHNS). Cauquenes:
Fundo EI Roble, Colhueco, E Chillan, Xll - 1965 , 24 (AMNH); Pelluhue, 31-VU1985, F. Silva, 5 (MHNS), 18120-1X- 1985, F. Silva, 1 (MHNS). Caulfn: Cherquenco,
1954, I (MHNS) . Chacabuco: Caleu, ex Salix viriminalis, 10-111-1990. M. Eigueta,
I (MHNS). Chilo,,: Aucar, 6115-1-1952, L. E. Pena, 2 (MHNS), 15-1-1952, L. E.
Pena, 20 (MHNS). Choapa: Cerro La Silla del Gobernador, 3 1-X- 1988. E. Maury,
I (MLP); km 272 Panam . Norte, ex Senna cumming;; var. coquimbensis, 14- V- 1985,
M. Elgueta,4 (MHNS). 23-1-1986. M. Eigucta . I (MHNS), ex Adesmia microphylla.
II -lll - 1986, M. Eiguela, 1 (MHNS); Quilimari, 13-IX -1972, 1 (MHNS). Colchagua:
Valle del Nilahue, 1-11-1915,2 (MHNS). Concepci6n: Concepci6n, 26/28-Xll-1926,
F. & M. Edwards, 1 (BMNH), 20-111-1952, 1 (MHNS). Cordillera: EI Alfalfal, 23-U1967, V. Perez, 4 (MHNS), 25-1-1968; J. Moroni, 3 (MHNS). Curic6: EI Coigal,
Xl-1955, L E. Pena, 5 (CWOB). Elqui: La Mazquesa, 350 m, A. Aguilera coiL,
1996
THE WEEV IL GENUS RHYEPHENES
19
10
31
11
12
,r
17
,.
,.
l4
j
\
\
I
I
16
9
40
,,
15
f
"
42
~
41
44
46
47
48
4'
Figs. 29- 49. Rhyephenes spp.. female genitalia. 29 - 35. stern um 8. ve ntral view; 36--42.
hcmi stemite. ve ntral view : 43- 49. spcrmatheca . 29. 36. 43. R. clathram.s; 30. 37. 44, R. ga)';:
31. 38. 45. R. goureaui; 32. 39. 46. R. hume ralis : 33, 40. 47 . R. la teraJi.s: 34. 4 1, 48. R. maille;:
35 , 42. 49. R. squamiger.
V1I1- 1973,4 (MHNS); Pt a. Lengua de Vaca, 3 1-VU - 197 1, L. E. Peila, I (M HNS).
Los Andes: Estero Leiva, X- 1953, 5 (MHNS); Rio Blanco, 30-XII - 19 17, 2 (M HNS).
M alleco: Angol, 8-1X- 1949, Cerda , 6 (MHNS), 18-111 - 1953, I (MHNS); CuracautIn,
13-lll - 1952, 2 (MHNS). Melipill a: Cuesta de Barriga, 7- 1- 1952, I (MHNS); EI Cancio, 30-Xl - 1970 , G. Barri a. I (MHNS ). Petorca: Quebrada del Chi vato. 30-X- 1988.
E. Maury. I (MLP); Quebrada del Talanquen. 9-Xl I- 1984. H. Niemeyer. 3 (MHNS);
Q uebrada EI Ti gre, 8-X- 1988. E. Maury. 5 (MLP). Quillota: Palmas de Oco~. 27-X1988. E. Maury. I (MLP); Quebrada Escobares. 22- U- 1964. J . Soler vicens. 3
(MHNS ); Quillota. M . Frit z. I (MACN). San Fe lipe de Aconcagua: La 0 11 ita.
Cantillana. 2000 m. 18-XIl- 1969. L. E. Peila, I (MHNS). Santiago: Farellones. 2500
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOC I ET Y Vol. 104(1 - 2)
m, 4-XlI-1969, J. Valencia, I (CWOB ). 26-111-1971, J. Valencia, 2 (MHNS ); Oasis,
11- 1966, R. Perez, 17 (MHNS); Piche Alhue , 1800 m, 18-XlI - 1969, L. E. Pena. 3
(MHNS); San CrisI6bal, 14-XII- 1966, R. Perez, I (MHNS); San Jose de Maipo,
17-U- 1970, L. Alfaro, I (MHNS ). Talagante: Talagante, 7-1X-1970, M . Elgueta. 4
(MHNS). Talca; Vilches Alto, Cord. de Talca, 1- 1990, Arce. 5 (MHNS ). Valdivia:
La Uni 6n. Cocule, 6-111-1952, 2 (MHNS). Valparaiso; Marga Marga, 26-IV-1983,
A. Mann. 2 (MHNS). 3 (BMNH); Quillota. sobre nispero, I (MHNS); Quintero,
"bajo Ambrosia" , 29-Vlll-198I, I (MHNS); Valparaiso, XU - I925. A. Faz, 3
(B MNH ).
Rlzyep henes gOllreau; (Gay and Soher, 1839)
(Figs. 5,6, 19.20,3 1. 38, 45.51)
Physorhorus gourea u; Gay and Solier, 1839:26.
Rhyephenes gourea ,,;; Solier. 1839:L.
This spec ies could be confused with R. maille;, the other species lacking humeral
stripes in the e lytra; the latter has the prothorax wider than the c lytra, and the elytral
tubercles on intervals morc developed than those on striae.
Redescription: Habitus (Figs. 5, 6). Body clothed with scales. Prothorax slightly
g lobose, as wide as e lytra; disc with anterior impression , and large and deep punctures. Elytra with basal area strong ly impressed; with rounded tubercles; tubercles
o n intervals small , similar to those on striae. and partially fu sed to them; humeri
lacking stripes of white scales. Male. Rostrum lacking tubercles. Aedeagus (Figs.
t 9, 20) robust in lateral view; ostiolar sc lerites ro unded; parameres short , broad.
Female. RoslTUm with broad, abundant punctures. Sternum 8 (Fig. 31) with plate as
long as wide. and sides with setae; arms broad. Hemi sternile (Fig. 38). Spermatheca
(Fig. 45) with nodulu s developed and ramus not developed. Length 8.0- 15 .2 nun.
Material examined: ARGENTINA. Chubu\: EI Maiten, 28-XJI-1958. A . Kovacs,
14 (AMNH ). Neuquen: CatAn-Lil. 25-11- 1963. I (IADIZA); Lago Alumine. 25-111964, 4 (IADIZA). 14-1- 1993, G. Debandi, I (IADIZA); La Victoria , X- 1963, I
(lAD1ZA ); without more precise data , 1 (MLP) . C HILE. Without more precise data:
48 ( I AMNH , 2 MACN. 43 MHNS. 2 MLP). Araueo: P N. Contulmo, 2-XJ-1904.
I (CWOB ). Bio- Bio: Abanico, 19-1-1979. M . Elgueta. I (MHNS ); San Carlos de
Puren. 22-1- 1973, I (MHNS). Cautin: Cherquenco. 1954, L. E. Pena. 3 (MHNS ).
Chiloe: Ancud , 1O-IV- 19 14. I (AMNH ), 19-Xll-I 926, E & M . Edwards. I (BMNH);
Pivehue. 16-11- 1992. S. Roig-Junent . I (IADIZA). Curic6: Potrero Grande, 23-X1976. G. Arriagada, I (MHNS). Llanquihue: Puen o Monn. 3 (MLP). Malleeo: Cordillera Lonquimay, Sierra Nevada. 2- 11-1962. 1. Valencia. I (CWOB ); P. N. Nahuel
Buta, 1260 m, 23-Xll-1985, S. Roig-Junent , 5 (IADIZA). Nuble; Cabrena, 31-11977, F Rodriguez, I (MHNS). Valdivia: Huellelhue. 24-1-1973. M . EJgueta, I
(MHNS).
Rhyephenes Imm e ralis (G uerin , 1830)
(Figs. 7, 8, 21, 22, 32, 39, 46, 52)
Tylodes humeralis Guerin. 1830: 124.
CryplOrhynchus araclmodes Erichson. 1834:264.
1996
THE WEEVIL GENUS RHYEPHENES
"
Rhyephenes incas Gyllenhal, 1837 :3 14.
Rhyephenes aracJ1Ilodes; GyllcnhaJ, 1837:3 14 (=R. ;!leas); Gemminger & Harold,
1871 :2563 ( = R. humeralis); Berg, 1899: 154 (Luz6n, erroneous type locality).
Physothorus bayeri Gay and Solier. 1839:27 .
Rhyephenes boyeri; Sclier, 1839:L (= R. incas); Lacordaire, 1866: 108 (reinstated);
Kuschel , 1955 :287 (=R. humera/is).
Rhyepilenes humeralis ; Schoenherr, 1844:402 (=R. ;"cas); Gemminger & Harold,
1871 :2563 (reinslaled).
Rhyephenes inca Lacordaire, 1866: 108 (error noted by Berg , 1899: 154).
Rhyephenes aequalis F Philippi, 1899:90; Kuschel , 1993 :64 (=R. humeralis).
This widespread species is variable in length . Its diagnostic strongly globose and
protuberant prothorax can be less notable in so me of the smaller specimens.
Redescription: Habitus (Figs. 7. 8). Body clothed with scales. Prothorax strongly
g lobose, protuberant, wider than elytra; disc lacking anterior impression , and large
and deep punctures. Elytra with basal area strongly impressed; with subconical tubercles; tubercles on intervals large. more developed than to those on striae, and
partially fused to them; humeri with three short stripes of white scales. Male. Rostrum with tubercles on dorsum and sides. Aedeagus (Figs. 21, 22) robust in lateral
view; ostiolar sc lerites rounded; parameres short, broad. Female. Rostrum with fine
punctures. Sternum 8 (Fig. 32) with plate longer than wide, and sides with setae;
arms narrow. Hemisternite (Fig. 39). Spermatheca (Fig. 46) with nodulu s and ramus
not developed. Length 5.5-6.7 mm.
Type material: Lectotype male (here designated): [LECTOTYPFJRhyephenesl
aequa/islF. Philippi] [= Rhyephe"eslhumeralisl(G uerin)1 del. G. Ku schel/ 1990]
(MHNS).
Other material examined: ARGENTINA. Without more precise data: 10 (9
BMNH. I MACN) . Mendoza: without more precise data, C. Bruch. I (MLP). Neuquen: without more precise data, I (MLP). Rio Negro: Bariloche, 161l8-Xl-1926.
F. & M . Edwards, 4 (BMNH). CHILE. Aisen: Puerto Cisnes, ll - 1961 , L. E. Pena, 6
(BMNH). Arauco: Arauco, 16-Xll-1985, S. Roig-Junent, 2 (IADIZA), Baier, 5
(MACN); Temuco, 1- 1975, S. Roig-Junent , 2 (IADIZA), 2 (MLP). Bio-Bfo: Pemehue, al night, 1-11- 1968, L. & C. W. O'Brien, 6 (CWO B), 20-1-1993, G. Debandi, I
(lADIZA). Cachapoal: Mollo., Xl - 1967, I (CWOB); Rengo, 11- 199 1,4 (MLP). Cardenal Cam: Rosario, 23-ll-1967, J. Numhauser, 2 (BMNH). Cautin: without more
precise data, L. E. Pena, 5 (CWOB). Chacabuco: Caleu, ex Salix vitiminalis,
10-111-1990, M . Elgueta, I (MHNS). Chiloe: 25 !un S Castro, V-1968, L. & c. W.
O'Brien, I (CWO B); 9 !un E Chepu, 4-11- 1968, L. & c. W. O'Brien, 2 (CWOB ).
Choapa: I km S Huentelauquen, C. W. & L. O'Brien, I (CWOB); Los Vilos, 18-1X1969, A. Tobar, 13 (CWOB); Palo Colorado, N Quilimari, 27-X - 1991 , M . Elgueta,
2 (MHNS). Coihaique: Coihaique, 8-111-1972, A. Tobar, 2 (CWOB). Concepci6n:
Concepci6n, 25-VIII-1968, J. Apablaza & C. W. 0' Brien, 2 (CWO B), 21-I11-1970,
E. Ho lsten, I (MHNS) , E. Reed, I (BMNH); 40 km E Concepci6n, 25-VIII - 1968,
J . Apablaza & c. W. O'Brien, I (CWOB). Cordillera: San Jose de Maipo, 1-1969,
R. Dulovitch, I (MLP). Curic6: EI Coigo, cordillera Curic6, X- 1964, 7 (AMNH ),
X-1974, 8 (AMNH); EI Coiquel, Xl-1955, L. E. Pena, 8 (CWOB); Quebrada Mala,
32 Km E Molina, 2-Xl-1991, M . Elguela, I (MHNS). Linares: 3 !un SE Agua de
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY
Vol. 104(1 - 2)
La Gloria. 4-XI- 1967. 5 (CWOB); 13 km SE Agua de la Gloria, 4-X I- 1967. L. &
C. W. O'Brien, 4 (CWOB). 24-V IU - 1968, L. & C. W. O'Brien, I (CW OB); La
Pasarela, 28 km de Linares, 1- 1984, S. Roig-Junen\, 2 (IA DlZA); La Vega, 6 km S
Parral. 8-XI-1991, M . ElgueJa. I (MHNS); Panamavida. 20-X- 1968. D. Correa. 2
(CWOB ); Romehual , cordi ll era Parral. 5/10-X I- 1960, L. E. Peiia. I (IPCN). Los
Andes: Los Andes. 22-V-1979. Gordon. 2 (CWOB ); Rio Blanco, 25-1- 1974. L. E.
Pena. 2 (M HNS). Malleco: Angol, 3-lX- I927. E. Reed. I (AMNH). 22- IV-1946. 2
(A MNH ). 12-X-1946. S. Perez. 2 (AMNH), 20-V II - 1947, W. Mason. 2 (CWOB ); 6
km W Angol, al nighl , 2-X I- 1967, 5 (CWOB); cordi llera Lonquimay. 4-1- 1962, I
(CWOB); P. N. Nahuel BUla. 38 km W Angol. 4 ,300', I 2- U- I 968. L. & C. W.
O·Brien. I (CWOB); Pichinahuel. Cordi llera Nahuel Buta. 1/10-1- 1959. L. E. Peiia.
I (BMNH). Nuble: Fundo Las Cruces. Cordi llera Parral, V- 1958, L. E. Pena, 6
(BMNH). I2I14-X II - 196O, L. E. Peiia, I ( BMNH); Las Trancas, 8 km W Termas de
ChillAn. 1300 m, 15-X I- 1981. R. T Schuh & N. I. Plalnick., I (A MNH); 15 km SE
Recinlo. al night. I-XI - 1967. L. & c. W. O·Brien. 3 (CWOB). Petorca: Cachagua,
19-1X-1971, M. Elguela. I (M HNS ); 7 km NE Papudo, 8-V IIJ - 1968. L. & c. W.
O'Brien, 2 (CWOB ); 13 km S Quinquimo. 12-Xl- 1967, L. & c. W. O'Brien, I
(CWOB); 10 km SE Zapallar. 11 -11- 1967. L. & C. W. O·Brien. 15 (CWOB). 23-1X1967, C. W. O'Brien, I (C WO B). Q uillola: Limache, Cerro de la Cruz, 1975 . S.
ROig-Junenl , I (IAD IZA); Cerro Macaya, 18-X I- 1962, A. Tobar, I (CWOB); 19 km
E Manzanar. 3-Xl- 1967 , L. & C. W. O'Brien. I (CWOB). San An lonio: anles de
Los Quci'ies, 15-1- 1984. S . Roig-Junent . 2 (lAD IZA). San Felipe de Aconcagua: 15
km SE LJay-LJay, 18-IX-1967. L. & c. W. O'Brien. 3 (CWOB). Sanliago: 15 km E
ArrayAn , al nighl, C. W. O'Brien, 2 (CWOB); Cerro Robles. 100' . at nighl,
24-XII- 1968, L. & C. W. O'Brien, 6 (C WO B). 6300'. 24-X II -1967. L. & c. W.
O'Brien, I (CWOB); Curacavf. 14-X- 1967. L. & c. W. O'Brie n, I (CWOB ),23-Xl1967, C. W. O'Brien, I (C WO B); EI Cancio. 5-X- 1963, J. Val enc ia. 2 (CWO B),
I 9-X- I 963, J. Valencia, 4 (CWO B), 2-X I- I 963. J. Valencia, I (C WO B), 23-111 - I 968,
D. Correa, 2 (CWOB ); EI Manzano, 23-X II - 1967. D. Correa, 4 (CWOB), 6-111 - 1968,
D. Correa. I (CWOB), 4-X- 1970, I (MHNS); 2 km E EI Manzano, 1-11- 1969. D.
Correa, I (CWOB); EI Saho. 20-XIl - 1967. D. Correa. I (C WOB); Farellones. 8,600',
under dung. 25-X II - 1968. L. & c. W. O'Brien. 7 (CWOB ), 2500 m. 4-XIJ-1969. J .
Valencia. 2 (CWOB ): La Reina. lOl l 5-X I- I 967. J. Barros, I (C WOB), 6-XII - I 967,
C. Reyes, I (CWOB); Las Condes, 6-111- 1938, 4 (B MNH); Las Cruces. 3- 111 - 1974,
E. Arriagada, 10 (CWOB); 5 km S Melipilla. al nighl , 16-IX-1967. L. & c. W.
O'Brien,7 (CWOB); I I km S Melipilla, 300'. al nighl. 16- IX-1967. J . Apab laza &
C . W. O'Brien, 5 (CWOB) ; 8 km W Padre Hunado , 3-lX- 1967, L. & c. W. O'Brien,
2 (CWOB ); 15 km W Padre Hunado, al nig hl. 6- IX- 1969, L. & c. W. O'Brien, 4
(eWOS); P. N. La Campana. 14- 111- 1985. C . Vivan. 1 ( MHNS); Rocas 5 10 . Domin-
go, on lupine. 23-X I- 1968. L. & c. W. O·Brien. 103 (C WOB); 4 km W Rung ue.
6-lX-1968. L. & C. W. O'Brien, I (CWOB). Talagante: Nahagua, 10-11- 198 1. J. E.
Barriga. 7 (MLP). Talca: Alt os de ViJches. 30-X-1969. J . Rozen & L. E. Peiia. 2
(AMNH ), 27-XII - 1969, J. Valencia, 2 (CWO B). 10-X- 1970, A. Tobar. 4 (CWO B),
13-X- 1970, A. Tobar. I (C WO B), 1280 m, 10/1 I-XI- 197 1, J. Valencia, 3 (CWO B).
1200 m. 517-X- 1972, Coscar6n, Pena & Wygodzi nsky, 7 (AMNH ); Consritu ci6n,
22-V-1969, E. Holsten. 3 (AMN H), 13-V- 1970. E. Ho lsten. 2 (MH NS); 7 km W
1996
THE WEEV IL GENUS RHY£PH£N£S
13
Molina, 1-1968, C. W. O'Brien, 10 (C WOB ), 20-1-1968, D. Correa, I (CWOB ), I-U1968, D. Correa, 30 (C WOB ), 25- U- 1968, o n Salix trunk, D. Correa. 6 (C WOB ).
26-XlI - 1968, D. Correa. 6 (CWOB ); Valle Empedrado. 6-X-1970. E. Holsten. I
(AM NH ); Vilches Allo. 14-XlI - 1973. 2 (MHNS). 2 1-XlI - 1973, I (MHNS). Valmvia:
Valdivia, D. Sharp, 2 (BMNH). Valparaiso: A lgarrobo, on Baccharis, 24-XI- 1968,
L. & c. W. O'Brien, I (CWOB ), IX - 1969. I (MHNS); 15 km NW Casablanca,
1000', at night, 9- IX-1967, L. & C. W. O'Brien, I (C WOB); EI Bato, E lI\ apel ,
25-X-1991, M. Elgueta,4 (MHNS ): EI Bato, a farm east of llI apel, 19-X-1966, I
(AMNH); La Caneta, 29-X- 199 1, M . Elgueta , I (MHNS); Valparaiso. 1903 , Crawfo rd expedition, I (BMNH), Walker, I (B MNH ), E. Reed , 2 (AMNH ), 3 (B MNH );
19 km SE Villa Alemana, 1.000', at night , 1X- 1967, L. & c. W. O ' Brien, 3 (CWOB );
Vina del Mar, 17- IlI - 1938. I (B MNH); 13 km S Vina del Mar, 650'. 3-1X- 1967, J.
Barros. 1 (eWOS ). Without more precise data: Araucanfa, R . M . Middleton , 13
(BMNH); A. Tobar, 6 (CWOB ); Reed , 3 (B MNH); C. Darwin, I (B MNH ); 11 6 (2
AMNH, 19 BMNH, 2 1 MACN, 70 MHNS . 4 MLP).
RhyeplreIJes laleralis (Guerin, 1830)
(Figs. 9, 10.23, 24. 33, 40, 47, 52)
Tylodes lateralis Gu~rin. 1830: 126.
Rhyephenes latera/is; Schoenherr, 1844:404.
This rather scarce species is eas il y recognized by the two short and one longer
e lytral humeral stripes.
Redescription: Habitus (Figs. 9. 10). Body clolhed with scales. Prothorax S)jghll y
globose, as wide as elytra. wider than elytra; disc lacking anterior impression. and
large and deep punctures. Elytra wit h basal area slightl y impressed; with rounded
tubercles; tubercles o n intervals small . similar to those on stri ae, and partially fused
to them; humeri with three stripes of white scales. two short and o nc longer. Male.
Rostrum with tubercles on dorsum and sides. Aedeagus (Figs. 23. 24) s lender in
lat eral view; ostio lar sclcritcs rounded; paramcres long. narrow. Female. Rostrum
with fine, sparse punctures. Sternum 8 (Fig. 33) wi th plate as lo ng as wide . and sides
lacki ng setae; arms narrow. Hemisternite (Fig . 40). Spennatheca (Fig. 47) with nodulus and ramus not developed. Length 9.1 - 13.2 mm.
Material examined: C HIL E. Without more precise data: 3 (MHNS). Nuble: Cobquecura, 13-1- 1967 , P. Ram;rez. I (C WOB): Quirihue, 1- 1979, C. Vidal , 3 (MHNS).
Talca: Alto Vilches, 14- UI - 197 1, L. E. Pena, 2 (MHNS), 2-X1- 1973, I (MHNS),
21-X1 1- 1973, 2 (MHNS ); Constituci6n : 27- U- 1948, 2 (MHNS), 25-1X- 1970. E. Ho lsten, 5 (AMNH ). Valmvia: Panguipulli , 14-U- 1966, I (MHNS).
Rhyephell es maille; (Gay and Solier, 1839)
(Figs. II , 12. 25, 26, 34. 41, 48, 53)
Pltysotltorus maille; Gay and Solicr. 1839:24.
Physolho rus laeviroslris Gay and So li er, 1839:25 (probably synonym of P. maillei).
Rhyepltenes maille;; Solier, 1839:L; Angu lo, 1970:3 13 (larva).
Rhyepltenes laevirostr;s; Solicr, 1839 :L; Lacordaire, 1866: 108 (not synonym of R.
maillei); Berg, 1899: 153 (= R. maillei); F. Philippi, 1899:82 (= R. maillei); Hustache. 1936:235 (rein stated); Morrone, 1994:95 (= R. maillei) .
14
JO URNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2)
Rhyephefles immaeu/alus Blanchard, 1853:251 , pI. 14, Fig. 16 (= R. laevirostris);
Berg, 1899: 153 (=R. moillei); Hu stache, 1936:235 (= R. laeviroslris): M orrone.
1994:95 (=R. maillei).
Rhyephenes sulcalUs F. Philippi, 1899:86; Morrone. 1994:95 (= R. mail/ei).
Rhyepi1elles philippi; Fiedler, 1942:279; Kuschel, 1950: 18 (=R. sulcatus); M orrone,
1994:95 (= R. maillei).
This widespread species is variab le in len gth. The names R. laevirostris. R. imand R. philipp;; have been applied to variants of this species.
Redescription: Habitus (Figs. 11, 12), Body clothed with scales. Prothorax. slightl y
maeu/allls, R. su/calUs.
globose, wider than elytra; disc lacking anterior impression, and large and deep
punctures. Elytra with basal area strong ly impressed; with subconical tuberc les; tu-
bercles on interval s large. more developed than to those on stri ae; humeri lacking
stripes of white scales. M ale. Rostrum with tubercles on dorsum and sides. Aedeagus
(Figs. 25. 26) slender in lateral view; ostiolar sclerites rounded; parameres long,
narrow. Female. Rostrum wi th fine punctures. Sternum 8 (Fig. 34) wi th plate longer
than wide, and sides with setae; arms narrow. Hemjstemite (Fig. 41). Spermatheca
(Fig. 48) with nodulus developed and ramus not developed. Length 6.3- 15.4 mm.
Type material: Lectotype male (here designated): [LECTOTYPEJRhyephenes/
sulcatus!F. Philippil fRhyepheneslmailleilvarianl Gay & Solier/del. G. Ku scheUI990J
(MHNS).
Other material examined: ARGENT INA . Chubut: EI Hoyo, 5-V-1959, A . Kovacs.
4 (BMNH), 18-X- 1960. A. Kovacs, I (BMN H); Lago Verde, 560 m , 4-11- 1983, M .
& P. Gen tili, I ([PCN); Rio Correntoso. 1 (MACN); withou t more precise data. 2
(MLP). Mendoza: without more precise data, I (A MNH). Neuquen: Alumine, 25-11 1964, I ( IADlZA), ll - 1977, O. de Ferrari s, 3 (IPCN), 8- 1- 199 1, G. Debandi, 2 (lA DIZA); Chapelco. 7- 111 - 1964, 2 ( IADIZA), 1,700 m, 15-111- 1964, M . Gentili , 3
(IPCN), 1750 m, Xll - 1965, M . Gentili. I (IPCN), 1400 m, 30-111- 1982, M. Gentili,
I (IPCN); Copahue, 1,925 m, 18-Xll- 1963, M. Genli li, I (IPCN), 2000 m, 3-11- 1987,
M . Gentili. I (!PCN); Huiliches, 13-XlI-199I , G. Debandi, I (IADlZA ); Kilca,
4- 1- 1972, I (IPCN); Lago Hennoso, Xll-1949, M . Barrera, 3 (MACN); Lago LaCM,
23 km W San Martin de los Andes , 25- 1-1972, L. Hennan, I (CWO B); Lago LaCM,
Nonthue, 640 m, 17-X I- 1992, M. Genti li . 2 (IPCN); Lago Queni, 875 m, M . Gentili,
I (IPCN); Lago Tromen, 1,000 m, 15-Xll-1962. M . Gentili, I (lPCN); Pucara, 1949.
S . Schajovskoi, 2 (MACN), 23-XI- 1957, M. Gentili, I (IPCN), 21-Xll-1965, L.
Grosso, 3 (MLP), 6-11- 1972, L. Herman, I (CWOB); Rio Alumino, 16-XIl- 1965, M .
Gentili, 1 (lpeN); road between Pucara and Lago Venado, 24125-1- 1972. L. Herman,
2 (CWOB ); San Martin de los Andes. 4-W - 1942, M. BirabCn, I (MLP), 17-llJ - 1964,
I (lADlZA), 9-XlI-1976, O. de Ferraris, I (lPCN), 1000 m, 2-Xl-1986, M . & P.
Gentili, I (IPCN), 640 m , M. Genti li , 5 (lPCN), 1O-Xl-1991. G. Debandi, 2 (IADIZA); valle del Rio Limay, n - 1947, J. Navas. 3 (MLP); without more precise data,
28 (24 MACN, 4 MLP). Rio Negro: Bariloche, 25128-X-1926, F & M. Edwards, 2
(BMNH), 11-1938, Bi.rabCn & Scott, I (MLP), 1- 1942, Rossi , 3 (MACN), 26-XI1963, A. Kovacs. I (A MNH), Richler, 5 (MLP); EI Bols60, 20-LX-1956, A. Kovacs,
2 (BMNH). 12-VTl-1958, A. Kovacs, I (BMNH), 25-Xll- 1958, A. Kovacs, 16
(BMNH), 18-XlI-1959, A. Kovacs, I (BMNH), 22-X!I-1959, A. Kovacs, I ( BMNH),
29-Xl-1959. A. Kovacs, I (BMNH), 5-X-1960, A. Kovacs , 6 (BMNH), 18-1- 1961 ,
1996
THE WEEVIL GENUS RHYEPHt:NES
15
I (BMNH), IO-VUJ - 1961. A. Kovacs, I (BMNH), 20-X- 1961, 2 (BMNH), I-X1I1961, A. Kovacs, 4 (BMNH), 9-XU - 1962, A. Kovacs, I (BMNH), 22-Xl I- 1962, A .
Kovacs, I (BMNH), 16-1 - 1963, A. Kovacs , I (BMNH), 19-1- 1965, A . Kovacs, II
(AMNH); isla Victoria, 15-XI-1969, I (MACN), 2-XlI- 1969, I (MACN); isla Victoria, IV-1958, 2 (IADlZA); Norquinco, IO-U- 1964, A. Kovacs , I (AMNH); P. N .
Los Alerces. S. Roig-Jufient, I (lADlZA); Pto. Radal , isla Victoria, I (MACN) . Santa
Cruz: Lago Argentino , 1959, P. W. James , 3 (BMNH), 16127-11- 1974, C. Bord6n, I
(C WOB); valle del Lago Blanco, 19m, 2 (BMNH); Valle Tunel, 3 (MLP). Tierra
del Fuego: Without more precise data, 1900. C. Bruch, 6 (MACN). Without more
precise data: Patagonia, 69 (MLP); 1935, 5 (BMNH), 1- 1972, L. Herman, I (CWOB ).
CHILE. Without more precise dala: 50 (5 AMNH, 12 BMNH, 5 MACN, 26 MHNS ,
2 MLP). Ai sen: Lago J. A. Rios, Tailao, 8-XII-1956 , C. Riffart, I (MHNS); Plo.
Aisen, 12-1II-1943, R. Maldonado, 39 (MLP), 21 -1- 1968, L. & c. W. O ' Brien, I
(C WOB); Taitao, Base no. 5, 5-XlI- 1956, C. Biuart, 2 (MHNS); withoul more precise data, 11 - 1934, I (MHNS). Arauco: Arauco, 24- IJJ - 1969, G. Billings, 7 (MHNS);
C anete, 24-ill-1969, R. Billings, 6 (MHNS), pi no insigne, 12-V-1969, D. Edelman,
I (AMNH); Caram~vida, 31 -XII -1957, L. E. Peila, I (MHNS), 18-X-1969, G. Barria,
I (MHNS), 16-XII- 1985, S. Roig-Junent, 2 (IADlZA); Curanilahue, I -IV- 1970, D.
Edelman, I (AMNH); is la Mocha, Xl - 1982, Bullock, 3 (MHNS); P. N . Contulmo,
19-X- 1969, G . Barria, I (MHNS), 15-XU-1985, S. Roig-Juilenl , 5 (IADlZA); withoul
more precise data, Baier, I (MACN). Bio-Bio: Mulchen, 16-11 - 1969, F. Rojas , 1
(MHNS). Capil~n Pratl: Dos Lagunas, night, 13-1- 1968, C. w. & L. O'Brien, I
(CWOB), under wood, 14-1-1968, C. W. & L. O' Brien, 5 (CWOB ). Cautin: rio
Cautln , L. E. Peila, 2 (CWOB); 50 km S Temuco, II-X1I-1984, S. Roig-Junenl , I
(lADlZA); Villarica, 3-11-1968, B. Heineman, I (AMNH). Chiloe: Aucar, 18-1- 1952,
L. E. Peila, 7 (MHNS), 6-V- 1952, L. E. Peila, 2 (MHNS); 9 km N Castro, 7-11-1968,
L. & C. W. O'Brien, I (CWOB); 14 Ian E Chepu, 4-11- 1968, L. & C. W. O ' Brien,
3 (C WOB); Daicahue , 18-1- 1962, R, Usinger, I (CWOB); 20 km N Pichue, on
[Nothojag us j nitida, 16-Xl - 1992, S. Roig-Junenl. I (IADlZA); 6 km N Quell6n, 6-11 1968, L. & c. W. O ' Brien, I (CWOB ); 21 km N Quell6n, 6- lJ - 1968, L. & c. W.
O' Brie n. t (eWOS); without more precise data, C. Darwin. I (BMNH). Coihaique:
Coihaique, 1-1934, 9 (MHNS); 17 km SE Coihaique, 20-1- 1968, L. & c. W. O'Brien,
4 (CWOB); 56 Ian SE Coihaique, 20-1- 1968, L. & C . W. O ' Brien, 1 (C WOB); 64
Ian SE Coihaique, 20- 1-1968, L. & C. W. O ' Brien, 12 (CWOB); 7 km W Coihaique,
at nighl , 21-1-1968, L. & c. W. O ' Brien, I (C WOB ); 33 km W Coihaique, 23-11968, L. & c. W. O'Brien, 4 (CWOB ). Coichagua: 10 km S San Fernando, 12- Xl 1967, J . Undurraga, I (CWOB). Concepci6n: Concepci6n, 21-UJ-1969, E, Holstem,
I (AMNH ). L1anquihue: EI Manso, 22-1- 1959, A. Kovacs, 13 (BMNH), 1-IV-1960,
A . Kovacs, 52 (BMNH); Frutillar Bajo, 11- 1965, G . Silva, 2 (MHNS); Plo. Varas,
M . Richter, 2 (MACN). Magallanes: Cabo Negro, 28-XII - 1975 , Perez, 6 (MHNS):
cordillera Lonquimay. Sierra Nevada, 2-1- 1962. J. Valencia , I (eWOS), 4-1-1962 , 1
(CWOB ); Dawson island, 1916, I (BMNH); EI Canelo, 15-ill- 1969, L. E. Pena, 3
(MHNS ); Estancia Cane/o, 17-1-1968, L. & C. W. O'Brien, 12 (CWOB ); 26 km E
Estancia Canelo, on and under Nothofagus, 17-1-1968, L. & C. W. O'Brien , 37
(CWOB ); is la Navarino, 1935, J. Bird, 1 (AMN H); Laguna Amarga, 10-1- 1968, C.
W. & L. O'Brien, I (CWOB); Magallanes, 2-IV- 1953, T. Cekalovic, I (MHNS);
Punta Arenas , Walker, 2 (BMNH); 3 km W Punta Arenas, al nighl, 16-1- 1968, L. &
16
JOURNAL OF THE NEW YORK ENTOMO LOG IC AL SOCI ETY Vol. 104(1 -2)
c.
W. O'Brien, I (CWOB ). Malleeo: lealma, 11 - 1989. I (MHNS ); Pailahueque.
X- 1962. M . Frit z. 3 (MACN); P. N. Nahuel Buta. 1200 m. 23-XII - 1985. S. RoigJunent. 3 (lADIZA). 20-1- 1993. G . Debandi, 2 (IAD IZA). 21-1- 1993. G. Debandi . I
( IAD lZA); P. N. Tolhuaea, 13-1- I 993, G. Debandi, 2 (IADIZA); PiehinahueI. Nahuelbuta , 1200 m. 14-11- 1956. L. E. Peiia, I (CWOB ). Osom o: Antillanea, 3,700'.
Notha!aglls plimilia, 10-1l- 1968, C. W. O ' Brien , I (C WOB ); 18 km NW Antill anea.
1,800', at night , 9-11- 1968, L. & c. W. O ' Brien, 3 (C WOB ); Bahia Mansa. 26- IX 1968. C. Calder6n, 3 (MHNS); Osorno, 1- I 975. S. Roig-Juiient , 9 (IADIZA), X II 1984, S. Ro ig-Junent. I (MAC N); 5 I km E Osomo, Lago Puyehue. 800'. 9-11-1 968,
L. & C. W. O ' Brien, 8 (CWOB ); P. N. Pu yehue. 11-1 992, S. Roig-Juiient. I (IAD IZA); P. N. Puyehue, Agua Ca liente, XlI - 1984. S. Ro ig-Juiienl. I (LADI ZA); Puea'rihue. coastal to wn , 312 1-lLI- 1967, 2 (AMNH); Purranque, LX - 1985, J. E. Barriga,
I (MHNS). Santiago: Lago Array"', 20-XII - 1968, C. Barrientos. I (CWOB ). Talagante: Maipo, 7-X I- 1968, J. Marshall. I (CWOB). Talca: Alto de Vilches, 10·X1970, A . Tobar, I (CWOB). Tierra del Fuego: forest S of Bahia lml'i l (Useless Bay).
XII - I904. R. Crawshay. I (BMNH); rio MeKlelland, XUXII-1904, R. Crawshay. I
( BMNH). Ultima Esperanza: Caneha Carreras. 24- 1- 1988, J. J. Morrone, I (MLP);
Cerro Guido, Notha!aglls, 10-1- 1968, C. W. & L. O'Brien, 4 (CWOB ); Cerro Lc6n.
19-1- 1969. v. Perez. I (MHNS); P. N. Torres del Paine. 25-1- 1988. J . J. Morrone . I
(MLP). Valdivia: Laneo, 1-1 980. 4 (MLP); La Uni6n, 10 -lli - 1952, I (MHNS ); Panguipuli, 14-11- 1966, G. Ol1ega, 20 (C WOB), 11- 1984, L. E. Peiia. I (CWOB ); Pirehueieo, I-XII - 1985, S. Roig·Junent , I (lADI ZA); Valdivia, 30·XII-1968, D. Correa,
I (CWOB ), 3 1-XII - 1968, ni ght , D. Correa. 9 (C WO B), 3-1- 1969, D. Correa. 56
(C WOB ), 29-XlJ - I 986, E. Krammer. 2 (MHNS); 13 km SE Valdivia, 3- 11 - I 968. L.
& C . W. 0' Brien. I (CWOB ); 18 km N Valdivia. 4-1- 1969, D. Correa, II (CWOB ).
Rhyephell es squamiger F. Philippi. J899. reinstated name
(Figs. 13, 14, 27, 28,35. 42, 49.53)
Rhyephenes squamiger F. Philippi, 1899:88; Ku schel, 1993:63 ( = R. gayi).
This scarce spec ies. eas il y recogn ized by its body clothed with scales. has a rcIictual distribulion in central Chile. Ku schel (1993) incorrectl y considers lhis spec ies
as a geographkal race of R. gay;; however, the cladi stic anal ysis indicates that they
arc two different (and unrelated) species .
Redescription: Habi tus (Figs. 13. 14). Body lacki ng scales. Prolhorax sli ght ly globose, wider than e lytra; disc lacking anterior impression, and small and shallow
punctures. Elytra with basal area slightl y impressed; with subcon ical tubercles; tubercles on interva ls small , similar to those on striae. and partially fu sed to them;
humeri with two shon stri pes of white scales. Male. Rostrum lacki ng tubercl es.
Aedeagu s (Figs. 27, 28) slender in lateral view; ostiolar sclerites rounded; parameres
lo ng. narrow. Female. Rostrum with broad, abundant punctures. Sternum 8 (Fig. 35)
w ith plate as long as wide . and sides with setae; arms narrow. Hemistemite (Fig.
42). Spennatheca (Fig. 49) with nodulus and ramus developed. Length 9. 1- 10.9 mm .
Type material: Lectotype male (here designated): (883/aJ [squamigerl (= Rhyephenes/gayi/(G ue rin )/det . G. Kuschell 1990) rLECTOTYP FlRhyephenes/squlImigerlF.
Philippii (MHNS).
Other material examined: CHlL E. Choapa: Los Vilos, "ex Senna cumming i".
1996
THE WEEV IL GENUS RHYEPH ENES
17
50
52
Figs. 50- 53 . Rhyep /le rles s pp .. geographical di stribution . 50. R. c/athraw.\' (black circles)
and R . gay; (ope n circles): 51. R . gOllrealfi: 52. R. Iwmeralis (black circles) and R. ialeralis
(open ci rcles): 53. R. mllillei (black c irc les) and R. sqllamiger (open circles).
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1 - 2)
18
,-- roo.
••
, ,
R. squamiger
, , ",
10
,,
,,
"
11
18 20
1
~ ~
R. lateralis
,
, "I , " " "
,
~ '.' '.2 V 2.'
~~~~~;
1
,
1
0
,
1
"
0
I' '1
~ ~ ~
1f ';
~ ~
~ ~
':
i ; ;
(, i
R. clatbratu s
0
R.goureau;
R . gayl
,
, , • , 1 , "2 ,
2 , , , I
, 'l, 'i,
o 0 0
10
11
Fig. 54 .
R. bumeralis
R.maiUei
Cladogram of the species of Rllyephe"es. Black squares = synapomorphies: dotted
square::. = homoplasics.
11-1989. Barriga. 1 (MHNS); Nague. Los ViI os. 17-IX- 1971. L. E. Pena. 1 (MHNS);
km 27 Panamericana None, ex Bahia ambrosioides. 24-lX-1985, I (MHNS). Limari:
Fray Jorge, al nighl , 3-X-1967, L. & c. W. O ' Brien, 1 (CWOB). 20-VI-1968, L. &
C. W. O'Brien, 1 (CWOB), 21 -VI - 1968, L. & c. W. O ' Brien, 2 (CWOB), 7-X-1975,
C. Vidal , 1 (MHNS).
CLADISTICS AND BIOGEOGRAPI-IY
The analysis of the data matrix (Table 1) produced four c1adograrns (el = 0.55.
R1 = 0.51, length 43 steps) under equal weights. which after successive weighting
resulted in one c1adogram (e l = 0.82, R1 = 0.85, length III steps). In the c1adogram
(Fig. 54), the following phylogenetic sequence results: (R. sqllamiger, (R. lmeralis.
«R. cialhrallls, R. gOllreauj). (R. gay;. (R. humeraUs, R. maillei))))).
When the cladogram is compared with the distributional maps of the species of
Rhyephenes (Figs. 50-53), basal species (R. squamiger and R. latera lis) appear 10
be endemic to centra l Chi le, whereas the most apomorphic species are widespread
in cenlral and southern Chi le (R. gayi) or ex tend also to sou thern Argentina (R.
Iwmeralis and R. mailJei). Whether this pattern reHects a series of vicari ant or dispersal events cannot be deduced directly from the cladogram sequence. Future comparisons with other plant and animal taxa dlstributed in the same areas will help to
solve this problem.
ACKNOWLEDGMENTS
I thank Adrian Fortino for inking the drawings and Federico Ocampo for taking the photographs . This study was supponcd by gra nt 4662- 91 of the National Gcographic Society and
the Conscjo Nacional dc In vestigaciones Cie ntifi cas y Tccnicas (CONICET). Argentina .
1996
THE WEEVIL GENUS RHYEPHENES
19
LITERATURE C ITED
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Received 24 June 1996; accepted 27 February 1997.