Rev. Biol. Trop. ,45(2); 887-904, 1997

Systenotelus, a remarkable new of weevil (Coleoptera: ) associated witb (Cyclantbaceae) in Costa Rica and Panamá

Robert S. Anderson1 and Luis D. G6mez-p.2 IResearch Division, Canadian Museum of Nature, P.O. Box, 3443, Station D,Ottawa, ON. KIP 6P4, Canada 2 Las Cruces Biological Station,Organization for Tropical Studies, lnc. Apdo. 73-8257, San Vito, Coto Brus, Costa Rica

(Received l-V-1996. Corrected 15-VII-1996. Accepted 2-X-1996.)

Abstract: Systenotelus Anderson & Gómez, new genus, is described with three included species, S. carludovicae Anderson & Gómez, new species (Costa Rica), S. costaricensis Anderson & Gómez, new species (Costa Rica and Panama), and S. stockwelli Anderson & Gómez, new species (Panama). Adults of Systenotelusare distinguished by very elongate-narrow genitalia in females, female pygydium constricted near the midlength, and tumescent mesostemal process. They are relatively large in body size (4.9-12.4 mm), are pale to dad: yellow in color and possess various black dorsal and lateral maculations. Adults are remarkable in that the apex oC the abdomen is moderately to markedly tapered in females and the genitalia and associated structures of both sexes are modified and very elongate-narrow. Adults of S. carludovicae have been collected on freshly opened inflorescences of Ruíz & Pavón and C. drudei Masters () in Costa Rica; adults of S. costaricensis have been collected on inflorescences of Carludovica drudei in Panama; and, adults of S. stockwelli have been collected on inflorescences of Carludovica palmatain Panama. Systenotelus is a member of the tribe of the (sensu Kuschel), many other species of which attend flowers of various Cyclanthaceae and . The closest relative of Systenotelus appears to be an undescribed monotypic genus associated with Sphaeradenia hamata Harl.(Cyclanthaceae) in .

Key words: , Coleoptera, Curculioninae, Derelomini, Systenotelus, Cyclanthaceae, Neotropical.

The floral biology of Cyclanthaceae was the monotypic genus ) and the first studied by Drude (1877) under cultivated Carludovicoideae, with the remainder of the conditions. Harling (1958) confirmed many of species. Beach (1982) and Seres and Ramírez these early findings through field studies, (1995) reported that observed association of weevils with Poiteau is attended and pollinated by scarab Cyclanthaceae, found their larvae in dissected beetIes of the genus Cyclocephala in Costa inflorescences, and assumed their adults to be Rica and respectively. However, one the primary pollinators of these . Recent of us (LOO) has found Cyclocephala to be publications concerning the poIlination of the rather infrequent in the inflorescences of severaI Cyclanthaceae confmn them to be hundred populations of Carludovica aOO cantharophilous (Beach 1982; Gottsberger Cyclanthus in SW Costa Rica, from sea level 1991; Eriksson 1994; Seres and Ramírez to 2000 m elevation. 1995). In the Carludovicoideae on the other hand, Two subfamilies constitute this weevils (Curculionidae) are the primary family, the Cyclanthoideae (representedonIy by poIlinators. Gottsberger (1991) in Perú, 888 REVISTA DE BIOLOGIA TROPICAL reported that weevils of the tribe Derelomini reclassification of Curculionoidea by Kuschel are the pol1inators of two species, (1995) recognízes only six subfamilies of funifer (Poit.) Lindman and Curculionidae with most traditional Carludovica palmata Ruiz & Pavón. In subfamilies of Curculionidae relegated to tribal Venezuela, Seres and Ramírez (1995) also status in Curculioninae. This classification record derelomine weevils as the primary would appear to recognize Derelomini as a tribe pol1inators of fom Asplundia species and within Curculioninae although this is not Evodianthus funifer. Similarly, Erikkson explicitly stated. Thompson (1992) on the (1994) in Colombia, found that species of this other hand, places them as a subfamily of same tríbe of weevils are pollinators of Curculionidae, a position not inconsistent with Sphaeradenia hamata Barl. Kuschel's presmned placement (given that Studies on the po1lination biology of tribes within Curculioninae of Kuschel are Carludovica palmata and C. drudei Masters in more or less equivalent to subfamilies within Costa Rica by one of us (LDG)· have found Cmculionidae of Thompson). Regardless, both of these plant species also to be attended derelomines clearly are not Erirhininae sensu and pollinated by a variety. of derelomine Kuschel (or Erirhinidae sensll Thompson) curculionids. Collections made by Henry P. given their apomorphic non-orthocerous maJe Stockwell confirm that this same group of genitalia which contrasts with the weevils attend C. palmata in Panama as well. plesiomorphic orthocerous maJe genitalia of Gottsberger (1991), Eriksson (1994) and Seres trueErirhininae. We regard derelomÍne weevils and RamÍrez (1995) report that at least sorne of presently as best placed in the system of the derelomine weevils in their studies Kuschel within Curculioninae as the distinct tentatively were placed as members of the tribe Derelomini (or as Derelominae within genus Schoenherr. One of us (RSA) Curculionidae [system of Thompson]). has examined vOllcher specimens of the weevil species involved in the first two Codons for collections follow Amett aforementioned studies and (as for the species et al. (1993). of weevils attending Costa Rican and Panamanian Carludovica species), would not Systenotelus New Genus,Anderson & assign any of the species ínvolved to Gómez Phyllotrox. Rather, an of these curculionids attendingcyclanths areconsidered to represent a Size: Large to very large (for Derelomini). variety of undescribed genera and wldescribed Total body length of male, 4.9-10.9 mm; species near to, but distinct from Phyllotrox. female, 5.4-12.4 mm. 'Ibe only species which could be confident1y identified was Perelleschus carludovicae Forro: Elongate-narrow to elongate-robust; (Günther) from Carludovica palmata in Costa greatest width al about midlength, variously Rica, Panama and Pem. tapered both anteriorly and posterior1y, In order to pen1'1it an evolutionary extremely so in female specimens of S. perspective of the poBination biology ofthese carludovicae. plants based on . the evolutionary history of their weevil associates, we here describe the Color: Pale yellow, with pronotum with most remarkable and distinctive taxon, species black median line; elytra with various dorsal of which are found in association with andlor lateral black maculations, or Carludovica species in Costa Rica and maculations lacking. Panama. We also present notes about the nature of the.association of these weevils with Rostrum: Slightly shorter than length these plants and discuss their phylogenetic pronotum in mate; slightly shorter than length relationships. pronotum to subequal in length to pronotum in Higher taxonomic ranking and placement of female. Narrow, straight to very slightly derelomine . weevils has been cOllfused. arcuate, of uniform thickness throughout TraditionaUy they have been regarded as length; in dorsal view, widest at base, very Curculionidae: Erirhininae; however, the recent slightly tapered to apex. Antennae inserted at Allael'Sllll &; Gómez: Sy,nenOielus, a remarkabie new' germs 889

about midlength or slightly anterior to Thorack Sienta: Pale yeHow midlength in female, at apical one-third in throughout Procoxal cavities contiguous, maleo Cuticle shiny, densely and finely coxae inserted very close to margin of punctate; with fine indistinct appressed hair­ prostenmrn, distant fmm anterior margino like vestiture, vestiture longer, more erect Prosternum in front of coxae with shOlt mediaHy on venter. Scrobe well-defined, deep, but acuminate posterior intercoxal projectiono continued anteriorly past point of antennal Mesosternum short, markedly f;umescent at inserti.on as fine, narrow moderately deep middle between mesocoxaf:. M�¡aste:mum long, groove. fineJ.y shallowiy pUrlctate, w¡in dense, indistinct appressed hair-like vestíture; Antermae: Smali, compact and robust; posteriorly mediaHy fulTowed in female, scape subequal to ur very slightly Jonger than broadly concave in maleo length of funicle and club combined. Fl.lnide oE seven artieles; artiele 1 davate, as as AbdOhlltm: Fa!';o articles 2-4 combinen; articie 2 slightly clavate, Cutide duH to shiny, ílneJy slightly longer than aruele 3; artides 3-7 punctate, 'vvith dense, fine, indistinct appressed transverse, widened towards apex oí funide. hair-l.ike vestiture. Visible sterna oí V8Iious Club smaH" compact, of three artieles; basal lengths; sternum V long, wñ�h lateral margins and apical artideslonger than rniddle artide. rurected dorsaHy, partially ellclosing latero­ dorsal portions of pygydiuffi in female. Eyes: Large, round, slightly protruded; Pygydium of female flat to ver)' slightly widely separated dorsally. convex, moderately long to ver}' .in dorsal view varioll:>ly constncted at fmm rruddle to Head: Cuticle shiny, coarsely punctate. basal one-third, variouslye:Kpanúed suhapicaHy, Pronotum. Cuticle sbiny, smooth; bhmtly lobed at apex; of male, fla!: dorsaIly, uniformly, very finely fu'1d very densel.y unfOImiy rounded and sorllcwbat bulbous punctate, with dense, fine, indistinctappressed apically, ventrally wilh "v" shaped hair-like vestiture. Fonn robust to elongate emargination. (Uw=O.8-1.4); widest ai or imrnediately anterior to base, uniform!y tapered anteriody to apex, Leg§:: Paje yellow throughout Short and width al apex subeql.lal to width heaa; with very stout; hind iegs "ery sHghtly longer than slight lateral and ventral subapical COl'lstriction. middle iegs, middle legs s1i.ghtly ionger than Basal margin bilobed posteriorly at tuiddle. front legs, ü:tide shi,ny, very finely and Anterior margin lacking lateral pOSí:ocular densely punctate, wit,'1 dense, fine, indistínct lobes. appressed hair-jike vesüture. Femur robust to very vanously ¡ateraJJy flattened, with Elytra: Cutide shiny, smooth. Fonn or 1/áthoutlarge bhmtJy rocmded tooth or¡ inner e10ngate (lhv=2.5-3.0), widest at or margin near apex. Tibia straight, more or less irnmediately anterior to humeri, variously unifmm in width iliroughout; apical comb of tapered posteliody to apex. Striae ten; 1··8 very ilume:rous, sman $opines. Tarsus of five distinct slight1y impressed, 9-10 more deeply artides (usuaBy conceaJ.ed 3i"ticle 4 of impressed; finely and dense!y punctate, Curculionidae distinctIy visible between ¡obes Intervals impunctate to minutely and densely of artide 3); artick 1 stout, very sHghtiy punctate throughout, with or wi.thout dense, 10nger than artide 2; artide 2 stout, shorter fine, indistinct appressed hair-like vestiture. than artide 3; mtide 3 with elangate íateral Elytral apices separateiy rounded, pygydium apicaUy projecíed lobes; rutide 4 very short, moderately to extensiveiy exposed in dorsal narrow, visible between bases of !:obes of view. Humen subquadrate. mide 3; artide 5 elongate, projected beyond apex of Jobes of ru:tide 3 by very more Scutel!um: Visible, large, with fine, than length of articie 3. Tarsa] daws lurge and indistinct appressed nair-likevestiture. robust, simple.

Wings: Present, long. 890 REVISTA DE BIOLOGIA TROPICAL

Genitalia and associated structures: Phylogenetic relationships: Monophyly Genitalia of female with tergum VIII moderate of Systenote/us is indicated by the character to elongate-narrow; sternum VID well­ states of very elongate-narrow genitalia in sc1erotized, elongate-narrow, extended to base females, medially constricted form of the gonocoxite n, apodeme elongate-narrow, "y" female pygydium, and tumescent mesosternal shaped at base; foretube sclerotized or not; process. The closest relative of Systenotelus is gonocoxite TI well sclerotized, moderate to presumed to be an undescribed large derelornine elongate-narrow, styius lacking; vagina collected in ovaries of infructescences of elongate, membranous, irregular in form Sphaeradenia hamata in Colombia. A sister throughout length; bursa copulatrix small to group relationship with this taxon is presurned large, membranous, with spermathecal duct on the basis of the character states of relatively inserted on venter of bursa copulatrix al large body size for Derelomini, pattero of darle confluence with vagina and common oviduct; maculations on a paJe yellow background, and spermathecal duct short or long, spermathecal general habitus. Along with various other gland small or large, spermatheca somewhat Derelomini, the two taxa also share the globose, bluntly rounded, point of insertion of character state of a small compact antennal duct and gland apicaL Genitalia of male with funicle. sternum VID represented by smaU lateral, triangular sclerotized lobes; sternum IX well­ Type species: Systenotelus carludovicae sclerotized, "y" shaped, apodeme moderately Anderson & Gómez, by present designation. long; aedeagus very long and slender. Natural history: AH specirnens of Recognition: Species of Systenotelus are Systenotelus carludovicae were collected on recognized easily by their relatively large body opening inflorescences of Carludovica size (4.9-12.4 mm) for Derelomini, paJe to palmata and C. drudei in December and dark yellow color with various black dorsal and January in the vicinity of San Vito, Costa lateral maculations and general dorsal habitus. Rica. Adults of S. carludovicae were observed They are recognized further by the tumescent copulating OIl, and females were observed mesosternal process, tapered apex of the ovipositing in, inflorescences of C. palmata. abdomen and associated very elongate-narrow AH adult specimens of S. stockwelli were genitalia in females, and by the form of the collected collected on freshly opened female pygydium which in dorsal víew is inflorescences of Carludovica palmata in constricted at the midlength. February, April and June near Gamboa in With the exception of the primarily South Panama. At this same site in June, larvae and American genus Cele tes Schoenherr, species pupae of S. stockwelli were collected in an that are associated prímarily with paIms, other older spadíx oi" C. palmata which was green and Derolomini have shorter total body lengths firm. AduIts of S. costaricensis were collected than species of Systenotellus. Ce/efes species in Panarna in July on opening inflorescences of can easily be distinguished from Systenotelus C. drudei. No information is available on the species by the much more elongate antennal plant associations ol' S. costaricensis in Costa funicle (with the individual artieles much Rica although it is expected also to be longer than wide), the non-tumescent associated with species of Carludovica. In rnesosternal process, form of fernale pygydium Costa Rica, S. costaricensis has becn collected which is not constrictd, the generaHy more from June through August. elongate, cylindrical and curved rostrum and -in sorne species- by the eJongate front legs of Included spedes: There are three species males. AH other Derolomini (including presently recognized in this genus; S. Phyllotrox) which posses a short, stout carludovicae Anderson & Gómez, S. antennal funicle can be separated from costaricensis Anderson & Gómez, and S. Systenotelus species by their smaller size, stockwelli Anderson & GÓmez. non-tumescent mesostemalprocess, and by the form of the gemale pygidium which in dorsal view is not constricted. Anderson & Gómez: Systenotelus, a remarkable new genus 891

Key to adults of the known species of Systenotelus

Pronotum longer than wide (l/w=1.12-1.15 in maje [Fig. 3]; 1.42-1.46 in female [Fig. 1]). Rostrum relatively long (0.77-0.78 times the length of pronotum in male [Figs. 3,4]; 0.96-1.04 times the length of the pronotum in female [Figs. 1,2]). Each femnr of maJe and middle and hind femora of female with large subapical blunt angulation. Abdominal stemllm V of female long and narrow, much longer than wide; abou! as long as stema 1 and n combined (Fig. 13). Elytra of female in dorsal view with posterior one-half tapered to narrow apex (Fig. 1). Aedeagus in dorsal view with margins uniform, not sinuate (Fig. 6)...... S. carludovicae Anderson & Gómez (Costa Rica; on Carludovica palmata Ruíz & Pavón and C. drudei Masters ) l' Pronotum wider than long (l/w=0.84-0.95 in male [Figs. 16, 29]; l/w=0.89-0.94 in female [Figs. 14, 27]). Rostrum short (0.69-0.79 the length of pronotum in male [Figs. 16, 17, 29, 30]; 0.75-0.82 in female [Figs.14, 15, 27, 28]). Femora withollt subapical angulation, uniformIy rounded. Abdominal stemum V of female robust, as long as wide; about as long as stema I-IV combined (Figs. 26, 39). Elytra of female in dorsal view with posterior one-quarter tapered to broad apex (Figs. 14, 27) . Aedeagus in dorsal view with margins sinuate (Figs. 19, 32 ...... 2

2 (1') Body size 7.7-9.9 mm . Dorsal elytral black maculations present, on intervals 3- 4 near base (Iacking in sorne specimens), interval 4 al anterior one-third, posthumeral region on intervals 7-9, posterior portion of intervals 5-8, and intervals 10-11 throughout length (Figs. 14-17). RoslrUm in lateral view slightly ventrally arcuate in female, slightly tapered to apex in both sexes (Figs. 15, 17). Aedeagus in dorsal view uniform1y tapered from base to near apex (Fig. 19) ...... ::;"" ...... _S. costaricensis Anderson & Gómez (Costa Rica, Panama; on Carludovica drudei Masters)

2' Body size 4.9-6.9 mm. Dorsal elytral black maculation either entirely lacking or Iimited to lateral marginal region and entire posterolateral one-halfto two-fifths (Figs. 27-30). Rostrum in lateral view straight in female, distinctly tapered to apex in both sexes but especially so in femaJe (Figs. 28, 30). Aedeagus in dorsal view constricted near base, then tapered to near apex (Fig. 32)...... S. stockwelli Anderson & Gómez (Panama; on Carludovica palmata Ruíz & Pavón)

Systenotelus carludovicae New Species, Anderson & Gómez (Figures 1-13, 40-41)

Description: (Figs. 1-13). Total body dense punctures and fine hair-like vestiture. length of male 9.9-10.8 mm (n:::lO); female, Elytra of female in dorsal view with posterior 11.2-12.4 mm (n=lO). Maximum body width one-half tapered to narrow apex. Mesosternum (across widest point of elytra) of male 3.7-4.1 markedly tuberculate at middle between mm; female, 3.8-4.1 mm. Color pale yellow mesocoxae. Abdominal sternites I-II of male with following areas black: rostrum; midline of subequal in length, each about twice the length head near posterior margin; flanks andmidline of each of sternites III-IV; sternites ID-IV of pronotum; scutellar region; elytral sutural relatively long, subequal in length; sternum V line, base of intervals 4-5, intervals 4-5 at about as long as wide, about as long as sterna 1 anterior one-third, humeral region, posterior and TI combined. Abdominal sternites I-IV of one-tbird to two-fifths of intervals 4-8, and femalemore or less similar in length, stemum intervals lO-U throughout length; and coxae. 1 slight1y longer than II, n slightly longer than Rostrum 0.77-0.78 times the length of m, III slightly shorter than IV;sternum V long pronotum in male; 0.96-1.04 times the length and narrow, much longer than wide, about as of pronotum in female. Rostrum in lateral long as sterna 1 and n combined. Pygydium of view not to only slightly tapered to apex, more female very long, in dorsal view markedly so in the maleo Antennal insertion at anterior constricted at middle, expanded subapically, one-trurd in male, submedial in female. bluntly lobed at apex; of male, flat dorsally, Antennae with scape not reaching eye by at unfonnly rounded and somewhat bulbous least width of antennal club or more. Pronotum apically, ventrally with "v" shaped incision. longer than wide (l/w=1.12-1.l5 in maJe; 1.42- Each femur of male with large blunt subapical 1.46 in female). Elytra with various black tooth; front femora of femate lacking dorsal and lateral markings; intervals with fine angulation, unifonnly rounded, middle and bind REVISTA DE BiOLOGIA TROPICAL Anderson & Gómez: Systenotelus, a remarkable new genus 893

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11 13 894 REVISTA DE BIOLOGIA TROPICAL Anderson & Gómez: Systenotelus, a remarkable new genus 895

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24 26 8% REVISTA DE BIOLOGIA TROPICAL Aaderson & G6mez: Systenotelus. a relllllrlcable new genus 897

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35 898 REVISTA DE BIOLOGIA TROPICAL

femora of female with subapical bIunt km.S.E.) de Tajo Viejo, Fila Cal, 330m. angulation. Genitalia of female with tergum 16XU.l994,(941309) L. D. Gómez, on vm extremely elongate-narrow, much longer Carludovica palmata". Paratypes, 2 females, 5 than wide, apically very heavily sclerotized, males, "COSTA RICA: PunL Coto Brus, Las black in color; sternum VIII weU-sclerotized, Cruces 1 100m. 21.XIJ.1994, L. D. Gómez , elongate-narrow, extended to base gonocoxite on Carludovica palmata". Paratypes, IS n, apodeme similarly elongate-narrow, "y" females, 21 males, "COSTA RICA: Punt.Coto shaped at base; foretube sclerotized; gonocoxite Brus, Las Cruces 1l00m. XII.l994, L. D. II very elongate, length many times width, Gómez, on Carludovica pa/mata". Paratypes, with laterally directed small heavily sclerotized 17 females, lS males, "COSTA RICA: Punt. apica! tooth; bursa copulatrix short. Genitalia Coto Brus, Las Cruces, l100m. 2SXU.l994 of maje with sternum IX "y" shaped, apodeme Luis D. Gómez, on Carludovica palmata". moderately long, slender; aedeagus extremely Paratype male, "COSTA RICA: Punt.Osa long and slender (length about 11 time the Península, Chacarita, > 100m, 20.III.l995, L. width at base), in dorsal view tapered from base D. Gómez, on Carludovica palmata ". to about apical one-quarter then slightly Paratypes in AMNH (4), BMNH (4), CASC swollen laterally then tapered to bluntly (2), CMNC (51), CNCI (4), CWOB (6), rounded apex; in lateral view uniformly FMNH (2), HAHC (4), INBC (20), MUCR slightly arcuate, apex slightly deflexed; no (8), SEMC (4) and personal collection of LDG internal sac or internal sclerotization evident; (14). Total 59 femates, 66 males. apodemes relatively short, about one-thírd length aedeagus; tegmen narrow, basal apodeme Variation: Specimens vary slightly in the as long as each apodeme of aedeagus. extent of black markings on the elytra, particularlythose on intervals 4 and 5 near the Type material: Holotype male, "COSTA base and at the anterior one-third. RICA: Punto Las Cruces Biol. Stn. Coto Brus, 1l00m. 1.1994 , L. D. Gómez P. on opening Distribution: This species is known only inflorescence of Carludovica pa/mata" from the Valle de Coto Brus and the Osa (INBC). Allotype female (INBC) and 7 female Península in Prov. Puntarenas, in southern and 3 male paratypes labelled as holotype. Costa Rica (Fig. 41). Paratypes, 6 females, 10 males, "COSTA RICA: Punto Las Cruces Biol. Stn. Coto Brus, Natural history: AH specímens of 1 100m. , XII.1994 , L. D. Gómez P. on Systenotelus carludovicae were coUected on opening inflorescence of Carludovica opening inflorescences of Carludovica palmata". Paratype female, "COSTA RICA: palmata and C. drudei (Cyclanthaceae) in Punto , San Vito, 1200m , May 1993, Luis D. December and January in tile Valle de Coto GÓmez-P. inflores. of Carludovica drudei & Brus and in March in the Osa Península. C. palmata". Paratypes, 2 males, "COSTA Adults of S. carludovicae were observed RICA: Punt. Coto Brus, Cerro Paraguas, 1430 copulating on, and females were observed m, lS.XU.l994 , (941316) L. D. Gómez, on ovipositing in, flowers of C. pa/mata (Fig. 40) Carludovica palmata". Paratypes, 1 female, 2 in the vicinity of San Vito. males, "COSTA RICA: Punt. Coto Brus, A briefdescription of the natural history of Cerro Paraguas, 1600m, lSXU.1994, Carludovica species at Coto Brus, Puntarenas (941314) L.D. Gómez, on Carludovica Province, Costa Rica follows. Although plants drudei". Paratypes, 2 females, "COSTA of Carludovica bloom sporadically throughout RICA: Punt. Cerro NeiUy (3 km.N.W.), the year, their populations are flowering from 200m. lSXU.1994,(941313) L.D. Gómez , on December to May, with peak production Carludovica drudei". Paratypes, 3 females, 1 between April and May. Clumps of male, "COSTA RICA: Punt. Las Cruces (15 Carludovica pa/mata and C. drudei produce, km.S.E.) del Tajo Viejo, Fila de Cal 560m. on average, five stalked inflorescences during 16.xU.1994 ,C 941305) L. D. Gómez, on the peak of their flowering period (late-April to Carludovica drudei". Paratypes, 1 female, 2 early-May) at l100m in cloud forest habitat. males, "COSTA RICA: Punt.Las Cruces (15 Inflorescences do not mature syncmonously and Anderson & Gómez: Systellotelus, a rematkable new genus 899

simultaneous anthesis of more than two is very rareo UsuaBy, inflorescences open in a sequence Description: (Figs. 14-26). Total body of 24-36 hour intervals. In both species, the length of male 7.7-9.2 mm (n=5); female 8.8- inflorescence is a peduncled spadix with flowers 9.9 mm (0=8). Maximum body width (across arranged in very compact, spira} rows. When widest point of elytra) of male 3.5-4.2 mm; maturity is reached, and through the initial female 3.4-4.2 mm. Color pale yellow with stages of anthesis, the inflorescence is following areas blaek: rostrum; midline of head demonstrably thermogenetic and by the time near posterior margin; flanks and midline of the spathal bracts open and the osmophoric pronotum; scutellar region; intervals 3-4 near staminodes expand, volatilízed scents are base (lacking in sorne specimens), interval 4 at released. The aroma is a mixture of spicy and anterior one-third, posthumeral regíon on fruity, with a distinct lemon component. intervals 7-9, posterior portion oí intervals 5-8, Anthesis begins at dusk with a bulging of the and intervals 10-11 throughout length; and bracts and an increase in temperature and is eoxae. Rostrum 0.75-0.79 time the length of completed in the early morning with the full pronotum in male; 0.75-0.80 the length of the expansion of the staminodes. With the first pronotum in female. Rostrum in lateral view openings between the bracts, start to slightly tapered to apex, slightly ventraUy enter the inflorescence and continue to increase areuate in female, straight in maleo Antennal in number as anthesis procedes. insertion at anterior two-fifths in male and Examination of half-opened inflorescences female. Antennae with scape noí reaching eye sometimes reveals the presence of several by much less than width of antennal club. inseets such as Dermaptera and Coleoptera Pronotum wider than long (1Jw = 0.93-0.95 in (Nitidulidae and Staphylinidae), particu1arly male; lIw=0.89-0.90 in female). Elytra with when the cyc1anths are in proximity to speeies various black dorsal and lateral markings; of Araceae. In more isolated Carludovica intervals with fine dense punctures and fine clumps, one may find a few Scarabaeidae hair-like vestiture. Elytra of female in dorsal (Cyclocephala spp.), but almost exclusively view with posterior one-quarter tapered to the area variety of Curculioninae of t.!-¡e relatively broad apex. Mesostemum markedly tribe Derelomini. At the outset of their tuberculate at middle between mesoeoxae. aggregation, the weevils (which may be very Abdominal stemites I-ll of mate subequal in Iíumerous) move over the inflorescenee length, each about twice length of each of scurrying around the bases of the extruded sternites ID-IV; sternites ID-IV short, subequal staminodes. Mating, as evidenced by many in length; sternum V slightly wider than long, mounted and copulating pairs of weevils about as long as stema I and II combined. (incIuding S. carludovicae and vanous other Abdominal sternites l-ll of female subequal in derelomines), oecurs when the staminodes start length, eaeh about 1.5 the length of sternites to faH off the flowers. By the time the ID-N combined; sternites ID-N very short, staminodes have ful1y dropped, many weevils subequal in length; sternum V robust, as long are burrowing into the inflorescenee; however, as wide; about as long as sternaI-IV combined. other weevils may fall to the ground or may fly Pygydium of female long, in dorsal view to nearby infloreseences. Other vlsItors constrieted at basal one-third, very slightly attracted by the seent and the availability of expanded subapically, bluntly lobed at apex.; of pollen arestingless bees (Trigonafulviventris) male, flat dorsally, unfornlly rounded and and an oceassional euglossine beco somewhat bulbous apically, ventraUy with "v" shaped incision. Femora of maJe and female Phylogenetic relationships. This species lacking subapical blunt angulatioll, uniforrnly is presumed to be the sister speeies to fue rounded. Genitalia of female with tergum VID lineage comprised of S. stockwelli and S. elongate-narrow, muen longer than wide, costaricensis. Systenotelus costaricensis apically very heavily sclerotized, black in New Speeies, Anderson & Gómez color, with . lateral arms extended to base (Figures 14-26, 41) gonocoxite n, somewhat expanded, heaviIy sclerotized; stemum VID well-sclerotized, elongate-narrow, extended to base gonocoxite 900 REVISTA DE BIOLOGIA TROPICAL

n, apodeme similarly elongate-narrow, "y" laterally then tapered to bluntly rounded apex; shaped at base; foretube sclerotized; gonocoxíte margins somewhat irregularIy sinuate II elongate, length many times width, lacking throughout apical one-half; in lateral view apical tooth; bursa copulatrix moderate. uniforrnly slightly arcuate, apex slíghtly Genitalia of male with sternum IX "y" shaped, deflexed; no internal sac or internal apodeme moderately long; aedeagus extremely sclerotization evident; apodemes relatively long and sIender (length about 11 x width at short, about one-trurd length aedeagus; tegmen base), in dorsal view tapered from base to about narrow, basal apodeme as long as each apodeme apical one-quarter then slightly swollen of aedeagus.

Figure 41. Collection localities of Systenotelus species. S. carludovicae, cueles; S. costaricensis, squares; S. stockwelli, triangles.

Type material: Holotype male, "Ref. Barrios". Paratypes in CMNC (8), CWOB Nac. Fauna Silv. Tapantí, 125Om. Prov. Cart. (11), INBC (5), STRI (4). Total 16 females, 14 COSTA RICA, G. Mora, Ago. 1991, L-N- males. 194000, 559800" (INBC). AHotype female (INBC) and 2 female and 3 maJe paratypes Variation: There is slight variation in the labelled as holotype. Paratypes, 4 females, 1 pattern of black elytral maculations. In most male, "Est. Hitoy Cerere, 100m. R.Cerere, specimens the pattem is typical; however, in a Res. Biol. Hitoy-Cerere, Prov. Limón, Costa few specimens the median portion of the elytral Rica, 30 Jul.- 20 Jun. 1992, F.A.Quesada L­ disk is more extensively maculated. In a few N-1842oo, 643300" Paratype female, "COSTA specimens, the midline of the pronotum is RICA Cart. Prov. Turrialba, CATIE grounds, black only in the anterior one-half. ca. 2000' raiofan, II-12-1990, J. Rifkínd & P. Gum". Paratypes, 6 females, 7 males, Distribution: This species is known from "PANAMA. Colón, C.Z. Ft. Sherman, OId three proximal sites in the provinces of Ft. San Lorenzo Rd. 7-25-1995 , C.W. & L.B. Cartago, Limón and Turrialba, Costa Rica, and O'Brien" "on Carludovica drudei flower, from the Canal Zone in Panama (Fig. 41). (Cyclanthaceae)". Paratypes, 1 femate, 3 males, "Panama. Provincia de Colón, Fuerte Sherman Natural bistory: Adult specimens of , Carludovica drudei I 25.VII.1995, H. Systenotelus costaricensis were collected on Anderson & Gómez: Systenotelus, a remarkable new genus 901

freshly opened inflorescences of Carludovica very short, subequal in length; sternum V drudei (Cyclanthaceae) in July near Fort robust, as long as wide; about as long as stema Sherrnan, Panama in lowland rain foresto There I-IV combined. Pygydiuníc'of female long, in is no inforrnation on the plant associations of dorsal view constricted at basal one-third, very this species in Costa Rica; adults have been slightlyexpanded subapically, bluntly lobed at collected in Costa Rica from June through apex; of male, flat dorsally, unforrnly rounded August. and somewhat bulbous apically, ventrally with "v" shaped incision. Femora of male and Phylogenetic relationships: Systenotelus female lacking subapical blunt angulation, stockwelli and S. costaricensis are presumed uniformly rounded. Genitalia of female with to be sister species based on the shared tergum VIn elongate-narrow, much longer than apomorphic character states of aedeagus in wide, apically heavily sclerotized, dark brown dorsal view with lateral margins sinuate, and in color, with lateral arms extended to base the very short abdominal sternites lIT and IV. gonocoxite n, somewhat expanded, heavily Systenotelus stockwelli New Species, sclerotized; sternum vm well-sclerotized, Anderson & Gómez elongate-narrow, extended to base gonocoxite (Figures 27-39, 41) n, apodeme similarly elongate-narrow, "y" shaped at base; foretube sclerotized; gonocoxite Description: (Figs. 27-39). Total body n elongate, length many times width, lacking length of male 4.9-6.2 mm (n=lO); female 5.4- apical tooth; bursa copulatrix moderate. 6.9 mm (n=10). Maximum body width (across Genitalia of male with sternum IX "y" shaped, widest point of elytra) of male 2.3-3.0 mm; apodeme moderately long; aedeagus extremely female 2.5-3.0 mm. Color pale yellow with long and slender (length about 11 times the following areas black: midline of head near width at base), in dorsal view constricted at posterior margin (lacking in sorne specimens); about basal one-sixth then tapered to about midline of pronotum; elytral intervals 10-11 apical one-sixth then very slightly swollen throughout length and intervals 2-9 or 3-9 in laterally then tapered to bluntly rounded apex; posterior one-half to two-fifths (lacking in margins irregularly sinuate throughout apical sorne specimens). Rostrum 0.69-0.72 x length one-half; in lateral view uniforrnly slightly pronotum in male; 0.80-0.82 x length arcuate, apex not deflexed; no internal sac or pronotum in female. Rostrum in lateral view internal �clerotization evident; apodemes distinctly tapered to apex, especially so in relatively short, about one-third length female; straight in both sexes. Antennal aedeagus; tegmen narrow, basal apodeme as insertion at anterior two-fifths in male, long as each apodemeof aedeagus. sudmedian in female. Antennae with scape not reaching eye by much less than width of Type material: Holotype male, antennal club. Pronotum wider than long (lIw "PANAMA: Canal Area, Pipeline Road 4 km. = 0.84-0.92 in male; lIw=Ü.89-0.94 in female). N.W. Gamboa, 25 Feb. '95, Stockwell", "on Elytra with various black dorsal and lateral inflorescence of Carludovica palmata" markings (sorne specimens lacking black (CMNC). Allotype female (CMNC) and 2 maculations); intervals with fine dense female paratypes labelled as holotype. punctures and fine hair-like vestiture. Elytra of Paratypes, 6 females, 6 males, labelled as female in dorsal view with posterior one-quarter holotype except "3 km. N.W. Gamboa". tapered to relatively broad apex. Mesosternum Paratypes, 10 females, 14 males, labelled as markedly tuberculate at middle between holotype except "3 km. N.W. Gamboa, 2 Apr. mesocoxae. Abdominal sternites I-n of male ·' 95 ". Paratypes, 19 females, 18 males, labelled subequal in length. each about twice the length "PANAMA: Panama . Pipeline Rd. km.4.0- of each of sternites lIT-IV; sternites lIT-IV short, 6.0 nr. Gamboa, 4Om, 23.VI.1995, R.S. subequal in length; sternum V slightly wider Anderson, tropo lowland foro ex. flowers than long, about as long as sterna 1 and 11 Carludovica palmata, 95-040". Paratypes in combined. Abdominal sternites I-n of female AMNH (4), BMNH (4), CMNC (39), CWOB subequal in length, each about 1.5-2.0 x length (4 + 4), HAHC (4), HPS (12), USNM (4). of sternites m-IV combined; sternites m-IV Total 38 females, 39 males. 902 REVISTA DE BIOLOGIA TROPICAL

Variation: Specimens vary in the extent of DISCUSSION dorsal elytral black maculations which are almost entirely lacking in sorne specimens. Although there are no confmned fossil records of Cyclanthaceae, except for the Distribution: This species is known only "protocyclanthoid" structures described fromthe fromPipeline Road near Gamboa in the Canal Eocene of India, wruch could represent Area of Pan ama (Fig. 41). Arecaceae or Pandanaceae rather than Cyclanthaceae (the present dístribution of Natural bistory. AH specimens of which is excIusively New World), roost Systenotelus stockwelli were collected on systematists agree that Cyclanthaceae are an old freshly opened inflorescences of Carludovica group whose affinities are with the palms palmata (Cyclanthaceae) in February, April and (Arecaceae) and screw-pines. PaIros are well in June near Gamboa in Panama in lowland known as cantharoprulous plants, and have a rain forest. Flowering in C. palmata at this site variety of Curculionidae, including many takes place most frequently from March to Derelomini, arnong their associates. April and during intensive examination of Weevils of the Derelomini are also associated many individuals in June (most of which bore with Cyclanthaceae; however, the derelornines from 1-5 mature spadices bearing fruil) only recorded by us from cyclanths are absent from one flowerwas found. flowers of other plant families (including In June, immature stages of S. stockwelli paIms) when cyclanths are not blooming, were found in an older spadix wruch was green suggesting an exclusive association of these and firm and wruch had not yet started to taxa with cyclanths. Similarly, palm-associated decompose andexpose seeds and the bright red Derelomini do not appear to be found on fleshy fruit pulpo Each mature S. stockwelli cyclanths. Erikkson (1994) suggested that the larva was found to have fully consumed a relationship of weevils with Cyclanthaceae is package of seeds but did not appear to have not necessarily an aneient, co-evolved system, consumed or otherwise entered the surrounding de<·�;te the presence of Derelomini on tlowers red f1eshy fruit pulpo Although a number of of both cyclanths and paIms, and based on spadices were exarnined, only 12 mature larvae presently available data we agree. and 2 pupae were reeovered, all from a single Curculionoidea are an old group of beetles spadix. Pupation takes place in the seed cavity whose fossil record predates that of the and adult emergence, while not observed (but emergence of the early flowering plants (see see comments fol1owing Oil Perelleschus Kuschel et al. 1994 for a review of known carludovicae), likely awaits natural exposure Cretaceous [and Upper Jurassic] weevil fossils). of the spadix contents. AH of these early weevil fossils are assignable During dissection of green spadices while to the primitive curculionoid families looking for S. stockwelli, numerous adults of Nemonychidae, Belidae, and Brentidae (both Perelleschus carludovicae emerged, Carinae and Brentinae: Eurhynchini). Host indicating thateven though adults, they had not plants of fossil taxa in at least two of these exited the spadix and were likely awaiting faroilies (Nemonyerudae and Belidae) appear naturalexposure. likely to have been various gyronosperms, such as conifers and cycads (or related plants), Pbylogenetic relationsbips: Systenotelus as these are the among the known hosts of stockwelli and S. costaricensis are presumed extant members of these farnilies and are to be sister species based on the shared present as fossils in these assemblages. apomorphic character states of aedeagus in Possible hosts of the Brentidae are less cIear. dorsal view with lateral margins sinuate, and True Curculionidae first appear in Upper the very short abdominal sternites m and IV. Cretaceous deposits in North America, China and Chile, well after the presurned origin of angiosperms in the Lower Cretaceous; hosts of these laxa arenot known. However, among the Baltic amber Curculionidae of Oligocene age is Electrotribus weigangae (UIke) (Kuschel Anderson & Gómez: Systenotelus, a remarkable new genus 903

1992), a laxon apparent1y most closely related ACKNOWLEDGMENrS to the palm-associated Neotropical genus Ce/etes Schoenherr of the Derelomini. Since We thank Henry Stockwell and Hector paIms were weU-represented at fue time of Barrios of the Smithsonian Tropical Research formation of Baltic amber, E weigangae also Institute (STRl) in Panama, Angel Solis of the may have been associated with paIms. Thus the Instituto Nacional de Biodiversidad (INBio) in association of sorne Derelomini with paIms Costa Rica, and Charlie O'Brien of Florida would appear to extend to at least the A&M University, Tallahassee, Florida for the Oligocene and would make the prospect of loan of specimens. Susan Laurie-Bourque other early Cenozoic Dere10mini being preparedthe Une drawings and the plates. LDG associated with Cyclanthaceae at least possible. thanks GaiI Hewson Gómez for pointing out On the other hand, while their exact this pollination syndrome which she compared phylogenetic relationships are not clear, to the relationship between weevils of the weevils associated with paIms and cyclauths are genus Rhopalotria in the subfamiIy not primitive in their phylogenetic position Oxycoryninae (family Belidae) andthe cones of within Curculionidae despite traditionaI Zamia sp. (Cycadaceae). placement as Erirhininae (see Introduction). Apomorphic features of mate genítalic structure indicate placement with other Curculionidae RESUMEN having similar non-orthocerous genitaIia. Kuschel (1952) had assigned derelomines lo Se describe un nuevo género y tres nuevas especies de coleópteros (Curculionidae, Curculioninae, Petalochilinae hut later (1964) questioned this Derelomini), asociados con las inflorescencias de placement and in the recent checklists of New especies de las ciclantáceas, Carludovica palmala y C. World weevils (O'Brien and Wibmer 1982; drudei, en Costa Rica y Panamá.Aunque se conocía la polinización de estas plantas por coleópteros, el informe Wibmer and O'Brien 1986) they más reciente asignaba los insectos al género Phyllotrox. inappropriately (but traditionally) are placed Un estudio del material descrito nos confIrma que esa identifIcación es incorrecta y, además, que el material back in Erirhininae. costarricense y panameño incluye un número de géneros While it is possible that association of y especies aún no descritas. Aquí se describe el primer Derelomini with cydanths may date to the material obtenido de nuestros estudios y se presentan notas sobre la historia natUral de las tres especies del early Cenozoic, we do not expect parallel nuevo género Systenotelus: S. carludovicae, S. cladogenesis or a close coevolutionary history costaricensis yS. stockewelli. between these weevil species and the Cyclanthaceae-Arecaceae lineage of plants. REFERENCES Association of Cyclocephala scarabs with Cyclanthoideae, and lack of any Imown Anderson, RS, 1993. Weevils and plants: Phylogenetic derelomine weevil associates for versus ecological mediation of evolution of host plan! Cyclanthoideae, suggests fue independent associations in Curculioninae (Coleoptera: Curculionidae). Mem. Entomo!. SOCo Canada 165: 197- origin of associations of derelomines with each 232. of paIms and cyc1anths" perhaps because of similarities in floral structure and habitat Arnett, RH. Jr. GA Samuelson & G.M. Nishida. 1993. The lnsect and Spider Collections of the World. Fauna associations. In support of this view, Anderson & Flora Handbook No. 11. Sandhill Crane, Gainesville, (1993) in a general review of evidence for Florida, vi + 310 pp. cospeciation in weevils and plants, did not find Beach, J.H. 1982. Beede pollination of Cyclanthus evidence for cospeciation, rather factors bipartitus (Cyclanthaceae). Amer. 1. Bol. 69: 1074- mediating speciation in weevils appeared likely 1081.

to be primarily ecological in nature, a common Drude, O. 1877. Über den Bau und die systematische factor being similarity in habitat associations. Stellung der Gattung Car(udovica. Bot. Zeit. 35: 591- However, before cospeciation can be dismÍssed 594.

as the mode of evolution in these taxa, more Eriksson, R 1994. The remarkable weevil pollination of information on fue range of host plant the neotropical Carludovicoideae (Cyclanthaceae). associations and phylogenetic relationsmps in PlantSyst. Evo!. 189: 75-81. Derelomini is necessary. Gottsberger, G. 1991. Pollination of some species of the Carludovicoideae, and remarks on the origin and 904 REVISTA DE BIOLOGIA TROPICAL

evolution of the Cyclanthaeeae. Bol. Jahrb. Syst. 113: descriplions of a new genus and species and 221-235. phylogenetic and zoogeographical cornments (Coleoptera: Curculionoidea). Entomologica Harling, G. 1958. Monograph of the Cyclanthaeeae. Acta Scandinavica 25: 137-149. Horti Berg. 18: 1-428. O'Brien, C.W. & GJ. Wibmer. 1982. Annotated checklist Kuschel, G. 1952. Los Curculionidae de la cordillera of the weevils (Curculionidae sensu lato) of North chileno-argentina (l.a parte). Rev. Chilena Entorno!. 2: Ameriea, Central Ameriea, and Ihe West lndies 229-279. (Coleoptera: Curculionoidea). Mem. Amer. Entorno\. Institute 34: 1- 382 . Kuschel, G. 1964. Insects of Campbell Island. Coleoptera: Curculionídae of the subantarctic islands of New Seres, A. & N. Ramírez. 1995. Biología floral y plinización Zealand. Paco Insee. Monogr. 7: 416-493. de algunas cotiledóneas de un bosque nublado venezolano. Ann. Misouri Bol. Gard. 82: 61-81. Kuschel, G. 1992. Reappraisal of the Baltie Amber Curculionoidea described by E. Voss. Mitteíl. aus d. Thompson, R.T. 1992. Observations on Ihe morphology Geolog.-Palaonlolog. lnst. d. Universitat Hamburg 73: and classification of weevils (Coleoptera, 191-215. Cureulionoidea) wilh a key lo major groups. J. Nat. Hist. 26: 835-891. Kusehel, G. 1995. A phylogenetie classification of Curculionoidea to families and subfamilies. Mem. Wibmer, GJ. & C.W. O'Brien. 1986. Annotated checklist Entorno!.Soco Washington 14: 5-33. of Ihe weevils (Curculionidae sensu lato) of South AmeIica (Coleoptera: Curculionoidea). Mem. Amer. Kusehel. G. R.G. Oberprieler & R.J. Rayner. 1994. Entorno!. Institute 39: 1 - 563. Cretaceoll$ _wils from southem Afrier!., with