pharaxonotha portophylla (coleoptera: erotylidae), new species

Pharaxonotha portophylla (Coleoptera: Erotylidae), new species andpollinator of Zamia (Zamiaceae) in Puerto Rico

NICO M. FRANZ1,3 AND PAUL E. SKELLEY2

1Department of Biology, PO Box 9012, University of Puerto Rico, Mayagüez, PR 00681, U.S.A.2Florida State Collection of Arthropods, Florida Department of Agriculture – DPI, 1911 SW 34th St., PO Box 147100,

Gainesville, FL 32614-7100, U.S.A. 3 Corresponding author: [email protected]

ABSTRACT.—Pharaxonotha portophylla, new species (Insecta: Coleoptera: Erotylidae: Pharaxonothinae), isdescribed based on individuals collected on the cones of native populations of Zamia amblyphyllidia D. W.Stevenson and Z. portoricensis Urban (Cycadales: Zamiaceae) in the Puerto Rican Cambalache and SusúaCommonwealth Forests, respectively. The new species is a member of a complex centered around P. floridana(Casey), and may be distinguished from other Pharaxonotha species by the following traits: head lackingsupraocular striae, with coarse punctation; pronotum slightly transverse, anterolateral edges weakly pro-jected and angulate; protibia narrowly triangularly widened toward apex; elytra with an anterior marginalline, elytral punctures coarse, not impressed, elytral striae not grooved, and elytral intervals plane. Phara-xonotha portophylla is the second species in this genus reported for the Caribbean region, raising the numberof known Pharaxonotha species to 13 worldwide. Samples of insect visitors on Zamia cones of either genderand at various stages of anthesis strongly suggest that P. portophylla is the primary and sole pollinator ofthese cycad species at the two locations. The abundance of adult individuals was much lower on the receptivemegastrobili than on the pollen-releasing microstrobili which produced similar attracting odors. Larvaldevelopment was restricted to the microstrobili, possibly indicating the presence of defensive metabolites inthe megastrobili and a reproductive interaction with “pollination by deceit”.

KEYWORDS.—cycad pollination, insect systematics, Pharaxonothinae, pollinator reproduction

INTRODUCTION

Species in the predominantly pantropicalgenus Pharaxonotha Reitter (Coleoptera:Erotylidae: Pharaxonothinae sensu Leschen2003; see also Wegrzynowicz 2002, Robert-son et al. 2004, Leschen & Buckley 2007)represent an old lineage of Erotylidae withlongstanding reproductive associationswith the Cycadales (see, e.g., Tang 1987,Vovides et al. 2003, Chaves & Genaro 2005;and references therein). As presently cir-cumscribed the genus contains 12 species(Sen Gupta & Crowson 1971, Leschen2003), of which two occur in CentralAmerica, Mexico, and the southeasternUnited States (Leschen & Skelley 2002), twoadditional species in Costa Rica (Pakaluk1988), and one in Cuba (Chaves & Genaro2005). The remaining species are predomi-nantly (southeast) Asian. Together withweevils in the genera Parallocorynus Vossand Rhopalotria Chevrolat (Belidae: Oxyco-ryninae; see Marvaldi et al. 2006), Phara-xonotha are considered the primary pollina-

tors of many Central American andCaribbean cycads, and are therefore of in-terest to coevolutionary analyses and con-servation management policies (e.g., Ober-prieler 2004, Tang 2004).

The native Puerto Rican cycads areplaced in the prominent New World genusZamia Linnaeus (see Walters & Osborn2004), and include the endemic Z. portori-censis Urban, as well as Z. amblyphyllidiaD. W. Stevenson and Z. pumila Linnaeuswhich also inhabit other Caribbean islands(Eckenwalder 1980, Stevenson 1987, Liogier& Martorell 2000). In particular the popu-lation structure of Z. amblyphyllidia innorthern Puerto Rico is well understood(Negrón-Ortíz & Breckon 1989, Negrón-Ortíz et al. 1996). In contrast, the only pub-lished reference on the pollinating agents ofany Puerto Rican species of Zamia is a noteby Breckon & Negrón-Ortíz (1983) who ob-served mycetophilid gnats visiting and re-producing in the cones of the same species.Here we describe a new species of Phara-xonotha, and present natural history infor-

Caribbean Journal of Science, Vol. 44, No. 3, 321-333, 2008Copyright 2008 College of Arts and SciencesUniversity of Puerto Rico, Mayaguez

321

mation indicating that this species is theprimary pollinator of Z. amblyphyllidia andZ. portoricensis at two geographically sepa-rate Puerto Rican localities.

MATERIALS AND METHODS

Study Sites and Collecting Methods

Cones of Z. amblyphyllidia (Figure 1) andZ. portoricensis (Figure 2) were sampled forvisiting insects at the Cambalache andSusúa Commonwealth Forest Reserves, re-spectively, during the period of January 12to February 12, 2007. Cambalache is locatedcentrally near Arecibo along the northerncoast of Puerto Rico, and is classified as asubtropical moist forest habitat on lime-stone soils (see Negrón-Ortíz & Breckon1989, Lugo et al. 2001). On the other hand,Susúa is located near Sabana Grande on thesouthwestern edge of the Central Cordil-lera, and is classified primarily as a dryslope forest habitat on serpentine soils (seeEwel & Whitmore 1973, Figueroa Colón1996). While the species concepts for PuertoRican Zamia are not conclusively resolved,we herein adopt the relatively narrow con-ceptions proposed by Stevenson (1987).

Insect visitors were taken from both fe-male megastrobili (Figures 1A & 2A) andmale microstrobili (Figures 1B & 2B), atvarious stages of development, by cuttingoff entire cones and placing them immedi-ately in whirl-pak containers. Insects crawl-ing out of receptive cones were subse-quently transferred in separate vials with95% ethanol. Older cones were returned tothe laboratory and monitored for 1-3 weeksfor emerging immature and adult insects.The samples were pooled into three majorphases of development: (1) pre-anthesis, i.e.1-5 days prior to the appearance of fissuresbetween sporophylls and the production ofodors; (2) anthesis, i.e. the stage of dehis-cence, micropyle receptivity (�) or pollenrelease (�) and production of sweet, resinyodors, with a duration of approximatelythree days; and (3) post-anthesis, i.e. 5-30days past cone receptivity (�) or pollen re-lease (�), when odors are no longer percep-tible. At this stage, the megastrobili arebeginning to set fruit whereas the mi-crostrobili are slowly decomposing.

Systematics

The description mostly follows the for-mat of Franz (2003), with occasional refer-

FIG. 1. Zamia amblyphyllidia at Cambalache. (A) re-ceptive megastrobilus; (B) microstrobilus, approxi-mately 12 hours prior to releasing pollen. Photos takenat 10:00 a.m.

FIG. 2. Zamia portoricensis at Susúa. (A) receptivemegastrobilus; (B) microstrobilus, approximately 20hours past opening of anthers.

N. M. FRANZ AND P. E. SKELLEY322

ence to Chaves & Genaro (2005) to facilitatean understanding of similarities and differ-ences with the species described in thatpublication. The new species of Pharaxono-tha was identified through application ofthe phylogenetic species concept (empha-sizing unique character combinations; seeWheeler & Platnick 2000). The morphologi-cal terms used are in accordance withTorre-Bueno (Nichols 1989) and particu-larly McHugh et al. (1997) for adult internalstructures including male and female ter-minalia. Observations of external and inter-nal morphology were made with a LeicaMZ16 stereomicroscope (magnification:7-115x) and an Olympus BX41 compoundmicroscope (magnification: 20-400x), eachequipped with an ocular graticule for mea-surements of lengths and length/width re-lations. The overall length was measuredfrom the anterior margin of the head to theposterior margin of the elytra; measure-ments of the head width and dorsal in-terocular distances follow Pakaluk (1988)and Chaves & Genaro (2005). The numbersof measurements taken are given in paren-theses for the preceding values. The habituspictures of Pharaxonotha beetles were pro-duced with a Microptics XLT imaging sys-tem. The line drawings were prepared witha drawing tube attached to the stereomicro-scope (external structures) or compoundmicroscope (internal structures). The initialsketches were scanned and redrawn usingan illustration software program, empha-sizing structures with systematic or diag-nostic significance.

The identification labels for type speci-mens include the genus name and speciesepithet, a gender symbol (for holotype andallotype only), and the authors and year.They are colored red for the holotype,green for the allotype, and yellow for allparatypes. The insect collection codens areadopted from Arnett et al. (1993), and asused as follows:

AMNH – American Museum of NaturalHistory, New York, NY

BDVC – Barry D. Valentine collection, Sara-sota, FL

FSCA – Florida State Collection of Arthro-pods, Gainesville, FL

MAIC – Michael A. Ivie collection, Mon-tana State University, Bozeman, MT

MEBT – Museum of Entomology andTropical Biodiversity of the MayagüezCampus, University of Puerto Rico, RíoPiedras, PR

MUCR – Museo de Insectos, Universidadde Costa Rica, Ciudad Universitaria,Costa Rica

NMFC – Nico M. Franz collection, Univer-sity of Puerto Rico, Mayagüez, PR

NMNH – National Museum of NaturalHistory, Washington, D.C.

NZAC – New Zealand Arthropod Collec-tion, Landcare Research, Auckland, NewZealand

UGCA – Collection of Arthropods, Univer-sity of Georgia, Athens, GA

UPRM – Invertebrate Collection, Univer-sity of Puerto Rico, Mayagüez, PR

SYSTEMATICS

Pharaxonotha portophylla Franz & Skelley,new species

Diagnosis. Pharaxonotha portophylla isreadily identified as a member of Phara-xonotha using the diagnostic features in Le-schen & Skelley (2002: 344), viz. pronotumwith lateral margins subparallel, each sideposteriorly with a short sulcus, and exter-nally visible ventrite I lacking metasub-coxal lines, among others. This new speciesis a member of a complex centered aroundP. floridana (Casey), currently under studyby PES. Pharaxonotha portophylla may bedistinguished from other described and un-described species by a combination of char-acters; i.e., body dorsally with short, indis-tinct setae; clypeus and frons flattened;head with coarse punctation, lacking su-praocular striae, posteriorly with trans-verse line anterior of stridulatory files; pro-notum slightly transverse; anterolateraledges of pronotum weakly projected, angu-late, not evenly rounded; protibia narrowlytriangularly widened toward apex; outertooth of protibial apex separated from in-ner terminal row of spinules via small gap;elytra with anterior marginal line; elytralpunctures coarse, not impressed; elytralstriae not grooved, elytral intervals plane.

PHARAXONOTHA PORTOPHYLLA SP. NOV. FROM PUERTO RICO 323

Description – male. Body in dorsal viewelongate to oval (Figures 3A); small, length2.75-3.38 mm, width 1.03-1.30 mm, greatestwidth near anterior 1/3 of elytra, length/width ratio 2.53-2.80 (N=10); in lateral viewdepressed (Figure 3B), dorsally and ven-trally slightly convex; color reddish-brown,

head with darkest pigmentation, thorax(including pronotum and legs) and ven-ter intermediate, and elytra with lightestpigmentation, light reddish-brown andweakly transparent; sculpture uniformlyand fairly densely punctulate on head andpronotum, elsewhere more sparsely dis-

FIG. 3. Habitus of P. portophylla. (A) dorsal view; (B) lateral view; (C) head, dorsal view; (D) head, ventralview.


tributed; pubescence very fine and short,appressed, aurate, moderately densely ar-ranged, longer on femora and tibiae.

Head – mouthparts. Clypeus plane, api-cally slightly rotundate, with row of fairlylong setae. Mandible (Figure 4A) large, ar-cuate-triangular; apically with 2 small, nar-rowly superposed teeth, centrally variouslysulcate, outer margin with row of setae.Maxilla (Figure 4B) with cardo apically ex-panded and setose; stipes narrowly trian-gular, apically acute, partially overlappingwith palpiger; galea and lacinia with long,densely aligned setae; maxillary palp 4-seg-mented, each palpomere with row ofsparsely arranged, subapically positionedsetae; palpomere I very small, nearly 1/4 aslong as II, transverse; II large, longer thanIII, elongate-clavate; III fairly large, nearly1/2 as long as IV, equilateral; IV elongate-oval, apically densely papillate. Labium(Figure 4C) with mentum trapezoidal to tri-angular, apically distinctly narrowed, ven-tral surface with two larger subcentral se-tae, apical margin with 2 separate strongemarginations to accommodate ligula andlabial palps, centrally with narrow projec-tion; ligula with palpiger projecting beyondmentum, paraglossae elongate-lobulate,

apically with long, densely arranged setae;labial palps 3-segmented; palpomere Ifairly small, elongate; II small, globular; IIIlarge, elongate-oval, apically papillate.Head in dorsal view conical (Figure 3C),apically gradually narrowed, plane to veryslightly convex, width 0.64-0.74 mm; dorsalinterocular distance 0.38-0.46 mm, headwidth/dorsal intraocular distance ratio1.55-1.73 (N=10); ventral interocular dis-tance 0.30-0.35 mm, head width/ventral in-terocular distance ratio 2.12-2.30 (N=10);dorsolateral margins bisinuate, occipitalstridulatory carinae present, supraocularstriae absent (cf. Chaves & Genaro 2005),punctation coarse, with short, dark,obliquely oriented subcuticular suture(apodeme) on each side above antennal in-sertion, apical margin truncate, posteriorlywith transverse line anterior of stridulatoryfiles; in lateral view slightly conical, withseparate, angulate regions for eye (poste-rior) and antennal insertion (anterior), andwith obtuse, lobulate to tuberculate, anteri-orly directed projection at anteroventraledge; in ventral view slightly concave (Fig-ure 3D), centrally with transverse linear im-pression, ventrolateral margins emarginate,apically truncate. Antennae 11-segmented,

FIG. 4. Mouthparts, ventral view. (A) left mandible; (B) right maxilla; (C) labium.


inserted laterally near anterodorsal marginof eye, length similar to pronotum + 1/2 ofhead, antennomere I (scape) fairly large,elongate, antennomeres II + III, slightlysmaller, equilateral to elongate, antenno-meres IV-VIII small, progressing sequen-tially from equilateral to transverse; club3-segmented, antennomeres fairly large,light reddish-brown, with dense setation,IX + X similar in length, clavate, XI slightlylonger, globular, slightly depressed, withfine, dense pubescence. Eyes globular, pro-truded, with as many as 11 large black fac-ets spanning from anterior to posteriormargin, anterior margin slightly oblique,nearly straight.

Thorax. Pronotum in dorsal view slightlytransverse, length/width ratio 0.75-0.80(N=10), evenly convex, anterolateral edgesweakly projected, angulate, not evenlyrounded, posteriorly on each side with 1narrow sulcus positioned at half distancebetween midline and lateral margin (cf. Le-schen & Skelley 2002: 344); posterior mar-gin slightly arcuate, centrally projected; lat-eral margins straight to slightly arcuate,subparallel, with slightly elevated carina;anterior margin nearly straight; in lateralview with pleural region concave andstrongly directed inward underneath later-al margin, subglabrate. Epipleura glabrate;with mesepisternum triangular; mesepim-eron narrowly pentagonal; metepisternumlarge, posteriorly slightly narrowed; met-epimeron small, nearly entirely covered byelytra. Prosternum in ventral view nearlyevenly convex; lateral margins anteriorlydistinctly diverging along suture; anteriormargin straight to slightly emarginate, withrow of long, anteriorly directed setae; pro-coxal cavities separated by distance similarto breadth of antennal funicle, laterallyclosed yet posteriorly open. Mesosternumshort, at the same level as metasternum(not inflected); mesocoxal cavities laterallyclosed. Metasternum long, laterally convex,centrally slightly impressed, with midlinesuture extending along posterior 3/5, atposterior end between narrowly separatedmetacoxal cavities with small, rotundateemargination. Metendosternite (Figure 5A)with lateral margins of stalk slightly di-verging; lateral arms long and narrow, di-

rected lateroventrally; dorsal furcal armsdirected dorsally at a nearly 90° to lateralarms, apically slightly narrowed; anteriortendons removed from median point bynearly 1/3 distance towards furcal arms.

Legs fairly short and slender, similar inlength and shape; procoxae small, globular;mesocoxae globular; metacoxa transverselyelongate-oval, without distinct posteriorface; trochanters apically obliquely trun-cate; femora fairly stout, laterally com-pressed, ventrally with slightly concave,glabrate surface to receive tibiae; tibiaeshorter than femora, protibia/profemurlength ratio 0.78-0.86 (N=10); protibiaslightly arcuate, compressed, marginsweakly carinate, narrowly triangularlywidened towards apex, apical marginobliquely truncate, with row of ca. 15 fairlyshort and narrow spinules, with 2 spinulesat inner (mesal) angle slightly larger, and 1small tooth at outer (lateral) edge, toothseparated from row of spinules via smallgap; tarsi 5-segmented, similar in length totibiae, tarsomeres I-IV ventrally with long,fine, densely arranged, anteriorly directedsetae; tarsomere I large, longer than II, cla-vate; II and II fairly large, similar in length,clavate; IV small, nearly 1.2 as long as III,globular; V long and narrow, slightlyshorter than II + III, apically slightly ex-panded; tarsal claws paired, simple.

Scutellum visible, transverse, posteriormargin rotundate. Elytra in dorsal viewelongate-oval, length/width ratio 1.72-1.90(N=10), greatest width near anterior 1/3;anterior margins nearly straight, with nar-row anterior marginal line (cf. Chaves &Genaro 2005), angulate at midline; humerismall (not protruded), convex; lateral mar-gins slightly arcuate, subparallel along an-terior 1/2, posteriorly slightly converging,more strongly converging along posterior1/4; posterior margins subcontiguous; inlateral view slightly and evenly convex; 10complete striae + 1 short scutellary striole(with 8-10 punctures) extending along an-terior 1/4 near midline; striae I-IX nar-rower than intervals, not grooved, punc-tures coarse, not impressed, subcircular,centrally with 1 short seta, slightly irregu-larly spaced, size slightly increasing to-wards lateral margins; stria X nearly obso-


lete along anterior 1/3; elytral intervalsplane. Wings (Figure 5B) slightly longerthan body, wing/body ratio 1.05-1.13(N=4), elongate to oval, length/width ratio2.58-2.80 (N=4), greatest width near centralregion; anterior margin slightly sinuate,posteriorly slightly arcuate, with separateanal region along basal 1/6 and slight in-curvation near apical 2/5; all major veinspresent (compare with McHugh et al. 1997,

Chaves & Genaro 2005), except for anal cellwhich is indistinct; radial, medial, and api-cal fields with maculations; macrosetaedensely arranged along apical 1/2 of Sc,more sparsely arranged along posteriormargin (absent in anal region); microsetaedistributed densely throughout surface.

Abdomen. Venter in ventral view with 5visible segments; anterior margin mediallywith distinct acute-triangular projection be-

FIG. 5. Thorax. (A) metendosternite, posterior view; (B) right hind wing.


tween metacoxal cavities, lateral edgesslightly projected, lateral and posteriormargins arcuate-rotundate, posteriorlyconverging; ventral segments separated bystraight to laterally slightly arcuate line, atthe same level (not inflected); ventral seg-ment I longer than II, metasubcoxal linesabsent (cf. Leschen & Skelley 2002); II to IVsimilar in length; V slightly longer than IV,posterior margin with row of short to fairlylong, appressed, densely arranged setae.Pygidium in dorsal view transversely rect-angular, lateral margins slightly arcuate,posterior margin nearly straight, with sev-eral rows of short to long, sparsely ar-ranged, suberect setae; in lateral viewslightly longer than ventral segment V,slightly convex, narrow, parallel to orienta-tion of tergum.

Terminalia. Proctiger similar in length totergum VIII, anterior margin evenly emar-ginate, anterior edges projected, lateral andposterior margins slightly arcuate, withshort, densely arranged setae. Paraproctsposteriorly expanded, triangular, posteriormargin straight-truncate, connected viamembrane to small, rectangular, anteriorlyacutely narrowed sternum IX; spiculumgastrale with two lateral, long and narrow,anteriorly converging and connectedapodemes. Tegmen (Figure 6) with anteriorregion (2/5) ring-like, posterior region (3/5) sheath-like (slightly convex, enclosingmedian lobe), weakly sclerotized, triangu-lar, centrally with large oval perforation toreceive median lobe; lateral margins poste-riorly gradually converging; posterior mar-gin slightly arcuate, dorsally with 2 large,elongate, subparallel, apically setoseparameres. Aedeagus (Figure 6) with me-dian lobe in dorsal view elongate l/w =4.5-4.8 (N=3), lateral margins subparallel toslight sinuate, apically narrowed, rotun-date, symmetrical; apical 1/2 more stronglysclerotized; in lateral view slightly de-flexed, basal 3/5 very narrow (laminar),apical 2/3 broader, with distinct internalspace, dorsal margin narrowed at apex,acute; internal sac membranous, with verylong, variously wound flagellum; aedeagalapodemes, distinctly longer than medianlobe, narrow and nearly straight; originat-

ing from lateral margins, thereafter adja-cent to each other.

Description – female. Similar to male (seedescription above); with exception of thepygidium which is more strongly convexand very narrow, with the lateral marginsposteriorly gradually converging and with1 long, suberect setae near the posterior edge.

Terminalia (Figure 7A). Paraprocts (ter-gum IX) with oblique longitudinal separa-tion, lateral regions more strongly sclero-t i z e d . E p i p r o c t s ( t e r g u m X ) a n dgonocoxites triangular, posteriorly gradu-ally narrowed, apices of gonocoxites later-ally with concave impression and variouslonger, suberect setae; gonostyli large, nar-row, posteriorly diverging and veryslightly broadened, with several shorter se-tae and 1 long seta. Vulvar tube membra-nous, posteriorly expanded. Spermatheca(Figure 7B) fairly narrow, breadth similarthroughout; deflected by nearly 180°, C-shaped, basally and apically convex; ductstrongly wound.

Variation. Relatively little external mor-phological variation is apparent in the ex-amined specimens, apart from the differ-ences in size, a slightly lighter and moreheterogeneous pigmentation in teneralspecimens, and variation in the shape,depth, and color of the elytral punctures.

Type material. Male holotype “PuertoRico (USA), Bosque Estatal Susúa, Que-brada Peces, 800 ft., N 18°04.32’, W66°54.60’/ ex cones Zamia portoricensis, leg.N. Franz, I-15-2007” (UPRM); female allo-type, same label information as male holo-type (UPRM); paratypes (unsexed), samelabel information as male holotype(AMNH, 5; FSCA, 10; MEBT, 8; NMNH, 8;UGCA, 5); “Puerto Rico (USA), Bosque Es-tatal Susúa, Quebrada Peces, 400 ft., N18°03.75’, W 66°54.43’/ ex cones Zamia porto-ricensis, leg. N. Franz, I-12-2007” (MUCR, 8;NZAC, 8); “Puerto Rico (USA), Bosque Es-tatal Cambalache, trails near entrance, 120 ft.,N 18°27.15’, W 66°35.84’/ ex cones Zamia am-blyphyllidia, leg. N. Franz, II-12-2007” (FSCA,3; NMFC, 5; UPRM, 5); “PUERTO RICO: Res.For. Cambalache, 20 km. e. Arecibo, 1-iii-‘96,ex male cycad cone, coll. B&B Valentine”(BDVC, 2; FSCA, 4; MAIC, 3).

Etymology. The species epithet makes


reference to the two putative host plant spe-cies Z. portoricensis (“porto-“) and Z. ambly-phyllidia (“-phylla”) on which the examinedspecimens of P. portophylla were collected.

NATURAL HISTORY

Adults of Pharaxonotha portophylla werecollected consistently on the microstrobiliof Z. amblyphyllidia and Z. portoricensis,

with the greatest numbers of individualspresent at the stage of anthesis (Table 1).Neither adults nor larvae were obtained atthe earlier stage of pre-anthesis, suggestingthat odor production is critical to this spe-cies’ attraction to the cones. As many as 30small to medium-sized larvae (Fig. 8A)were retrieved from a single pollen-releasing microstrobilus of Z. portoricensisat a late stage of anthesis. Presumably theseimmature individuals had emerged from

FIG. 6. Male terminalia; tegmen and aedeagus. (A) ventral view; (B) lateral view.


eggs laid by early-anthesis visitors, and un-derwent several molts in a period of 2-3days. The number of adults sampled onpost-anthesis microstrobili was low (range:0-4 individuals); it remains unknown

whether these individuals had stayed onthe cones since arriving at anthesis, orwhether they recently emerged from thepupal stage. Conversely, the post-anthesismicrostrobili yielded the highest abun-

TABLE 1. Abundance of adults and larvae of P. portophylla on micro- and megastrobili of Z. amblyphyllidia andZ. portoricencis at the Susua and Cambalache Commonwealth Forest Reserves, respectively, during three stagesof development: pre-anthesis, anthesis, and post-anthesis (see also text). Mean values are provided ± standarddeviation, with the corresponding numbers of cones sampled shown in parentheses.

Zamia amblyphyllidia (Cambalache)Microstrobili (�) Megastrobili (�)

Adults Larvae Adults Larvae

Pre-anthesis 0.0 ± 0.0 0.0 ± 0.0 — —(N = 2) (N = 2)

Anthesis 5.5 ± 0.7 2.0 ± 2.8 0.3 ± 0.5 0.0 ± 0.0(N = 2) (N = 2) (N = 4) (N = 4)

Post-anthesis 1.0 ± 1.7 3.2 ± 3.3 — —(N = 6) (N = 6)

Zamia portoricensis (Susua)Microstrobili (�) Megastrobili (�)

Adults Larvae Adults Larvae

Pre-anthesis 0.0 ± 0.0 0.0 ± 0.0 — —(N = 4) (N = 4)

Anthesis 5.4 ± 4.6 4.0 ± 8.3 0.2 ± 0.4 0.0 ± 0.0(N = 12) (N = 12) (N = 6) (N = 6)

Post-anthesis 0.2 ± 0.4 12.2 ± 15.3 — —(N = 10) (N = 10)

FIG. 7. Female terminalia. (A) oviposition structures (anterior to posterior): paraprocts, epiprocts, gonocoxites,and gonostyli, dorsal view; (B) spermatheca.


dances of P. portophylla larvae, with a maxi-mum of 45 individuals on a single Z. por-toricensis cone.

Repeated sampling of receptive mega-strobili produced only one adult specimenof P. portophylla on Z. amblyphyllidia cones(N = 4) and on Z. portoricensis cones (N = 6),respectively (Table 1). No larvae were ob-served on the megastrobili. Overall,samples of Z. amblyphyllidia cones yieldedlower numbers of individuals than Z. por-toricensis cones.

The reproductive behavior of Pharaxono-tha portophylla beetles is difficult to observedirectly on Zamia strobili, due to their rela-tively low density, low apparent levels ofactivity at any given time, and cryptic loca-tion between the bases of the sporophylls.The adults are rarely seen on the surface,exiting only after the cones are repeatedlyperturbed or cut off. Nevertheless, exami-nation and rearing of microstrobili indi-cates that beetles mate and oviposit fre-quently, and that the larvae develop whilefeeding on the parenchyma of the sporo-phylls (Fig. 8B).

Other insects visitors sampled on Zamiacones were predominantly caterpillars ofunidentified species of snout moth (Lepi-doptera: Pyralidae), with as many as 80 in-dividuals on the post-anthesis microstrobiliof Z. amblyphyllidia (average 22.6 ± 25.3 in-dividuals on cones where present, N = 10),and up to 30 individuals on Z. portophylla(average 5.5 ± 7.5 individuals, N = 15).These lepidopteran species were not ob-

served on the megastrobili. Additional spo-radic visitors of rotting Zamia microstrobiliincluded myrmecine ants, drosophilid flies,bark beetles, and diplopods. These visitorswere not considered to play a significantrole in the plants’ reproduction.

DISCUSSION

Systematics

Pharaxonotha portophylla is the secondspecies assigned to this ancient lineage inthe Caribbean region (see Chaves & Genaro2005), thereby raising the number of knownspecies in the genus to 13 worldwide (Le-schen 2003). The new species furthermorerepresents a first record for Puerto Ricowhich has a depauperate erotylid fauna(Wolcott 1948). Pharaxonotha portophylla ap-pears most similar to P. floridana by virtueof a shared coarse punctation on the headand anterolateral angulation of the prono-tum. However, P. portophylla is readily dis-tinguished from other congeneric speciesby a weakly transverse pronotum and shal-lowly impressed strial punctures; P. flori-dana has a more strongly transverse prono-tum and distinctly impressed strialpunctures. Both P. floridana and P. portophyllaare restricted to Zamia host species in Floridaand Puerto Rico, respectively (Tang 1987, Le-schen & Skelley 2002). Geographically, P. por-tophylla is closest to the Cuban P. esperanzaeChaves & Genaro (2005), yet the latter speciesis unique among members of this complex in

FIG. 8. Development of P. portophylla on Z. portoricensis. (A) habitus of early, intermediate, and advancedlarval stages; (B) cross-section of microstrobilus at approximately three weeks past anthesis, with rotting mi-crosporophylls showing traces of damage (white arrows) produced by feeding P. portophylla larvae.


having supraocular striae, lacking an anteriormarginal line on the elytra, and in its asso-ciation with the endemic Cuban cycad Micro-cycas calocoma (Miquel) A. de Candolle(Chaves & Genaro 2005).

At least ten new species of Pharaxonothafrom the Caribbean archipelago, Mexicoand Central and South America are presentin research collections. Their systematictreatment is needed to establish a morecomplete picture of the relationshipsamong the Neotropical species in the genus.

Natural History

The sampling of insect visitors on conesof Z. amblyphyllidia at Cambalache and onZ. portoricensis at Susúa strongly suggeststhat P. portophylla is the primary and solepollinator of these cycad populations.Wind pollination, an early paradigm in cy-cad reproductive ecology (cf. Stevenson etal. 1998), has since been widely discardedin favor of insect pollination (e.g., Hall et al.2004, Oberprieler 2004, Tang 2004). It ishighly unlikely that anemophily can ac-count for any apparent seed set in the ob-served Zamia plants (Negrón-Ortíz et al.1996; NMF, personal observation). With re-gards to insect pollen vectors, P. portophyllawas the only agent present on open conesof either gender, as well as the only specieswhose adults were visibly covered withpollen and located inside the microstrobiliat the time of anthesis. The temporal andbehavioral synchronization of Pharaxonothabeetles with their cycad hosts is well docu-mented (e.g., Tang 1987, Norstog et al.1992, Chaves & Genaro 2005). Other insectvisitors of Zamia cones at the two siteslacked these attributes and thereforeshould not be regarded as pollinators.

The similar odors of male and femalestrobili yet differential rates of visitationand reproduction in the beetles may be in-dicative of “pollination by deceit” (Terry etal. 2004), where megastrobili contain sec-ondary metabolites that counteract ovipo-sition and larval growth (cf. Schneider et al.2002). However, the chemical ecology ofthe P. portophylla/Zamia interaction in Puer-to Rico remains entirely unexplored. Ahandful of one-minute probes of the central

axes of Z. portoricensis strobili (� and �)with a NiCr-Ni thermocouple element(Ahlborn) revealed no thermogenetic activ-ity (NMF, personal observation).

Species of Pharaxonotha and members ofthe Parallocorynus-Rhopalotria weevil lin-eage (Marvaldi et al. 2006) have evidentlyplayed a critical role in the evolutionary ra-diation of cycads in the Neotropical region.This contribution reduces one of many sig-nificant gaps in our knowledge of a long-standing coevolutionary interaction thatshould receive more attention from system-atists, chemical ecologists, and managers ofnatural resources.

Acknowledgments.—We thank GaryBreckon (University of Puerto Rico at May-agüez) for invaluable assistance in locatingpopulations of Zamia species with receptivecones at various Puerto Rican sites, partici-pating in the sampling process, providinghabitus pictures of the host plants, for in-formation and discussions on their repro-ductive biology, and for reviewing themanuscript. Insightful reviews by RichardLeschen and James Robertson led to an im-proved taxonomic account of P. portophylla.The Puerto Rican Department of NaturalResources and Environment (DRNA) pro-vided collecting permits for the Cambal-ache and Susúa habitats. This research wassupported in part through a Seed Moneygrant awarded to the senior author by theResearch & Development Center of theUniversity of Puerto Rico at Mayagüez.This is Florida Department of Agricultureand Consumer Services, Division of PlantIndustry, Entomology Contribution No.1090.

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pharaxonotha portophylla (coleoptera: erotylidae), new species

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