The Coleopterists Society

Oviposition Behavior and Associated Morphology of the Neotropical Anthribid Ptychoderes rugicollis Jordan (Coleoptera: Anthribidae) Author(s): Anne T. Howden Source: The Coleopterists Bulletin, Vol. 46, No. 1 (Mar., 1992), pp. 20-27 Published by: The Coleopterists Society Stable URL: http://www.jstor.org/stable/4008931 . Accessed: 02/05/2013 16:43
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The Coleopterists Bulletin, 46(l):20-27.

1992.

OVIPOSITION BEHAVIOR AND ASSOCIATED MORPHOLOGY OF THE NEOTROPICAL ANTHRIBID PTYCHODERES RUGICOLLIS JORDAN (COLEOPTERA: ANTHRIBIDAE)
ANNmT. HOWDEN Biology Department, Carleton University, Ottawa, Ontario KIS 5B6, Canada
ABSTRACr

Basedon observationsmadein Chiapas,Mexico,femalesof the Neotropicalanthribid Ptychoderes rugicollisJordanbore deep oviposition holes in dead wood usingonly their ovipositors, a behaviorapparently in Curculionoidea. unreported A single male attends each female rathercasually while she is boring the hole, but guardsher more closely aftercopulatingandwhileshe oviposits.Brushing the sawdustfromthe female'sabdomen may be a sensory or an allogroomingbehavior. Examinationof the ovipositor shows featuresassociatedwith this method of oviposition to include:the long, stiffovipositor; the form, position, and strengthof the teeth on the apical toothed plate;and the fusion of the toothed plate to the coxite.

Oviposition behavior and the functional morphology of the ovipositor of Curculionoidea is a relatively unexplored field with the potential of yielding information ofphylogenetic value. For example, in the majority ofthe Anthribidae the ovipositor terminates in conspicuous, distinctive, outwardly-directed teeth or scoop-like projections. Yet, conspicuous as they are, the role of these teeth in oviposition is virtually unknown. Oviposition in anthribids is apparently quite varied, although detailed observations are usually lacking in the literature. Among Anthribinae, Anthribus nebulosus Forster is reported to eat a hole in the semisoft exoskeleton of a female scale insect and then lay an egg inside the brood chamber of the scale closing the wound with a secretion (Schmutterer 1952), Euciodes suturalis Pascoe is suspected of using the ovipositor to make the oviposition hole in grass stems (Penman 1978), and Trigonorhinus limbatus (Say) uses the toothed ovipositor to simply push aside parts of a composite flower head, or it may oviposit in stems (observation by Valentine, see Howden, in press). Other reports vaguely note that anthribines oviposit in cracks and crevices in bark. The choragine Araecerus fasciculatus (Degeer) uses the ovipositor to bore a hole in the host (fruit, seed, or stalk) and then to deposit the egg in the bottom of the hole (El Sayed 1935, 1940). Herein the large Ptychoderes rugicollis Jordan (Anthribinae) is reported to bore an oviposition hole 10 mm deep in a log using only its ovipositor! Not only is this a remarkable feat, but several aspects of the oviposition behavior as well as the morphology of the ovipositor are unusual in Curculionoidea. Field Observations of Oviposition Behavior Ptychoderes rugicollis Jordan (Fig. 1) occurs in southern Mexico and Central America where it is one of the largest species of Anthribidae, 15 to 22 mm in 20

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THE COLEOPTERISTS BULLETIN 46(1), 1992

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Figs. 1-3. Ptychoderesrugicollis. 1, habitus, Female 1. 2, tarsal segment 1, right

hind leg, Male 2. 3, apex of ovipositor, dorso-lateral view; female from Mexico, Veracruz, Lake Catemaco.

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THE COLEOPTERISTS BULLETIN 46(1), 1992

length. It is densely covered with black, rust, and white scales arranged in a pattern which is very cryptic against the bark of a log. Observations were made at El Chorreodero, 8 km E Chiapa de Corzo, Chiapas, Mexico. This site is a densely vegetated ravine at an altitude of approximately 590 m in the Chiapan highlands. On the night of 1 June 1990, a heavy rain started at 1830 and eased off at 1930; a light misty rain occurred sporadically thereafter and the vegetation remained wet all evening. Adults of Ptychodereswere observed on the side of a large log (species undetermined) lying on a moderately steep slope. The wood of the log was sound and the thin bark was intact. A large rock provided a convenient seat during the hour and a half observation period. A battery-operated head lamp provided good illumination. An optical visor provided 1.75 x magnification and a hand lens provided an additional 1.25 x magnification; this was adequate for the large size of the beetles. The first observation began shortly before 2000 when a pair of P. rugicollis were located on a log. The female (Female 1) held her body parallel to the surface of the log. She was apparently boring in the log with her ovipositor, although the only indication of this was a slight raising and lowering of the pygidium. A male (Male 1) was standing obliquely beside Female 1 with his antennae draped motionlessly across Female l's body. Thereafter, at intervals of three to seven minutes, Female 1 withdrew her ovipositor from the log then partially retracted and extruded it several times; at each retraction and extrusion of the ovipositor the trickle of powdery sawdust that fell from the tip was clearly visible against the dark background. At 2115 Female 1 moved forward half a pace, lifted her abdomen slightly and defecated. Male 1 immediately mounted her back and very briskly brushed off the apex of her abdomen with his hind tarsi, scattering powdery sawdust. Male 1 then apparently copulated with Female 1 (though I could not get into position to verify this) and after a brief period walked away several paces, retracting his shiny aedeagus. Female 1 promptly backed up to the oviposition hole and Male 1 returned and assumed his former position, with the addition of placing a middle leg on the female's body. Oviposition followed immediately and was accomplished with a very regular and gentle pulsing in and out. Male 1 left the site while Female I was still occupied. At 2130 I had to terminate the observation. The oviposition site was cut out of the log with a knife. The hole was roughly circular, not quite perpendicular to the surface of the log, and the interior walls were surprisingly smooth. The creamy white egg was nearly spherical and 1.4 mm in diameter. It was embedded at the bottom of the 10 mm long hole. The channel above the egg was plugged for a short distance with a hard material which appeared to be a combination of bits of wood fibers and a binding substance which probably included feces. The initiation of oviposition was observed in another episode which began at 2045, only 20 cm away from the first pair. A solitary female (Female 2) walked around the vicinity tapping the log with her antennae and eventually paused at one spot and pulled at the thin outer bark with her mandibles as if feeding on it. In less than five minutes Female 2 walked away a short distance, returned, and backed up to the torn spot until the tip of her abdomen was touching it. Female 2 assumed a tilted position with the fore legs fully extended and the head and prothorax slightly deflected. Otherwise boring proceeded in a manner like that of Female 1. Another male (Male 2) that had walked around the area before Female 2's initial use of her mandibles returned when she began

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THE COLEOPTERISTS BULLETIN 46(1), 1992

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boring the oviposition hole and behaved in a manner like that of Male 1. This observation was also terminated at 2130, before Female 2 laid an egg. Throughout the observations my movements disturbed Male 1 more readily than Female 1; several times Male 1 backed away from Female 1. However, when another male (Male 3) approached the pair, Male 1 moved closer to Female 1, resting the apex of his rostrum across the base of her elytra. Male 1 was not disturbed when Female 2 investigated the area, nor was he disturbed by Male 2.

Laboratory Examination of Abdominal Structures Involved in Oviposition To study the female reproductive tract of P. rugicollis in the laboratory, the entire abdomen was removed and macerated in hot KOH. This destroyed the muscle tissue, but left the tracheae and relevant parts of the reproductive system intact. No staining was needed. Three females of P. rugicollis (Female 2 and two specimens from Veracruz, Mexico) and one female of P. nebulosus (Olivier) from Veracruz were dissected. In addition, the extruded ovipositor of a pinned specimen of P. rugicollis from Veracruz was examined. The structures of P. nebulosus are similar to those of P. rugicollis; the following discussion refers to P. rugicollis only. Terms for the parts of the female reproductive system follow Howden (in press) in which the most frequently used terms were chosen regardless of their scientific or etymological accuracy. The morphology and function of the structures involved in oviposition are discussed in the order of their position on the beetle, beginning with the apex of the dorsal surface of the abdomen. The pygidium (tergite 7) did not appear to be actively used in oviposition. However, the contour and thickness of the integument suggest that both tergite 7 and sternite 7 might assist the boring and ovipositing female to (1) maintain a purchase on the substrate, (2) block the work area from small predators or parasites, and (3) serve as a track for the extruded ovipositor. Tergite 8 is light brown, thinly sclerotized and approximately as long as sternite 8 without its apodeme. Tergite 9 characteristically is sclerotized in Anthribidae. In P. rugicollis tergite 9 (Fig. 5) has a pair of long, heavily sclerotized lateral rods. When the ovipositor is fully extruded these sclerotized rods lend rigidity to the area between tergite 8 and the proximal end of the coxites, a long membranous section which otherwise lacks rigidity. The ovipositor is comprised of the coxites plus styli; in Anthribidae especially, the coxites are frequently referred to as hemisternites. In P. rugicollis the coxites are heavily sclerotized, complex structures (Figs. 3, 4, 6), 7.2 to 7.5 mm in length, as long as to slightly longer than (1.08-1.2 x) the abdomen, in situ extending anteriad into the thorax. In dorsal view the coxites appear to be a pair of cylindrical tubes closely abutting medially. When pried apart with forceps the abutting surfaces are revealed to be vertical, flat, smooth, polished, black, and very heavily sclerotized. These median vertical surfaces are deep distally, gradually diminish in depth and amount of sclerotization proximally, and near the proximal end become fused and horizontally flattened with only a thin contiguous line darkly sclerotized. The coxites are connected by a membrane (the vagina) attached to

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THE COLEOPTERISTS BULLETIN 46(1), 1992

mm 1.0 Figs. 4-7. Ptychoderes rugicollis,Female 2. 4, ovipositor, ventralview. 5, tergite9 and hind gut, dorsal view. 6, ovipositor and part of reproductivetract, lateralview. 7, an = anus, bc = bursacopulatrix,co = common sternite8, ventralview. Abbreviations:
oviduct, sty = stylus, va = vagina.

the ventral edge of the vertical median surfaces; this permits separation of the coxites distally by a distance greater than the width of a coxite. Apically each coxite terminates in a very hard, black, "toothed plate" (Holloway 1982), which is a cluster of three subequal teeth plus a small ventrolateral tooth on a solid base (Fig. 3). The teeth are somewhat scoop-shaped, slightly convex on their dorsal and median surfaces and slightly concave on their anterior and lateral surfaces. There are many long setae arranged in series and tufts on and between the teeth. The stylus is long, cylindrical, distally setate, and is borne on a colorless papilla dorsally at the base of the toothed plate. Most of the basal edge of the toothed plate is separated from the apex of the coxite by connective tissue which would seem to allow movement of the plate as if it were articulated. However, laterally the plate and coxite are

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THE COLEOPTERISTS BULLETIN 46(1), 1992

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continuously sclerotized, hard, and inflexible. Manually it is impossible to move the plate independently of the coxite. At approximately the proximal third, the dorsal surfaces of the coxites diverge and each coxite gradually attenuates, becoming vertical, broad, bladelike, and slightly concave on its inner surface. This attenuation (the lateral rod of Holloway 1982) is reinforced throughout its entire length with a darkly sclerotized median streak. Ventrally the ovipositor is almost flat (Fig. 4). The membrane connecting the coxites medially is elastic, permitting great separation of the coxites laterad. Apically this connecting membrane terminates in three thin flat lobes: two lateral rectangular lobes positioned beneath the base of the toothed plate and one median elongate tongue-like lobe, the apex of which has a tendency to curl ventrad. The dorsal surface of the median lobe is densely covered with minute, recumbent, caudally directed setae. In the extruded ovipositor of a pinned specimen, the coxites are pulled together by the drying, shrinking membranes connecting them, and the median lobe is then situated over (dorsad of) the lateral lobes. The vulva is located just before the distal end of the coxite. To bore the oviposition hole in the log, only small twists or rotations of the ovipositor within the wood would give 3600 coverage with the teeth. The teeth of one coxite act as reamers, while the teeth of the other coxite act as compressors. It seems likely that the ovipositor is extruded with the coxites contiguous, then within the hole in the wood the coxites are distended and pulled back, the concave anterior surfaces of the teeth hauling out sawdust. In one female examined, the concavities of the teeth were packed with sawdust. Sternite 8 (Fig. 7) is light brown and thinly sclerotized; its apodeme is approximately as long as the body of sternite 8, dark, more thickly sclerotized, and anteriorly serves as a site for muscle attachment. A pair of very stout tracheae, 0.5 x as wide as the coxites, originate at the spiracle on tergite 7. These tracheae divide into several branches as they go between the coxites. One branch is readily visible along the dorso-median edge of each coxite to its extreme apex. The area of the log where the females were ovipositing had small black spots on the surface of the thin bark. The digestive tract of Female 2 was packed with bits of black material (Fig. 5). This female had pulled at the bark with her mandibles before beginning to bore the oviposition hole, and it is possible she was only feeding. Holloway (1982:32) observed that ". . . black vegetative growths and fruiting bodies of ascomycete fungi on or protruding through the
bark are ... eaten by many adult anthribids."

Other Structures Involved in Oviposition Two structures of P. rugicollis are sexually dimorphic in a fashion apparently related to oviposition behavior. Males have long antennae which they drape over the female in a passive form of guarding. The length of the antennae of the males varied with the size of the beetle. In Male 1 and Male 2 the antennae are distinctly longer than the beetle; in the very small Male 3 the antennae reach only to ventrite 3. The antennae of the females (Fig. 1) are short, reaching just beyond the middle of the prothorax. Neither the antennae nor other parts of the body were studied for sensory structures. Male I used its slightly modified hind tarsi to brush the female's abdomen immediately before copulating. The ventral surface of tarsal segments 1 and 2

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THE COLEOPTERISTS BULLETIN 46(1), 1992

(Fig. 2) are densely covered with stout recumbent, overlapping setae; in addition, on the inner edge there is a row of sparse longer setae. The tarsi of females do not bear these longer setae although the ventral surface is otherwise similarly clothed. Males of P. nebulosus do not have the longer tarsal setae and the vestiture is like that of the females. When Male 1 brushed Female 1's abdomen, the falling sawdust was conspicuous, giving the immediate impression of the male grooming the female (allogrooming). The brushing may include both grooming and sensory functions, perhaps analagous to the male bottle brush weevil, Rhinostomus barbirostris (Fabricius), "wiping" the female with its long rostral hairs (Eberhard 1983:242).

Comparisons with Other Curculionoidea Oviposition in dead or dying wood is not uncommon in some groups of Curculionoidea, notably, some Belidae, Brentidae, and the Curculilonidae subfamilies Cossoninae, Cryptorhynchinae, Rhynchophorinae, and Zygopinae. However, in all those weevils where oviposition has been documented, the oviposition hole is made in the wood using the rostrum exclusively. Using the ovipositor exclusively to bore the oviposition hole in wood, as did P. rugicollis, may be unique to Anthribidae. The non-articulated, heavily sclerotized apical toothed plate on the coxite is assumed to be essential to boring the oviposition hole. By way of comparison, consider the coxite of what is possibly the most primitive anthribine whose ovipositor has been studied-the New Zealand Gynarchaeus ornatus (Sharp) (Holloway 1982). In that species the coxite is unmodified (i.e., without a toothed plate) and simply rounded apically. The non-articulated toothed plate of Ptychoderes rugicollis may have evolved from such an unmodified coxite or it may represent a secondary fusion of an articulated plate. Various forms of heavily sclerotized articulated styli and non-articulated processes on the coxite are found in Curculionidae. Oviposition behavior is unknown for most of these taxa, but where it is known, oviposition is in the soil and not in dead wood. In both the brentid Arrhenodes minutus (Drury) (Sanborne 1983) and the curculionid Rhinostomus barbirostris (Fabricius) (Eberhard 1983), the male guards the female zealously while she bores the oviposition hole in the log with her rostrum. The male has copulated, or is in the act, as the female bores, and he continues protecting his investment until after the female has oviposited. Such parental assurance obviously requires less dilligence on the part of a P. rugicollis male than in either of the above cases, since no other anthribid male can copulate with a female while her ovipositor is boring in a log. Nevertheless, an example of aggressive guarding of the female has been reported in Anthribidae by Thompson (1963:37-38) in the Ghanaian Deuterocrates longicornis Fabricius. In that case the male faced the female and positioned his long antennae along each side of her body. Guarding males challenged stray males with their mandibles in "intense rivalry," but were never seen to inflict injury. Apparent lack of aggression in guarding male P. rugicollis may be only an artifact of the small sample size. This small study has generated more questions than it answererd, but hopefully, it will stimulate the careful reporting of oviposition and the illustration of the ovipositing structures.

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THE COLEOPTERISTS BULLETIN 46(1), 1992

ACKNOWLEDGMENTS

27

I have relied heavily upon B. A. Holloway's 1982 revision of the New Zealand Anthribidae with its comprehensive study of the female reproductive tract. Donald B. Thomas and family, USDA Screwworm Laboratory, Tuxtla Gutierrez, Chiapas, Mexico, provided field assistance and were the first to locate the anthribids. Barry Valentine, Ohio State University, Columbus, confirmed my identifications and assisted with literature; I thank him and Robert S. Anderson, Canadian Museum of Nature, Ottawa, for comments on earlier drafts of the manuscript. Karen Hamilton, Lewis Ling, and Linda Sealey, Carleton University, assisted with the plates, took the scanning electron micrographs, and took the habitus photograph, respectively. The assistance of all is greatly appreciated. LrrERATURE CUTED EBERHARD, W. G. 1983. Behavior of adult bottle brush weevils (Rhinostomusbarbirostris)(Coleoptera: Curculionidae). Rev. Biol. Trop. 31:233-244. M. T. 1935. On the biologyofAraecerusfasciculatusDe ELSAYED, Geer(Col.,Anthribidae), with special referenceto the effects of variations in the nature and water content of the food. Annals Appl. Biol. 32:557-577. 1940. The morphology,anatomy and biology of Araecerus fasciculatus De Geer. Bull. Soc. Fouad Ent. 24:82-143. HOLLOwAY, B. A. 1982. Anthribidae(Insecta:Coleoptera).Faunaof New ZealandNo. 3. 264 pp. A. T. In press. Structures HowDEN, relatedto oviposition in Curculionoidea.In: Anderson, R. S., and C. H. C. Lyal (editors).[Proceedings of special interestgroup, Biology and Phylogenyof Curculionoidea, XVIII Int. CongressEntomol.]Mem. Ent. Soc. Washington. D. R. 1978. Biology of Euciodessuturalis(Coleoptera: PENMAN, Anthribidae) infesting cocksfoot in Canterbury. N. Z. Ent. 6:421-425. SANBwRNE, M. 1983. Some observations on the behaviour of Arrhenodesminutus (Drury)(Coleoptera: Brentidae).Coleopts Bull. 37:106-113. H. 1952. Die Okologieder Cocciden(Homoptera,Coccoidea)Frankens SCHMUTrERER, (Pt. 3). Ztschr.f. angew. entomol. 33:65-100. G. H. 1963. Forest Coleopteraof Ghana. Biologicalnotes and host trees. THOMPSON, Oxford ForestryMem. 24. 78 pp. (Received 6 November 1990; accepted28 May 1991)

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