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52<br />

ne M a t o d e re l a t i o n s h i P s<br />

BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM<br />

Richter (1993) reported that Rhabditis<br />

stammeri (Völk), a carrion-dwelling<br />

nematode, is specifically associated with<br />

Nicrophorus vespilloides. Juveniles <strong>of</strong> the<br />

nematode use the adult beetle for transport<br />

to carrion where both adults and larvae <strong>of</strong><br />

beetles become infected. Inside the beetle<br />

larvae, the juvenile nematodes are transported<br />

to the pupal chamber via the larval<br />

gut. After pupation, the juvenile nematodes<br />

are found in the pupal chamber at protected<br />

places such as the exuvium or beneath the<br />

wings <strong>of</strong> the pupa. After emergence from<br />

the pupa, the nematodes migrate to the gut<br />

and genitalia <strong>of</strong> the adult beetles. Rhabditis<br />

stammeri is also transmitted from one adult<br />

beetle to another during copulation. Infection<br />

by nematodes probably occurs in many<br />

other species <strong>of</strong> Nicrophorus as well, and is<br />

an area needing additional study.<br />

st r i d u l a t i o n<br />

All species <strong>of</strong> Nicrophorus have a stridulatory<br />

structure in both males and females<br />

(Fig. 2). Stridulation is used during burial <strong>of</strong><br />

the carcass, copulation, and the interaction<br />

between the female and her brood (Niemitz<br />

1972, Niemetz and Krampei 1972). Huerta<br />

et al. (1992) conducted lab experiments<br />

with N. mexicanus and found that a lack <strong>of</strong><br />

stridulation in the female resulted in poor<br />

or no “bonding” between the female and her<br />

<strong>of</strong>fspring, which negatively affected larval<br />

survival. Inhibition <strong>of</strong> stridulation in the<br />

male affected and sometimes precluded<br />

copulation. Lack <strong>of</strong> stridulation in both nest<br />

partners may affect the coordination <strong>of</strong> nest<br />

preparation (Halffter 1982, Huerta et al.<br />

1992, Halffter et al. 1983).<br />

Pr e d a t i o n<br />

Most ground-feeding insectivorous<br />

birds are probably familiar with burying<br />

beetles, and these beetles have been recorded<br />

as among the food items <strong>of</strong> several species,<br />

especially crows that routinely visit carrion.<br />

Jones (1932) conducted a number <strong>of</strong> experiments<br />

using N. americanus, N. orbicollis,<br />

and N. pustulatus to determine if a variety<br />

<strong>of</strong> insectivorous birds would be deterred by<br />

these aposematically colored beetles. Although<br />

he tested only seven specimens, none<br />

were eaten by the birds, while 42 out <strong>of</strong> 46<br />

other beetles (representing seven species) <strong>of</strong><br />

comparable size were eaten by seven species<br />

<strong>of</strong> birds during 93 feeding events. Jones<br />

concluded that birds do avoid these brightly<br />

colored beetles, at least when other food is<br />

available.<br />

so c i a l i t y<br />

The activities <strong>of</strong> adult Nicrophorus<br />

species in rearing their young is the highest<br />

level <strong>of</strong> sociality attained in the Coleoptera<br />

(Wilson 1971, Wilson and Fudge 1984). On a<br />

large carcass, the mating system is variable.<br />

Larger carcasses can support greater numbers<br />

<strong>of</strong> larvae and support broods <strong>of</strong> greater<br />

total mass than smaller carcasses (Trumbo<br />

1992). Consexual adults <strong>of</strong>ten tolerate each<br />

other and <strong>of</strong>ten feed each other’s young in<br />

a quasisocial fashion (Scott and Traniello<br />

1990a, Eggert and Müller 1992, Trumbo<br />

1992, Scott and Williams 1993, Trumbo and<br />

Wilson 1993). According to Trumbo and Wilson<br />

(1993), females <strong>of</strong> smaller Nicrophorus<br />

(N. defodiens, N. tomentosus) species were<br />

much more likely to feed young cooperatively<br />

than females <strong>of</strong>, for example, N. orbicollis.<br />

They hypothesized that since adults cannot<br />

discriminate between related and unrelated<br />

young, they feed any larvae on the carcass<br />

to ensure adequate care for their own young.<br />

Larger carcasses were more difficult to exploit<br />

because: (a) they took longer to conceal<br />

beneath the leaf litter; (b) they were less<br />

likely to be rounded into brood balls; (c) they<br />

were more likely to be utilized by dipterans;<br />

and (d) they were occupied by greater numbers<br />

<strong>of</strong> congeners (Trumbo 1992).<br />

Scott (1994b) suggested that competition<br />

with flies promotes communal breeding<br />

in N. tomentosus. She demonstrated that

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