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Host Defense in Nicrophorus Quadripunctatus Against Brood Parasitism by Ptomascopus Morio (Coleoptera: Silphidae: Nicrophorinae)

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Host Defense in Nicrophorus Quadripunctatus Against Brood Parasitism by Ptomascopus Morio (Coleoptera: Silphidae: Nicrophorinae) Popul Ecol (2006) 48:167–171 DOI 10.1007/s10144-005-0245-9 NOTES AND COMMENTS Seizi Suzuki Æ Masahiro Nagano Host defense in Nicrophorus quadripunctatus against brood parasitism by Ptomascopus morio (Coleoptera: Silphidae: Nicrophorinae) Received: 24 June 2005 / Accepted: 21 October 2005 / Published online: 21 December 2005 Ó The Society of Population Ecology and Springer-Verlag Tokyo 2005 Abstract Few studies have been conducted on the host Cervo et al. 2004). For example, parasitic cuckoos lay defenses of insects against brood parasitism. We inves- their eggs in the nest of the host species that incubate tigated whether the silphid beetle Ptomascopus morio,a and rear the parasitic offspring; thereby, the hosts suffer brood parasite of related silphid species Nicrophorus severe reductions in reproductive success (Payne 1998; concolor, can also parasitize another silphid species Soler et al. 2001). Parasitism favors the evolution of host Nicrophorus quadripunctatus and the manner in which defenses such as egg rejection (Soler and Soler 2000). N. quadripunctatus defends itself against parasitism. Such host defense mechanisms may be specific to Successful brood parasitism under natural conditions changes in parasitic behavior and may lead to an evo- was not observed at the time of year when P. morio and lutionary arms race (Rothstein 1990). In contrast to the N. quadripunctatus are both reproductively active. Fol- studies of birds and eusocial hymenopterans (Davies and low-up experiments revealed that P. morio attempts to de Brooke 1998; Soler et al. 2001), little research has oviposit near N. quadripunctatus nests, but is rarely suc- been conducted on brood parasitism in noneusocial cessful if adult hosts are present. When P. morio larvae insects (Rasa and Heg 2004), and there is even less were experimentally introduced to N. quadripunctatus research on host defenses. broods, some P. morio larvae survived when the host and All species of the genus Nicrophorus (Silphidae, Cole- parasite larvae were at the same stage. We concluded that optera) care for their young, such as providing them with N. quadripunctatus defends itself against brood parasit- food (small vertebrate carrion), guarding against intrud- ism in two ways: (1) potential brood parasites are re- ers exclusively, and regurgitating food for their young pelled, thus limiting their access to the resource; and (2) (Scott 1998). The genus Ptomascopus is believed to be a the young of the parasitic species are killed. sister group of Nicrophorus (Dobler and Mu¨ ller 2000; Peck and Anderson 1985; Szalanski et al. 2000). Pto- Keywords Host rejection Æ Facultative brood mascopus morio shows simple parental care (Suzuki and parasitism Nagano, in press). Their larvae are able to grow without parents in laboratory conditions (Peck 1982; Trumbo et al. 2001), but it is difficult for them to grow without parents in field conditions because of predation and Introduction competition (Suzuki and Nagano, in press). At a study site near Kyoto, Japan, P. morio Kraatz was found to be a The relationship between host and parasite has been brood parasite of Nicrophorus concolor Kraatz (Trumbo extensively studied in birds (Rothstein 1990) and euso- et al. 2001). Larvae of P. morio live in the broods of N. cial hymenopterans (Wilson 1971; Sledge et al. 2001; concolor and grow under the care of host parents though the range of care is undetermined (Trumbo et al. 2001; S. Suzuki, personal observation). P. morio readily para- & S. Suzuki ( ) sitized N. concolor in the field, and parasitism by P. morio Ecology and Systematics, Graduate School of Agriculture, Hokkaido University, 060-8589 Sapporo, Japan reduced the brood mass of N. concolor in the laboratory E-mail: [email protected] (Trumbo et al. 2001). Since Nicrophorus beetles eliminate Tel.: +81-11-7062486 intruders exclusively for their brood, such as the fly, rove Fax: +81-11-7064939 beetle, and other Nicrophorus spp. (Mu¨ ller and Eggert M. Nagano 1990; Scott 1990; Suzuki 2000), the P. morio larvae will Echigo-Matsunoyama Museum of Natural Science, have ways to deceive N. concolor parents in order to stay in Niigata, Japan the latter’s brood. 168 In contrast, P. morio seldom parasitized another host A second experiment was conducted to determine the species, Nicrophorus quadripunctatus Kraatz, in the same potential of P. morio to parasitize N. quadripunctatus. locality (Trumbo et al. 2001). Since the reproductive One pair each of N. quadripunctatus and P. morio and seasons of N. quadripunctatus and P. morio differ slightly 15 g of chicken were placed in a polyethylene container in Honshu, Japan (Nagano and Suzuki 2003), whether (15·15·9cm3) that was half-filled with soil; the con- the low frequency of the parasitism of N. quadripuncta- tainer was covered to prevent further colonization by tus by P. morio was caused by host defenses or by sea- insects (n=21). All the containers were checked at sonal segregation has not been established. However, the 10 days after the introduction of N. quadripunctatus and phenology of P. morio in Otaru, Hokkaido is similar to P. morio, and the larvae were checked. The larvae were that of N. quadripunctatus (Ohara 1995). Therefore, the identified, and all adult beetles were checked for injuries aim of our study was to investigate the host defenses of to their legs and antennae. These containers were placed N. quadripunctatus in Otaru where seasonal segregation in scavenger-proof cages on the forest floor until the between these two species can be negligible. experiments were terminated. The aims of this paper are as follows: first to confirm All experiments were conducted between early to mid the presence or absence of N. quadripunctatus parasiti- July 2002. zation by P. morio in the conditions in which P. morio is expected to parasitize N. concolor, and second to investigate the mechanism by which N. quadripunctatus Parasitism of undefended carcasses defends itself against parasitization by P. morio. The rarity of parasitism by P. morio could be either due to few attempts at parasitism or due to the successful Materials and methods host defenses of N. quadripunctatus. In order to distin- guish between these two possibilities, a pair of N. All experiments were conducted in the forest of the quadripunctatus and a piece of chicken (15 g, n=10) Naebo Forest Park in Otaru, near Sapporo, southwest- were introduced into a soil-filled container ern Hokkaido, North Japan. (15·15·9cm3). After ensuring that the chicken was buried in the soil, the N. quadripunctatus pair was con- fined to a small, clear plastic cylinder (16 mm diameter, Phenology of N. quadripunctatus and P. morio 39 mm depth) with many small holes to allow the pas- sage of odor and sound; this cylinder was placed near Burying beetles were collected using hanging polyethyl- the chicken carcass. A P. morio female was introduced ene traps (97 mm diameter, 166 mm depth), with pieces into the container which held the undefended carcass. of chicken as bait. Three traps were installed at the site; The container was checked for P. morio larvae 5–7 days they were arranged linearly at intervals of at least 5 m later. and were suspended from trees at a height of 1.5–2.0 m above the ground to prevent their removal by verte- brates. The sampling site was a natural mixed forest Host defense by N. quadripunctatus (Fraxinus mandshurica and Ulmus davidiana were com- mon species). All traps were kept open for 5 days, and To determine whether the presence of N. quadripunctatus the beetles were collected on the last day. Nicrophorus adults prevented P. morio from oviposition near the and Ptomascopus were identified from among the carcasses, a pair each of N. quadripunctatus and P. morio collected beetles and were counted. Sampling was and 15 g of chicken were introduced into a polyethylene performed once or twice a month from May to container (15·15·9cm3) that was half-filled with soil September 2001. (n=22). After confirming oviposition by either species, all the adult beetles were removed. The larvae were identified after hatching. Presence of brood parasitism To investigate whether N. quadripunctatus is capable of distinguishing and excluding parasitic larvae, a pair of To assess brood parasitism under natural conditions, 36 N. quadripunctatus and 15 g of chicken were introduced pieces of chicken (15 g) were placed in soil-filled poly- into a soil-filled polyethylene container. When the hat- ethylene bottles (90 mm diameter, 117 mm depth) that ched larvae appeared on the carcass, each brood was were suspended from trees at a height of 1.5–2.0 m. All subjected to one of three treatments: bottles were open at the top to allow the beetles to freely enter and then colonize the resource. The containers 1. The same-stage treatment (n=14): five of the P. mo- were checked 10 days after baiting. Larvae and adult rio first instars were introduced into the container nicrophorines were morphologically identified, and with the N. quadripunctatus brood when these larvae adults with a brood were regarded as parents. The reached the carcass. presence of P. morio larvae in the Nicrophorus brood can 2. The different-stage treatment (n=12): five of the be regarded as evidence of brood parasitism. P. morio first instars were introduced into the con- 169 tainer with the N. quadripunctatus brood when these lifrons, two by an unknown Nicrophorus sp. (with larvae had reached the second instar. Nicrophorus larvae but no adults) and one by P. morio. 3. The parental-removal treatment (n=13): the pair of No P. morio demonstrated parasitism. N. quadripunctatus was removed when the N. quad- Brood parasitism was rare when P. morio was ripunctatus larvae appeared on the carcass and five experimentally introduced with N.
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