Department of Entomology, Comstock Hall Cornell University, Ithaca ...

The Coleopterists Bulletin, 56(1):50–67. 2002.

FIRST REPORT OF IMMATURES, GENITALIA AND MATERNAL CARE IN EUGENYSA COLUMBIANA (BOHEMAN) (COLEOPTERA: CHRYSOMELIDAE: CASSIDINAE: EUGENYSINI) CAROLINE S. CHABOO Department of Entomology, Comstock Hall Cornell University, Ithaca, NY 14853, U.S.A. Abstract Illustrated descriptions of larvae, pupae and adult genitalia are provided for Eugenysa columbiana (Boheman) in Costa Rica. These represent the first such descriptions for the tribe Eugenysini. Females are reported as exhibiting guarding of larvae and pupae. Male presence with the female and larval group is also described. The distribution range is extended to Peru, and host plant data is recorded. A new character from the spermatheca is presented and its potential usefulness in deducing eugenysine relationships is discussed.

Resu´men Se presentan descripciones ilustradas de larvas y pupas y genitalia de adultos para Eugenysa columbiana (Boheman) en Costa Rica que constituyen el primer registo para la tribu. Se reporta el cuidado maternal de larvas y pupas. Se describe tambień la presencıá del macho con la hembra y el grupo de larvas. El rango de distribucioń se extienda a` Peru y se presentan datos de la planta hospedejera. Se presenta un nuevo caracter de la spermateca y se discute uso el potencial para clarificar las relaciones filogeneticas de los escarabajos eugenysines.

Eugenysa columbiana (Boheman) is a member of the Neotropical tribe Eugenysini which consists of 33 species in three genera, Agenysa Spaeth (8 species), Eugenysa Chevrolat (19 species), and Miocalaspis Weise (6 species) (Blackwelder 1946; Seeno and Wilcox 1982; Borowiec 1999). The tribe extends from Argentina and Brazil to Mexico with one species, Agenysa guianiensis (Boheman), also recorded on the Caribbean island of Curaçao (Borowiec 1999). Viana (1968) revised the Eugenysini, with good keys and descriptions for adults of the species. Descriptions of genitalia and immatures were not included then and are still undescribed. Eugenysines are among the largest and showiest cassidines yet their natural history is poorly known. Host plants for only two species have been recorded: an Ipomoea sp. (Convolvulaceae) for Agenysa boliviana Spaeth (Viana 1968) and Mikania guaco Bonpland (Asteraceae) for Eugenysa coscaroni Viana (Windsor and Choe 1994). Maternal care has been reported in E. coscaroni (Windsor and Choe 1994). This remarkable behavior is a rare phenomenon in the Chrysomelidae; among the approximately 40,000 species in the family (Lawrence 1982), only 22 species in 8 genera from the subfamilies Chrysomelinae and Cassidinae have been reported as maternal care providers (Table 1). Knowledge of these species ranges from fairly detailed observations (Windsor 1987; Choe 1989; FrieiroCosta 1995; Kudo and Ishibashi 1995; Kudo et al. 1995), to brief notes (Monte 1932; Preston-Mafham 1993) or just a single photograph (O’Toole and Preston-Mafham 1985). In all these species, females show a stereotypical behavior 50

THE COLEOPTERISTS BULLETIN 56(1), 2002

51

Table 1. Comparison of maternal-care providing species of Chrysomelidae. A 5 viviparous (Chrysomelinae only); b 5 guarded; c1 5 shield present (Cassidinae only); e 5 male present with brood; ? 5 plant species not in Index Kewensis (1997); blank cells indicate missing information. Egg CHRYSOMELINAE Pterodunga miribile Daccordi

Larva

Pupa

Host plant

Reference

b

—

Buckinghamia celcissima Muell.; B. ferruginiflora Foreman & Hyland; Grevillea baileyana McGillivray (Proteaceae) Populus tremula Linn.; Salix spp. (Salicaceae) (Jolivet 1988)

C. Reid, pers. comm.

Gonioctena rufipes (Degeer)

a

b

—

Lu¨hmann 1940; Goidanich 1956

Gonioctena viminalis Linnaeus Gonioctena japonica Chujo & Kimoto Gonioctena sibirica Weise Labidomera suturella Chevrolat

a

b

—

a

b

—

Alnus spp. (Betulaceae)

Kudo & Ishibashi 1995

a

b

—

Salix bakko Kimura (Sal.)

Kudo et al. 1995

b

—

Witherigia hetero- Choe 1989 clita? (Solanaceae)

CASSIDINAE: Acromis sparsa (Boheman) Acromis spinifex (Linneaus) Paraselenis (Spaethiechoma) flava Linnaeus Paraselenis (S.) solieri (Boheman) Paraselenis (S.) dichroa (Germar) Paraselenis (S.) tersa (Boheman) Omaspides (Omaspides) bistriata Boheman

STOLAINI b b, c1

b

b

b, c1

b

b

b

b

Merremia umbellate (L.) (Convolvulaceae) Ipomoea batatas P.; Ipomoea spp. (Conv.) Convolvulus sp. (Conv.)

Jolivet 1988

Windsor 1987; PrestonMafham 1993 Fiebrig 1910; Buzzi 1980; PrestonMafham 1993 Weyenberg 1874

b

I. batatas P. (Conv.)

Monte 1932; Buzzi 1988

b

I. batatas P. (Conv.)


I. tiliacea (Willd.) (Conv.)

Windsor & Choe 1994

I. philomega H. (Conv.)

Windsor & Choe 1994

b

b

b

b

b

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THE COLEOPTERISTS BULLETIN 56(1), 2002 Table 1.

Continued. Egg

Omaspides (O.) convexicollis Spaeth Omaspides (O.) nigrolineata (Boheman)

b

Omaspides (O.) pallidipennis (Boheman) b Omaspides (O.) tricolorata (Boheman) b Omaspides (P.) sobrina (Boheman) Omaspides sp. b Cyrtonota thalassina (Boheman)

Larva

b

Pupa

Host plant

b

I. philomega H. (Conv.)

b

I. batatas P. (Conv.) Passiflora sp. Passifloraceae) (von Lengerken 1939); I. alba L. (Buzzi 1994) (Conv.)

Ohaus 1899–1900; Costa Lima 1914, 1955; von Lengerken 1939

I. alba L. (Conv.)

Frieiro-Costa 1995; Frieiro-Costa & Vasconcellos-Neto 2000

b

b, c1

b

b

b, c1

b

b

CASSIDINAE: EUGENYSINI Eugenysa columbiana (Boheman) b, c1, e b Eugenysa coscaroni Viana b b, c1, e b

Reference

Rodriguez 1994a


Jolivet 1988 O’Toole & PrestonMafham 1985; O’Toole 1995 I. batatas P. (Conv.) (Costa Lima, 1936)

Ohaus 1899–1900; von Lengerken 1939

Mikania sp. (Asteraceae)

Chaboo, this paper

M. guaco Bonpl. (Aster.)

Windsor & Choe 1994

of sitting on groups of eggs or larvae, and responding to intruders or to objects introduced by the observer. General ideas about the origins of maternal care in beetles (Brandmayr 1992) and specifically in chrysomelids (Windsor and Choe 1994) have been discussed. Windsor and Choe (1994) also reported that males of E. coscaroni were sometimes present with the larval group but they concluded that male presence was probably related to mate guarding. The possibility of paternal care has arisen from an illustration that shows a male Acromis spinifex (Linnaeus) sitting on an egg mass (von Lengerken 1939); this image has been reproduced in the literature (Linsenmaier 1972; Jolivet 1997) thus perpetuating the idea of male involvement in brood care (Jolivet 1997; Buzzi 1998). This paper reports the first observations of maternal care in E. columbiana from Costa Rica. The presence of adult males with females and larval groups is also reported. Distributional data, a range extension to Peru, and the first host plant record are provided for the species. Illustrated descriptions of the


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reproductive system, larvae and pupae are provided, representing the first such descriptions in the tribe Eugenysini. The implications of morphology and behavior in E. columbiana are discussed. Materials and Methods Aggregations of E. columbiana were observed at two sites in southern Costa Rica, the Las Cruces Biological Station and near the Rio Penãs Blancas, during June 2000. Specimens were collected and reared, under license number A-00200 from the Ministerio del Ambiente y Energia (MINAE). Voucher adult and larval specimens of E. columbiana have been deposited in the collections of CUIC and INBio; dissected vouchers are deposited in the AMNH collection. Insect identifications were made by comparisons with previously identified specimens from the collections listed below but types were not examined. Plant identification was by L. D. Gomez (Wilson Botanical Garden). Cassidine species names follow Borowiec (1999), and plant species names are in accordance with the Index Kewensis (1997). Larval age was determined by counting head capsules in the exuvio-fecal shield. Locality data are stated as on specimen labels. Genitalic study in this species involved both mature and teneral adults that were recently killed and preserved in alcohol. Specimens were dissected in distilled water; potassium hydroxide (KOH) was avoided as it almost instantly destroys the spermathecal muscles and, very quickly thereafter, the softer parts of the female reproductive system. Comparative genitalic studies involved older museum specimens of nine additional eugenysine species: A. caedemadens (Lichtenstein), A. guianiensis, A. connectens (Baly), A. parellina (Boheman), E. colossa (Boheman), E. coscaroni, E. grossa (Linnaeus), E. martae Borowiec, E. unicolor Borowiec and Dabrowska, and M. gentilis (Erichson). Spermathecal terminology follows Samuelson (1966). Specimen examinations and illustrations were made with a Wild M5A microscope and a Zeiss compound microscope, both with camera lucida attachments. The aedeagus, spermatheca, larvae and pupae were photographed with a Nikon D1 Digital camera with an Infinity K2 lens attachment and Microptics fiber optic flash unit. Measurements were taken with a digital stage micrometer; length was measured along the dorsal midline, and width across the metanotum. The following collections and individuals provided specimens for this study: AMNH, American Museum of Natural History, L. Herman and S. Lodhi; BMNH, British Museum of Natural History, M. Brendell and S. Shute; CSCC, Caroline S. Chaboo Collection; CUIC, Cornell University Insect Collection, J. K. Liebherr and R. Hoebeke; INBio, Instituto Nacional de Biodiversidad, A. Solı´s; MCZC, Museum of Comparative Zoology Collection, P. Perkins; MMUE, University of Manchester Collection, C. Johnson and P. Rispin; UCR, University of Costa Rica, P. Hanson. Field Observations On June 6, 2000, in the vicinity of the Rio Penãs Blancas in southern Costa Rica, I found two cassidines species, Stolas lebasii (Boheman) (Cassidinae: Stolaini) (one adult) and E. columbiana, on a woody vine that was wrapped around a small tree. Leaves of the vine were well-chewed, with large holes along margins and within the leaf surface (Fig. 1). The vine was later identified as a Mikania sp. (Asteraceae).

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Fig. 1.

Leaf of Mikania sp. showing feeding damage by E. columbiana.

Fig. 2.

Adult of E. columbiana attending pupal group on Mikania sp. in Costa Rica.

The specimens of E. columbiana consisted of 6 tightly-clustered larval broods (16 to 111 individuals) and 12 adults. There were two egg clusters, all with the upper portions broken, probably from hatching. Each cluster of ovalshaped whitish eggs encircled the stem, and individual eggs were attached to the stem at one end; this arrangement corresponds to the egg mass of E. coscaroni (Windsor et al. 1992). Larvae fell into two distinct size classes, and each group consisted of only one size class. No solitary larvae were observed outside groups or actively feeding. Six adults were in mating pairs that remained in copula for the entire period of observation. In addition there were two singletons, and two pairs of adults associated with the two largest larval broods. In one group, one adult sat atop larvae at one end while the other adult sat at the opposite end with its forelegs on larvae. In the second group, each adult sat opposite each other on the vine, both with the posterior part of their bodies covering larvae. At one point, the sole S. lebasii individual stepped on the elytra of an adult E. columbiana, spent about 10 min touching with the antenna and legs, and eventually moved on. Later it walked over one larval group and was quickly dislodged into the bushes below by the repeated and rapid flexing movements of exuvio-fecal shield. Three muscoid flies landed on one larval group but were unable to settle because individual larvae, along with their neighbors, quickly flexed their shields. I disturbed the vine by first lightly shaking and then by more violent shaking. Larvae responded with the reflex abdominal movements (a behavior called cycloalexy), single adults dropped-off into the bushes (this behavior is called thanatosis and is the usual reflex of cassidines when disturbed). Adults attending larvae did not move.


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Two larval groups were collected, one untended group of smaller-sized larvae (111 4th instar individuals), and one group of larger-sized larvae (42 5th instar individuals) along with its two attendant adults. When handled, both adults and larvae exuded an orange-brown liquid with a faint odor from the mouth. The adults were not obviously sexually dimorphic, however later dissections revealed them to be male and female. On June 11, 2000 at the Las Cruces Biological Station I observed another group of E. columbiana, a single adult on top of a group of 32 pupae (Fig. 2). When lightly touched with a pencil the pupae flexed their bodies, and the adult moved about but remained with the group. Rearing experiments. Eight large-sized larvae were maintained in a Tupperware y container with a moist cotton ball for one week. They continued eating Mikania leaves thus confirming that the vine is a host plant. In the container, larvae spread out for feeding but regrouped when not active. Fecal shields were quickly flexed over the abdomen whenever I brought a forceps within a few cm of the group. These specimens died before molting into pupae. Three pupae were collected and maintained in a Tupperwarey container, with a piece of the vine and a moist cotton ball. Like pupae in the field, these individuals responded to light touch with a reflex jerking motion during nine days of observation, then were unresponsive for two days prior to adults eclosion. Newly-emerged adults were very mobile. Descriptions Eugenysa columbiana (Boheman) Calaspidea columbiana Boheman, 1850:212 (original description); Boheman, 1856:35 (catalog); Boheman, 1862:91 (catalog); Gemminger and Harold, 1876:3629 (list); Wagener, 1877:53 (list); Spaeth, 1905:91 (description); Spaeth, 1914:31 (catalog); Spaeth, 1937:85 (description). Calaspidea columbina: Champion, 1893:135 (localities; misspelling); Borowiec, 1999:79 (cited as misspelling). Eugenysa columbiana: Blackwelder, 1946:737 (checklist); Viana, 1968:21 (description); Wilcox, 1975:150 (checklist); Windsor and Choe, 1994:113 (distribution); Borowiec, 1996:184 (list); Borowiec, 1999:79 (catalog). Description. Male genitalia (Fig. 3). Median lobe forming 908 angle with basal piece; ejaculatory guides present; tegmen with manubrium (5 basal piece) as long as arms (5 lateral lobes); lateral plates slightly projected from rim of apical foramen. Ejaculatory duct very long, about 8 times longer than aedeagus; junction of duct and seminal vesicle with sclerotized protuberant knob; seminal vesicle wide distally and tapered proximally, proximal portion deflexed and tapered into vas deferens; duct coiled, coils loose proximally and tight distally. Female Genitalia Ovipositor. spiculum lightly sclerotized, with emarginate apex, and widened distally; palpi shorter than T8. Reproductive tract. Whitish; each ovary with 11–15 ovarioles; calyx, lateral oviducts and common oviduct enclosed in layer of stringlike tissue; bursa copulatrix rounded; oviduct section distal to bursa copulatrix with paired lateral lobes and short paired mesal projections; mesal projections yellowish. Spermatheca (Figs. 4–5). Well-sclerotized, falcate, with muscle on inner margin; pump with diameter smaller than in receptacle, deflexed and apically narrowed; appendix present. Receptacle complex, with three chambers distinguishable externally. Gland present, elongate and basally positioned; spermathecal duct and gland entry into receptacle chambers clearly separated, with spermathecal duct entering chamber 1 and gland entering chamber 2. Duct very long, more than 15 times spermathecal length, entirely tightly coiled.

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Fig. 3.

Photograph of aedeagal system of E. columbiana.

Larva (Figs. 6–12), 5th instar: length 10.8–14.0 mm; width at greatest diameter 6.4– 8.0 mm, (n 5 10). Body color generally yellow, tergum greyish yellow, dorsum with faint yellow medial stripe; thoracic sterna whitish; head, pronotum, legs, scoli and caudal furcae brown. Shape ovate, widest across metathorax, dorso-ventrally compressed; 14 pairs of dorso-lateral scoli and 1 pair caudal furcae present. Dorsal surface densely microtrichiate; setae present, sparse medially, more dense in pleural area. Ventral surface densely microtrichiate; setae present; setae denser, longer and finer on medial surface of thoracic segments; sternal, tergal and pleural areas with short sparse setae. Spiracles annulate. Head: exposed in dorsal view, rounded; dorsal surface with sparse setae; asperities absent. Epicranial suture with stem short, endocarina incomplete, not reaching frontoclypeus. Stemmata convex, six in two rows, laterally positioned; ventral row smaller than dorsal row. Antenna with two ring-like sections, set in membrane. Frontoclypeus triangular, longer than wide. Labrum (Fig. 8) hemispherical, with distal medial emargination; exterior surface with single paired lateral setae on postero-lateral margin, spiny processes absent; interior surface with single paired medial setae and single paired antero-lateral setae. Mandible (Fig. 9) palmate, mesal margin quadri-dentate, with 1st and 4th teeth smaller than 2nd and 3rd. Maxilla (Fig. 10) with cardo longer than stipes, setae sparse; palpus two-segmented, segment 2 with conical sensory papilla; mala undifferentiated, with apical setae. Labium (Fig. 10) with palpus one-segmented; basal plate fused, ring-like; ligula inconspicuous; hypopharynx densely asperate. Epipharynx densely microtrichate. Thorax. Pronotum pigmented, overlapping head; margin with three paired scoli, three on each side, two antero-laterally positioned and paired, third posterolaterally positioned; mesonotum with two lateral scoli; metanotum with one pair scoli


57

Fig. 4. Photograph of spermatheca of E. columbiana (spermathecal muscle removed by KOH).

on postero-lateral angles. Pronotal anterior scoli and mesonotal anterior scoli shortest. Spiracles annulate, prominent, projecting, without setae on inner margin. Legs (Fig. 11) four-segmented, with sparse long setation; tarsungulus curved. Abdomen. Ten-segmented; segments 1–8 with paired lateral scoli; 1–7 with paired annulate spiracles in dorsal lateral lines. Lateral scoli (Fig. 12) curved, tapered; scoli on segments 1–2 anteriorly directed; scoli on segments 6–7 latero-posteriorly directed; scoli on segments 8 posteriorly directed and not visible in dorsal view. Scoli on segments 6–8 long, almost twice longer than abdominal scoli. Caudal furca (Fig. 13) paired, long, unbranched, tapered, dorsally projected; surface wrinkled, setae present, asperities absent; basal plate narrow, sclerotized. Thoracic and abdominal scoli with surface slightly wrinkled and setose; setation distinct, with basal portion long and bristle portion stiff and barb-like. Shield. Present, welldeveloped up to 7 mm in height, composed of feces and exuviae with four intact head capsules. Larva (Fig. 14), 4th instar: length 6.3–8.7 mm (n 5 10). Body color light yellow, without grey cast. Scoli number and arrangement, setation and trichiation pattern, and fecal shield with characteristics similar to 5th instar larvae. Shield (Fig. 15): present, well-developed, composed of feces and exuviae with three intact head capsules. Pupa (Figs. 16–17), length 13.0 mm (n 5 1). Color dark brown to black, dorsum with narrow medial cream-colored stripe extending from anterior pronotal margin to 1st abdominal tergum. Integument smooth, devoid of setation or trichiation. Head hidden by pronotum, not visible in dorsal view. Pronotum with well-developed explanate margin, anterior edge continuous; dorsal surface feebly wrinkled, without dorsal or lateral projections. Abdominal segments 1–3 with margins laterally extended into short lobes; other segments with rounded margins; abdominal segment 9 without urogomphi. Spira-

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Fig. 5.

Spermathecal system of E. columbiana.

cles elevated. Exuvio-fecal shield retained ventrally, compressed and forming point of attachment to stem of vine.

Distribution. Previously published localities for this species include Colombia, Costa Rica and Panama (Champion 1893; Viana 1968; Borowiec 1999). Viana (1968) provided localities for 17 specimens. One specimen (MCZ) examined for this study was collected in Peru, a new country record. Material Examined. 111 fourth instar larvae, 42 fifth instar larvae, 5 pupae, and 34 males and females were examined with the following label data: No data: Bowditch coll. (1, MCZ); Columbia: (2, MCZ); Cartagena, 1871, coll. Dr. Maak (1, MCZ); Panama: (1, MCZ); Peru: Lima (1, MCZ); Costa Rica: Provincia Puntarenas: Corcovado National Park, Osa Peninsula, 21–30 Nov 1977, coll. D. H. Janzen (3, INBio); Miramar, 5 May 1977, coll. F. Vargas (1, UCR); Est. Queb. Bonita, 50 m, Res. Biol. Carara, coll. E. Bello, 17 Mar30 Apr (no year), coll. P. Campos (1, INBio); Est. Queb. Bonita, 50 m, Res. Biol. Carara, June 1990, coll. E. Rojas (1, INBio); Est. Quebrada Bonita (Agui-


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Figs. 6–13. Larva of E. columbiana, 5th instar. 6) Ventral view, scale bar 5 1 mm; 7) Dorsal view, scale bar 5 1 mm; 8) Labrum, ventral view, scale bar 5 0.25 mm; 9) Mandible, dorsal view, scale bar 5 0.25 mm; 10) Maxilla and labium, ventral view (half shown), scale bar 5 0.25 mm; 11) Proleg, laterosternal view, scale bar 5 0.5 mm; 12) First abdominal scoli, dorsal view, scale bar 5 0.5mm; 13) Caudal furca, posterior view, scale bar 5 0.5 mm.

rre), R. B. Carara, A. C. Pacifico Central, 100 m, Oct 1994, coll. R. Guzmań (1, INBio); Est. Quebrada Bonita, R. B. Carara, 100 m, Feb 1995, coll. R. Guzmań (1, INBio); La Palma Coopeunioro, Alfrededores del Albergue Unioro, 250 m, 13–22 Mar 1995, coll. L. Angula (2, INBio); Cerro de Oro, ACOSA, 270 m, 14–18 Mar 1995, coll. M. Madrigal (1, INBio); 300 m, 4–5 Nov 1995, coll A. Azofeifa (2, INBio); Est. Agujas, Sendero Zamia, 300 m, 26– 30 Nov 1995, coll. A. Azofeifa (1, INBio); Est. Agujas, Sendero Zamia, Rio Agujas, 300 m, 9–8 Mar 1996, coll. A. Azofeifa (1, INBio); Est. Rio Bonito, 2.3 Km al O. del Cerro la Gamba, 110 m, 10–30 Apr 1996, coll. E. Fletes (1, INBio); San Vito, Las Cruces Biol. Stn., ex. Mikania sp., 12 June 2000, coll. C. S. Chaboo (3 pupae; CSCC); Provincia Cartago: Parque del Este, Tres Rios, 11 July 1990, coll. J. Segura (1, UCR); Provincia San Jose´: Parque del Este, June 1990, coll. J. Seguro (1, UCR); Rio Penãs Blancas, north bank, ex. Mi-

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Figs. 14–15.

Larva of E. columbiana. 14) 4th instar; 15) Exuvio-fecal shield.

Figs. 16–17.

Pupa of E. columbiana. 16) Dorsal view; 17) Ventral view.


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kania sp., 6.VI.2000, colls. C.S. Chaboo and H. Rowell (7 adults, 153 larvae; AMNH, CSCC, CUIC, INBio). Remarks. The testes varied in size and texture, from soft small rounded structures to large, stiff structures. This variation may be related to maturity of particular individuals. I was unable to determine if the endophallus surface is spiny or not. The 4th instar larva is similar to the 5th instar larva in the number and arrangement of scoli, furcae, setation and trichiation. The only obvious distinctions are in color and size. The shield has a pyramidal shape composed of layers of exuvial skins. Each exuviae was folded over once, with head capsules neatly stacked posteriorly, and the older caudal furcal integument enclosing the more recent one. The head capsules thus provide a convenient count for determining the larval instar. Discussion These observations of E. columbiana, supported by larval and adult reproductive morphology, suggest a complex defense system that is consistent with reports for other chrysomelids (summarized in Jolivet 1997), and an unusual mating system. Several levels of defense are discernable: grouping of eggs, larvae and pupae, and reflex behaviors like thanatosis and cycloalexy can be defensive (Vasconcellos-Neto and Jolivet 1989; Damman and Cappuccino 1996); larval scoli and trichiation can provide a physical armor for mechanical defense; setation can increase detection of intruders (Eisner et al. 1967); and red and black adult coloration may provide some aposematic protection. The fecal shield, host plant choice and maternal care suggest more sophisticated defensive strategies. Cassidine fecal shields have well-demonstrated defensive functions (Eisner et al. 1967; Olmstead and Denno 1992, 1993; Olmstead 1994; Go´mez et al. 1999; Eisner 2000), although Mu¨ller and Hilker (1999) have found contrary data. Vencl and Morton (1999) hypothesised that in species with maternal care the larvae have reduced or temporary fecal shields. However both E. coscaroni and E. columbiana have well-developed shields, constructed of feces and exuvial skins, and are obvious contradictions to this hypothesis. Host plant choice might also provide a chemical defense strategy. Could the liquid regurgitated upon handling be a plant-derivative with repellent effects, as has been demonstrated for many chrysomelids (Pasteels et al. 1998) or might the fecal shield incorporate plant compounds, as in other chrysomelid species (Go´mez 1997; Pasteels et al. 1998; Vencl and Morton 1998; Go´mez et al. 1999; Mu¨ller and Hilker 1999)? Interestingly, Mikania is now reported as a host plant for five cassidine species, E. columbiana, E. coscaroni, S. lebasii, Echoma clypeata (Panzer) (5 Echoma marginata (Linneaus)), and Echoma anaglyptoides Borowiec (Windsor and Choe 1994; Windsor et al. 1995). The plant recurs in the traditional folk medicines of diverse indigenous peoples (Bishayee 1994; Lentz et al. 1998; Facey et al. 1999). Consequent systematic explorations have revealed chemicals that are new (e.g., mikanolidae, dihydromikanolides, melampolides) (Ruengler et al. 2001) or rare (e.g., sesquiterpinoids, nerolidol derivatives) (Diaz et al. 1992) and which show insecticidal, anti-bacterial, anti-protozoal, and anti-inflammatory properties (Fiero 1999; Muelas et al. 2000). It is highly intriguing if these beetle species are exploiting this unusual chemistry. Prolonged presence of males with the female and larval brood is now re-

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corded for two eugenysines, E. coscaroni (Windsor and Choe 1994) and E. columbiana. Male presence may be a function of territoriality, which would increase mating opportunities, or of mate guarding which would assure paternity (Reid and Roitberg 1994). Paternal guarding of the larval brood is unlikely simply because of its rarity in insects but if it does indeed occur, this would represent another level of sophistication in this species’ defense system. Cassidine reproductive structures have been documented in several works (Spett and Lewitt 1925, 1926; Sanderson 1948; Varma 1955; Pajni et al. 1987; Bordy and Doguet 1987; Mann and Crowson 1983, 1996; Mann 1988; Verma 1985, 1988, 1996). In these studies, where a few genera and species were examined, and in my doctoral dissertation research involving more than 300 species from all cassidine tribes, the system of E. columbiana is unusual in three aspects. First, the ejaculatory duct of this species is exceptionally long and tightly folded into numerous coils. Second, the spermathecal duct in the female is also exceptionally long and coiled, as predicted for such correlated reproductive structures (Suzuki 1989). Third, the spermathecae has three receptacle chambers. This discovery in E. columbiana led me to sample other eugenysines species, and it appears that the three-chambered receptacle is more widespread within the tribe. Its function and its potential as a phylogeneticallyinformative character needs further exploration. The only clues we have about the physiological mechanisms operating in the cassidine spermatheca and the aedeagus come from the elegant studies of V. Rodriguez (1994b, 1995) with Chelymorpha cribraria (Fabricius) (5 Chelymorpha alternans Boheman). In that species the flagellum penetrates far up the spermathecal duct, even reaching into the spermatheca in one case (Rodriguez 1994b; Eberhard 1996:6–7, 352). Should this mechanism apply to other cassidines, the flagellum may navigate the very long and highly coiled spermathecal duct in E. columbiana. All the morphological and behavioral features discussed here fit with known data on the remarkable biology of cassidines; each aspect presents an interesting avenue of research. Among species exhibiting maternal care (Table 1), comparative data for particular aspects is quite patchy. Given that eight cassidine species have been reported as maternal care providers only in the last 12 years, then it is highly probable that more species with this unusual behavior await discovery. In order to clarify the nature of parental involvement and other complex behaviors in E. columbiana and other cassidine species, an interdisciplinary approach to observations and experimentation is necessary. Acknowledgments I thank the curators and collections for providing specimens for study. Field work was funded by a Rawlins Grant from the Department of Entomology, Cornell University and a field study grant from the American Museum of Natural History. F. Campos (OTS, San Jose´), facilitated my collection and study permits in Costa Rica and H. Cochran (INBio) facilitated my export permit. I thank H. Rowell for the field excursion, J. and R. Figueroa Sandi for help in the field, M. Schlaeper for taking field photos and L. D. Gomez for the plant identification and his many kindnesses during my stay at the Wilson Botanical Garden. J. Pruski (U.S. National Herbarium) provided information on Mikania while N. Upton (Green Umbrella, nature documentary production) discussed his observations of A. sparsa. T. Nguyen and C. Johnson generously provided technical help and F. Merino, D. Silva and J. Stark provided German


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